Deep-Sea BiologySIO-277 • Instructor – Lisa Levin • Contact: firstname.lastname@example.org, 534-3579 • Room 2236 Sv - call before visiting • Tues & Thurs, 9:30-10:50, in Vaughan 100 • Website:http://cmbc.ucsd.edu/Students/Current_Students/SIO277/ • Series of 20 lectures ~ 70 min w/discussion • Assigned readings on electronic reserve prior to each lecture, also some book chapters • Texts: • Gage and Tyler - Deep-Sea Biology • Koslow - The Silent Deep
Student Requirements: • Read assigned papers before each lecture • Attend each lecture • Student Cruise - Oct. 31 (Saturday) • Mid term assignment Challenger Forward – due Oct. 29th • Research Proposal (Deep-Sea Future) Develop research needed for stewardship of the deep-sea in the face of climate change, mining, fishing, energy extraction etc. Read about subject, identify unanswered problem/issue, write a hypothesis-based research proposal (5 page max). • Abstract due Nov. 3 or earlier • Proposal due Nov. 24th • Oral presentation of proposal in a symposium during final exam week Dec. 10th
Course EvaluationGrading: Letter or S/U • Take Home Mid-Term Assignment (30%) Challenger Forward http://19thcenturyscience.org/HMSC/HMSC-INDEX/index-linked.htm • Written Research Proposal (40%) • Oral Presentation (25%) • Participation in Discussion/Cruise (5%)
Reading for Sept. 29, Oct.1 Sept. 29 Physical Environment: • Gage and Tyler 1991. Chapter 2. Oct. 1 Faunal composition, depth zonation: • Carney, R.S. Zonation of deep biota on continental margins. 2005. Oceanogr. and Mar. Biol: An Annual Review 43: 211-278. • Gage and Tyler 1991. Skim through images on pages 61-162.
Orientation – The global ocean A blank slate
Most of the Earth is Ocean and most of that is Deep Sea • Percent distribution of earth’s surface assessed in vertical relief. • Hypsographic curve
Defining the deep sea> 200 m(beyond the shelf break)or > 1000 m (hard core) Definitions can be important - much human impact is < 1000 m
Zonation Terminology meters 200 500 1000 3000 4000 6000 Epipelagic (euphotic) Continental Shelf Upper Continental Slope Mesopelagic (disphotic) Bathyal Lower Continental Slope Bathypelagic (aphotic) Continental Rise Benthopelagic Abysss Hadal
Continental Margins Abyss
History: Before Exploration Socrates (600 BC): The beginning of wisdom is to know that one knows nothing Aristotle (300 BC):The ocean (deep sea) is a frontier to be explored [180 spp, recorded from the Aegean Sea) Pliny (50 BC): The deep sea is an inferior world. All we know of it is all there is to be known. Posidonius(1 BC) Mediterranean Sea is 2000 m deep.
History of Deep-Sea Biology • Eric Mills – Problems in Deep-Sea Biology: an historical perspective (In: The Sea, Vo. 8 – the deep –sea edited by G. Rowe – 1983) – book on reserve • Tony Koslow - Chapter 1 in The Silent Deep. “The rise of deep-sea Exploration: Early paradigms” pp 8-22 Chapter 2. “On the shoulders of giants: The Challenger Expedition”pp. 23-39.
The beginning John Ross – (1818 – 1819) Baffin Bay –deep sea ‘clamm’ - 4 samples from 850 to 2000 m [crustaceans, corals, shellfish, worms, basket star from 1.6 km) In searching the NW Passage 2. James Clark Ross (1841-1847) Tasman Sea/Antarctic – fauna to 750 m Noted similarity with high latitude fauna and concluded uniform cold temperature at seafloor. 3. Harry Goodsir – (1845) Davis Strait (Arctic)– fauna dredged to 550 m
Edward Forbes (1815-1854) • Described marine faunas of European Seas • Described major biogeographic provinces from Arctic to Mediterranean and Caspian seas • Vertical zonation of benthos - established the science of marine benthic ecology - pattern in species distributions • Aegean Sea to 420 m - unfortunate focus. Unproductive waters, little in deep water. • Ignored work of earlier scientists
Azoic Hypothesis (1859) life absent > 550 m (300 fathoms). • Deep sea is • Dark • Cold • High Pressure • Stagnant and anoxic How could life survive?
