SIO 277Deep-Sea Biology

Deep-Sea BiologySIO-277

• Instructor – Lisa Levin• Contact: llevin@ucsd.edu, 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.1Sept. 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

Heterogeneous bathymetry

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

Continental Shelf

Upper Continental Slope

Lower ContinentalSlope

Continental Rise

Abysss

meters200

500

1000

3000

4000

6000

Epipelagic (euphotic)

Mesopelagic (disphotic)

Bathypelagic (aphotic)

Benthopelagic

Bathyal

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 beginning1. 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 questionL.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 scientistsWest of Ireland - dredged to 2700 mSouth of England/France - 4450 mMediterranean 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 thermometerLed 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

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

Evolved from a one page proposal!

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. 1872South 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 Results1. Animals collected throughout the ocean to 5500 m depth2. Many deep-sea species of many taxa, high proportion of

rare species, many with direct development3. Decreasing abundance and diversity with depth -

generated paradigm of the Depauperate Deep. 4. Different taxa in deep than shallow water (zonation)5. Stability of deep-sea environment (temperature, chemical

composition, lack of seasonal changes)6. Constant seawater constituent ratios.7. Deep-sea sediments (calcareous, siliceous oozes) of

pelagic origin (foraminifera, radiolarians, coccolithophores, pteropods, diatoms, etc)

8. Red clay of terrestrial origin in central oceans, manganese nodules rich in metals

9. Description of water masses based on T and S10. Description of shelf , cont. slope upper and base

Theories Laid to Rest

1. Azoic Theory disproved. Animals present throughout the deep sea to 5500 m one sample at 7000 m Japan Trench.

2. Huxley’s Bathybius -(artifact of preservation) addition of alcohol to seawater caused precipitation of gypsum.

3. 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.

4. 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 platesSamples distributed to experts globally

a. high species diversity in deep-seab. common taxa in high latitudesc. greater depth ranges for deeper species, sharper zonation in shallow waterd. endemism common in deep watere. first proof of deepwater plankton (between 915 and 1830 m depth)f. Discovery of mid Atlantic Ridge

Post-Challenger Expeditions

Alexander Agassiz (1877- 1888) USAa. 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 depthd. existence of midwater planktone. 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

1. Travailleur – (1880) – Bay of Biscay

2. 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.

• 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 of Monaco

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 WWIIDanish 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 eraRussian 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 era1949: there were less than 100 oceanographers in

the USA1959: Oceanography budget 21 million1969: Oceanography budget 221 million. In those 10 years: 20 new vessels and 8 new

laboratoriesAmerican 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

Observatories(e.g., Neptune Canada)