2

The physical environment of the deep sea

THE SHAPEOFTHE

SEAFLOOR

The topography of the floor of the deep ocean (Fig. 2.1) is a balance between the parameters of seafloor spreading and sedimentation of inorganic and organic particles. Around the periphery of the ocean basins lies a continental shelf of varying width ending at the shelfbreak, usually at a depth of c. 200 m, below which plant lifeis supposed to be absent. ln the Antarctic, owing to the weight of the ice cap, the shelf edge is at c. 500 m. If we accept topographic criteria, the deep sea may be said to begin at the shelf break. This is a safer distinction than one based on photosynthetic depth since attached seaweeds have been found as deep as 268 m off the Bahamas (Littler et al., 1985). Seaward of the shelf edge there is a marked increase in the downward gradient of the seabed indicating the continental slope (Fig. 2.2). The slope may be a simple structure where the isobaths are parallel to each other and evenly spaced or it may contain a series of irregularities to give a very uneven gradient. The slope marks the underlying boundary between oceanic and continental crust. The theory of plate tectonics tells us that these areas of crust consist of a dynamic system of plates where crust is both being formed at mid-ocean ridge spreading centres and consumed by subduction at 'active', or seismic margins typical in the Pacifie (Fig. 2.3). 'Passive', or aseismic margins are typical of the Atlantic (Leeder, 1985). The gradient of the slope may be interrupted by terraces and submarine canyons. The latter appear as irregular, fissure-like channels cut down the continental slope which may act as conduits for transport to the deep ocean basin beyond, but probably were most active as such during glacial periods when sea levels were lower, and downslope processes far more intense than today. However, bottom currents, intense enough to resuspend the sediment, may occur from internal tides focussed along the canyon axis (Gardner, 1989).Their V-shaped profiles are probably the result of erosion by turbidity currents (see p. 24).Many can be traced on to the adjacent continental shelf, often at the mouths of major rivers.

10

1. HISTORY, ENVIRONMENT

AND METHODS
FSC
WTR

NB
RP

Fig. 2.1. Topography of a section of the northern N.E. Atlantic lying to the west of the British Isles showing sorne of the chief physiographic features of the deep ocean. The view looks northeast and shows the rugged topography of the Reykjanes Ridge (RR) extending southwards frorn Iceland (I), and the E.-W. trending Charlie-Gibbs Fracture Zone (GFZ) that separa tes it frorn the northern section of the MidAtlantic Ridge (MAR).The westward flank of the Reykjanes Ridge extends into the part of the Labrador Basin lying east of southern Greenland; while the eastward flank of the Ridge rnerges into the gentler topography of the Iceland Basin (IB).This basin is bounded to the E. by a chunk of alrnost cornpletely subrnerged continental crust, the Rockall Plateau (RP) and its associated northern Banks. continued

At the base of the slope of passive margins there is typically a thick wedge of slope-derived sediment termed the continental rise (Fig. 2.2) or, if the base of the slope is offshore from a large river, there may be a mu ch larger formation of alluvial sediment called a submarine fan. The topography of the rise is usually much smoother than the slope, but may be cut by channels extending from canyoned slopes. By a depth of c.4 km the seabed has levelled off to give a wide expanse of relatively flat abyssal plain which extends gently from 4 km to 6 km depth. These are often undulating or quite featureless, or they may be interrupted by numerous flat-topped guyots or seamounts (Fig. 2.1), which are inactive ocean-floor volcanoes that do not rise above sea level, and sometimes occur in chains (Epp & Smoot, 1989), Seamounts can rise several kilometres above the ocean floor and their profiles show declivities as great as 25, much steeper th an any major seafloor feature elsewhere in the ocean. The abyssal plains do not extend across the oceans but are separated by the mid-ocean ridge (Figs 2.1,2.3,15.2). The ridge is the site of formation of new ocean crust and is a more or less continuous system occupying about 33% of the area of ocean floor. The generally symmetrical process of extrusion of new crust along both sides of the mid-ocean ridge results in the separation of the flanking lithospheric plates. The solid geometry of the Earth necessitates the formation of a series of cracks, called transform faults, which appear as great slashes at right angles to the main axis (Fig.

PHYSICAL ENVIRONMENT
This 'microcontinent' is separated from the continental crust of northern Europe by the Rockall Trough (RT). Shallow sills, including the lceland-Faeroe Rise (IFR) and the Wyville Thomson Ridge (WTR) that separate the Norwegian Basin (NB) from the more southerly Atlantic basins allow overflow of cold Arcticcooled water from the Norwegian Sea into the Iceland Basin and Rockall Trough to contribute to the deep-sea water mass via the Faroe Shetland Channel (FSC) (see p. 15).Two seamounts, the flat-topped Anton Dohrn Seamount (ADS) and Hebridean Seamount (HS) lie on the continental rise of the eastern Rockall Trough. The latter basin opens into the deeper Porcupine Abyssal Plain (PAP). The steep continental margin lying southwest of Ireland is broken by a large bightlike terrace, the Porcupine Seabight (PSB)and further . by numerous canyons on the continental slope of the Bay of Biscay (BB). (Computer-generated chart courtesy Dr G. Robinson, NERC Unit for Thematic Information Systems, Reading University.)