Azoic Hypothesis • Attacked by G.C. Wallich • 1860 - 13 starfish recovered from a sounding line at 1260 fathoms (2300 m) in the N. Atlantic off Greenland • 1861 - Allman and Milne Edwards – 15 species recovered from a broken telegraph cable in the Mediterranean between Sardinia and N. Africa at 2300 m including stony coral. Yet the azoic paradigm persisted
John Jeffreys (1861 – 1868) Shetland Island dredging to 311m 204 species • 51-ft sailing vessel • No auxillary power • Hemp lines for dredging Osprey
Azoic theory in question Otto Torell (1861-1865) a. Benthic fauna at 2560m off Spitzbergen Michael Sars and G.O. Sars (1864-1868) a. Dredging to 550 m in Norwegian fjords b. 427 species of invertebrates including asteroids and crinoids (connections to fossil record) Led to the idea of deep sea a refuge for extinct faunas.
Azoic theory in question L.F. de Portales & L. Agassiz (1867-1868) dredged fauna off Grand Bahama Bank to 1555 m depth. W.B. Carpenter & C. Wyville Thomson • H.M.S. Lightning (1868) • Dredged fauna to 1189 m N.E. Atlantic • Deep water temperatures low (0 – 8.5oC)
Beginning of Big Science • Government funding • Networking • Politicking • Manipulating the public’s imagination
Wyville Thompson and William CarpenterHMS Lightening 1868 • North of Scotland between Shetlands and Faroes • Dredged 10 d of 6 wk at sea! To 1180 m • Abundant life everywhere, relict species • Brisingid starfishes, Hexactinellid sponges • Arctic outflows and NA Deep Water (0oC) separated by a ridge from warmer (6.4oC) Gulf-stream-influenced Atlantic waters. Fauna varied with water temperature
H.M.S. Porcupine (1869-70) First ship specifically equipped for oceanographic studies in deep water (first fully organized oceanographic expedition) Carpenter and Thomson - chief scientists West of Ireland - dredged to 2700 m South of England/France - 4450 m Mediterranean Sea w of Spain Recovered all major groups (mollusks, crustaceans, echinoderms, sponges, stalked crinoids, banks of Lophelia pertusa, primitive urchins - Living fossils 1st use of protected thermometer Led to hypothesis of density-driven, deep-water circulation. W. Thompson
H.M.S. Challenger (1872 – 1876)Beginning of modern oceanography(Institutional, collaborative, multidisciplinary) C. Wyville Thomson – Expedition leader + 4 naturalists (Murray, Buchanan, von Willimoes-Suhm, Moseley) + 1 artist (Wild) • 3.5 years • 68,890 nautical miles • 362 stations • 40% time spent in ports 19th century equivalent of the US space program in 20th century
Challenger Expedition Evolved from a one page proposal! Objectives - To map: 1. Global patterns of deep-water circulation 2. Chemistry of world’s oceans 3. Geology of the deep-sea floor 4. Distribution/abundance/origin of deep-sea organisms Determine: chemical composition of seawater, physical conditions of the deep sea, characteristics of sediment deposits, distribution of organic life
H.M.S. Challenger (226 ft corvette) • Dredging equipment • Winch • Dredging platform • Accumulators (safety springs) • Beam trawl
H.M.S. Challenger • Natural history laboratory
Primordial ooze • Origin of life from inanimate matter • Bathybius haeckelii primordial ooze • Missing evolutionary links between fossil and modern organisms In 1858 Huxley had found gelatinous matter - identified as Monera and called Bathybius haeckelii - formed living sheet over much of the ocean and provided food source for higher organisms Bathybius haeckelii, 1868-76. Viewed under the microscope the small discoids are the exoskeletons of tiny sea creatures, while the jelly within which these are suspended is the gelatinous gypsum precipitate. www.creationism.org/ books/TaylorInMindsMen Alcohol + water yielded (Gypsum - Ca Sulfate)