11

2.3).Mid-ocean ridges are usually about 2.5 km below sea level, but, with increasing distance from the ridge, depth increases to about 5 to 6 km with the depression of the thin ocean crust by an ever-increasing accumulation of pelagie sediment blanketing the uneven topography of the oceanic crust. Here lie the featureless expanses of the abyssal plains with gradients typically in the region of 1 : 1000. Trenches occur if the abyssal plain is bordered by an active margin, when the oceanic crust (the lithosphere) buckles and deepens as it is eventually destroyed by subduction beneath an adjacent continent (Fig.2.3).Trenches are best developed in the Pacific Ocean where 'active' continental margins are dominated by subduction zones. As a result, the Pacifie, in contrast to the Atlantic, is shrinking in size despite the relatively high seafloor spreading rates found there. Trenches are seismically active areas that are best developed when associated with island arc systems where one plate is subducted under an adjacent oceanic plate (Fig. 2.3).Trenches are greater than 6 km deep and can extend to depths greater than 11 km in the case of the Challenger Deep in the Mariana Trench. The relative proportions of the Earth' s surface at any given level may be shown by a hypsographic curve (Fig. 2.4(b)) which should not be confused with its superficial resemblance to the profile of a passive continental margin (Fig. 2.2). The terms applied so far are physiographical whilst the ecological depth zones associated with them are labelled: (i) (ii) (iii) (iv)
sublittoral, or subtidal- Low water mark to 0.2 km bathyal, or archibenthal- 0.2 to 2 km abyssal- 2 to 6 km hadal, or ultra-abyssal- >6 km .

This terminology will be used throughout this book for the general description of depth zones. However, as we shall see, this depth zone
Vertical exaggeration
1
1 1

x4
f f

0
1 1..
1

100

200

3bo:
1
1

17
1 1

Iii

400

500

600

700

1000 1100

----

1 1
1

1 1
1 1 1

Continental margin

1 1

E ..:.:
..<::'

1 2 3 4

D..
<I.J

Shelf /' break Continental Shelf ]

1:::

el

Vertical exaggeration
<I.J

x 50

.::
0
U

Fig. 2.2. Profile of typical passive (aseismic) continental margin. (From Anikouchine & Sternberg,
1973.)

5 6 0 100 200 300

Abyssal Plain 800 900 1000 1100

12

I. HISTORY, ENVIRONMENT
ccr

AND METHODS

Fig. 2.3. Tectonic structure of ocean basin showing movement (arrows) of oceanic crust (ocr) overlying the mantle (ma), that is in tum overlain by the rafted continental crust (ccr) of the continent (co) along the passive margin shown on the right. The linear pattern of the spreading centre at the mid-ocean ridge (mor) may be offset by transform faults (tf). The zone of crust subduction found at the trench (tr) along the active margin on the left is shown bordered by the volcanoes and volcanic islands (vi) typical of the western
Pacifie,

terminology cannot be rigidly applied, and vertical zonation of fauna, which in the deep sea seems determined much more by a complex of sometimes interacting ecological factors th an by simple physical variables associated with the depth gradient, needs to be described in the deep sea by multivariate quantitative methods (see Chapter 9). DEEP WATER MASSES AND THEIR FORMATION ln terms of topography, the deep sea starts at the edge of the continental shelfbut in terms of hydrography it is usually considered to be that region below the permanent thermocline (Fig. 2.5). This latter is the transition layer in the water column in which temperature falls rapidly with increasing depth until values below 4C are reached and the downward temperature gradients become small. ln most of the world ocean this more stable temperature regime is entered at 0.8 to 1.3 km depth, except in the N. Atlantic where the injection of Mediterranean outflow at intermediate depths depresses the 4C isotherm to about 4 km. As

10 8
6

(a)

10 8
6

(b)

4
2

km

Maximum height (8.85 km) High mountain ranges Mean height of land (0.8 km) ../ Continental shelf
1

2
~--,--::-::,.--J

km 0
Fg. 2.4. Distribution of Earth' s surface lying at different levels: (a) frequency distribution, (b) cumulative-frequency curve based on (a) termed the Hypsographic Curve. This should not be confused with the superficially similar profile of the continental margin shown inFig.2.2. 2

Sea level

4
6

1 1 L

Mean depth of oceans (3.2 km)

~--'---="::':";

Ir:--=-----'

8 10L--L--~__L--L __ 10 20 30 40 Percentage of earth's surface

20

40 60 Cumulative percentage

80

PHYSICAL

ENVIRONMENT
Temperature, C

13

-5 0.5 1.0

la

15

20

25

la

15

20

--' Seasanal thermacline

--

E
..r:::'

~
QJ

1.5

el 2.0
Fig. 2.5. Vertical temperature profiles for different latitudinal zones.