H.M.S. Challenger Expedition Departed Portsmouth – 21 Dec. 1872 South to Madeira, St. Thomas, Bermuda, Halifax, Azores, Cape Town, Kerguelen to Antarctic pack ice, Melbourne, Fiji, Hong Kong. Philippines, Japan, Hawaii, Tahiti, Valparaiso, Falkland Islands. Arrived Portsmouth – 24 May 1876
Challenger Expedition Results • Animals collected throughout the ocean to 5500 m depth • Many deep-sea species of many taxa, high proportion of rare species, many with direct development • Decreasing abundance and diversity with depth - generated paradigm of the Depauperate Deep. • Different taxa in deep than shallow water (zonation) • Stability of deep-sea environment (temperature, chemical composition, lack of seasonal changes) • Constant seawater constituent ratios. • Deep-sea sediments (calcareous, siliceous oozes) of pelagic origin (foraminifera, radiolarians, coccolithophores, pteropods, diatoms, etc) • Red clay of terrestrial origin in central oceans, manganese nodules rich in metals • Description of water masses based on T and S • Description of shelf , cont. slope upper and base
Theories Laid to Rest • Azoic Theory disproved. Animals present throughout the deep sea to 5500 m one sample at 7000 m Japan Trench. • Huxley’s Bathybius -(artifact of preservation) addition of alcohol to seawater caused precipitation of gypsum. • No living fossils - trilobites or Belemnites [extinct cephalopods] Rather deep fauna evolved from continental shelf and slope forms … relatively recent - with onset of Glaciation in Cenozoic. • No large, cosmopolitan deep-sea species, but genera widely distributed (5-7% at high and low latitudes, 3-4% had bipolar distributions). 1 species in common between Pacific and Atlantic at mid equatorial latitudes.
Challenger Results Scientific results published in 50 volumes with final summary by John Murray (1895- 13 years after Thomson’s death at 53). 29,500 pages 3,000 plates Samples distributed to experts globally a. high species diversity in deep-sea b. common taxa in high latitudes c. greater depth ranges for deeper species, sharper zonation in shallow water d. endemism common in deep water e. first proof of deepwater plankton (between 915 and 1830 m depth) f. Discovery of mid Atlantic Ridge
Post-Challenger Expeditions Alexander Agassiz (1877- 1888) USA a. Steamer Blake in Gulf of Mexico and Caribbean. b. replacement of hemp with wire rope (less deck volume, more efficient handling) c. abundant fauna dredged to 3567 m depth d. existence of midwater plankton e. reiterated question of food supply to deep sea – sinking of plankton, role of terrestrial debris. f. caught nothing below 200 m (100 fathoms) with Sigsbee gravitating plankton net
Italian circumnavigation by the Vetter Pisani (1882-1885) • Chierchia and Palumbo using a crude opening-closing net discovered deep-water plankton (i.e. siphonophores) to 2300 m depth. • Chun, inspired by these studies, sampled off Naples finding a rich pelagic fauna exists to 1400 m depth – hypothesized a “ladder of vertical migrations” (Chun, 1887).
Valdivia expedition (1898-1899) • Carl Chun expedition leader • European and African coasts to Antarctica, Indian Ocean, Red Sea, Suez Canal, Mediterranean Sea to Hamburg • Opening/closing nets revealed extensive deepwater pelagic fauna to 2000 m depth – below which fauna became sparse.
French Oceanographic Expeditions • Travailleur – (1880) – Bay of Biscay • Talisman – (1888-1927) – E. Atlantic, Mediterranean. • Distinctive deep-water ichthyofauna, with vertical migrators • Distinctive abyssal molluscan fauna originating at high latitudes • Homogeneity of deepwater fauna across the Atlantic • Ancient annelids are widespread, modern ones are not.
Prince Albert of Monaco • Series of cruises on Hirondelle, Princess Alice, Princess Alice II, between 1885 and 1914. • Technological advances using wire rope, steam winches, large closing trawls, baited traps to abyssal depths • Areas of study – N.E. Atlantic, Mediterranean • Circulation of N. Atlantic studied with drifters • Sardine fishery off N. Spain, marine mammals • Successful trawling to 6035 m* off Cape Verde • Confirmed vertical migrations by deep pelagic fauna. • Used baited traps - confirmed existence of scavengers at depth (lysiannasid amphipod 14 cm long) • Once worked a 120 h continuous station at 5940 m off Portugal *deepest until 1947
Prince Albert - cont. • Significantly advanced technology to study deep-sea communities • Migrating bathypelagic fauna exists supporting Chun’s “ladder of migrations” hypothesis concerning transport-food supply to deep ocean • Unique combination of creativity and financial independence.
Pre WWI Michael Sars cruise (1910) • a European collaboration • John Murray (Great Britain) and Johan Hjort (Norway) – N.E. Atlantic a. Extensive pelagic trawling, minimal benthic trawling (5160 m) b. Further speculation on sources of nutrition for deep sea- phytoplankton – zooplankton – detritus-dissolved organic matter c. Influential text on oceanography (Murray and Hjort –1912- The Depths of the Ocean.
WWI through WWII • From 1910 until after WWII – de-emphasis of deep-sea research • More emphasis on coastal fisheries problems and plankton dynamics
Post WWII 1. Swedish Deep-Sea Expedition(1947-1948) • Circumnavigation in Albatross – • Seismic studies, piston coring, bottom water sampling • only last 3 months devoted to deep-sea benthic trawling • Benthic trawling in Puerto Rican Trench – to 7900 m – deepest trawl in N. Atlantic
Post WWII Danish Deep-Sea Expedition – Galathea(1950 – 1952) around the world. - Anton Bruun – expedition leader • 1st thorough study of abyss & hadal zones • 1st microbiology in the deep sea • 1st quantitative abyssal samples (grab) • Trawled to 10,190 m in Philippine Trench (sea anemones, amphipods, isopods, bivalves, holothurians). • - Trawled in five trenches recovering 115 species > 6000m depth. Found a distinct hadal fauna. • - Isolation of barophilic bacteria from deep-sea (Zobell and Morita, 1959) - First use of 14C to estimate primary productivity (Steeman Nielsen) Reports published even in 1980’s
Cold war era Russian deep-sea expeditions (1950s) - Vitiaz – extensive grab sampling to determine benthic biomass in deep basins and trenches of Atlantic, Pacific and Indian Oceans (Zenkevitch, 1963; Belyaev, 1972). Emphasis on feeding ecology
Modern era 1949: there were less than 100 oceanographers in the USA 1959: Oceanography budget 21 million 1969: Oceanography budget 221 million. In those 10 years: 20 new vessels and 8 new laboratories American non-expedition studies (1960’s) - Strong financial support from U.S. government - Establishment of Gay Head Bermuda transect (55 to 5000 m depth) – Howard Sanders and Bob Hessler in 1965 - Anchor dredge, epibenthic sled, finer sieves - Higher abundance, greater species diversity
Galathea 3 - Danish Vaedderen “the ram” ( 2006-2007) Ram’) 3 • Antifreeze protein systems in Antarctic fish • Avian diversification • Biodiversity in protista • Biological interaction on islands • Collecting poisonous sea snakes • Deep-sea fish at the Antarcti c • Dissolved Organic Matter • Enzymes from the Ikaite columns in Greenland • Fluorescent proteins • Gingers • Oceanic oxygen deficiency zones • Parasites in zooplankton • Plankton dynamics in the Andaman Sea • Plant communities in the Galapagos Islands • Roseobacter bacteria – the ocean’s stars • Sea turtels in the major sea current systems • Sound in the Oceans • The Benthic Fauna of the Solomon Sea • The DNA of the Polar Seas • The European Eel • The Horseshoe Crab • The Marine Carbon Cycle • The origin of the vertebrate immune system • The physiology of antarctic fish • The significance of the climate the degree of isolation on biological interplay and biodiversity in lakes