2.5 3.0

High latitudes

Law latitudes

the upper waters are heated by solar radiation (in mid-latitudes with a seasonal cycle to form a more superficial seasonal thermocline), freshened by coastal runoff, and mixed by wind, the permanent thermocline isolates the deep ocean from the direct effect of surface parameters. Because of the much greater depth of the mixed layer as a result of wind stress (particularly evident in the N.E. Atlantic), the deep water mass below the permanent thermocline deepens at mid-latitudes, with a concomitant depression in isotherms away from the Equator. They shoal again at higher latitudes towards the areas of deep water formation nearer the poles (Fig. 2.6). This has had a marked bearing on interpreting the depth-related distributions of deep-sea animals in terms of temperature (see Chapter 9). However, the water below the permanent thermocline is not of constant temperature and salinity. Most of the floor of the world's major oceans is bathed in water masses which originally formed in either the Antarctic or in the Greenland/Norwegian Seas of the Arctic Ocean (Fig. 2.7). For surface water to become dense enough for it to sink to the bottom of the ocean, it must become either more saline by evaporation and ice formation or colder by heat loss. The water th en sinks to its new density

a
Fig. 2.6. North-South section of western Atlantic showing distribution of temperature (0C). This illustra tes the sharp gradient in temperature near the Equator with a trend to submergence of isotherms towards the poles where the water mass becomes increasingly isothermal. (From Svedrup
et al., 1942.)
/

,
1
/

,,

- -,
..;L. ..r:::

2 3 4 5
S

::---4---,,-

1
/

- "

/1'

.......

el

QJ

70

60

50

40

30

20

la

10

20

30

40

50

60 ON

14

1. HISTORY, ENVIRONMENT

AND METHODS

Fig. 2.7. Main movements of water masses in a NorthSouth section of the Atlantic Ocean, showing the origin of dense, cold Antarctic Bottom Water (AABW)from the Weddell Sea (left), which can be traced as far as 40 N, and the overlying cold, less dense Antarctic Intermediate Water (AAIW).North Atlantic Deep Water (NADW) originates in the Norwegian Sea (right) near Greenland from sinking of a mixture of dense surface water from the Gulf Stream with cold Arctic water. Warm, dense Mediterranean Water (M) intrudes from the E. from the Straits of Gibraltar. A similar northward-flowing pattern exists in the Pacifie, but deep flow from the N. is only poorly developed. (From Turekian, 1976.)

level where it spreads out. To form the very deepest waters in the ocean, the surface layers must become very cold. It eventually returns to the surface by gradual upwelling until, in time, it all returns to the surface as return flows from low to high latitudes. The deepest waters found in the ocean are formed in the cold surface layers close to the coast of Antarctica especially in the Weddell Sea where winter-cooled surface water is exceptionally cold (-1.9 "C). This water (Fig. 2.8) mixes with the upper part of the saline warm deep water (Circumpolar Water) to give a modified deep water that flows along the western shelf edge of the Weddell Sea along the Palmer Peninsula, mixing with Western Shelf Water. This proceeds to sink downslope as Weddell Sea Bottom Water. This water then mixes with deeper saline waters of the Circumpolar Water to form Antarctic Bottom Water (AABW)(Mantyla & Reid, 1983). Antarctic Bottom Water is generally accepted as a generic term for water masses of very similar characteristics (-0.4 "C: 34.660/00) which are formed in the Ross Sea and off the Adelie Coast and contribute to this bottom water. AABW sinks to form a circumpolar bottom water with branches penetrating all the main oceans (Fig. 2.7). ln the Atlantic this water flows up the west side at depths >5 km but is prevented from doing so on the east side by the Walvis Ridge. Sorne of this water will penetrate the N.E. Atlantic by passing through the major fracture zones in the Mid-Atlantic Ridge. Branches of AABW extend into the lndian and Pacifie Oceans basins unless its spread is impeded by ridges such as the East Pacifie Rise.

Equator

..Y.

E 2
3 4

.. d
(j)

0..

-,-----

- ---

PHYSICAL ENVIRONMENT

15

,'"

\ ., .. :.>. ... '

z .............

~s"'~ , ,....

AABW

>2 km depth

Fig. 2.8. Formation of Antarctic Bottom Water (AABW) in the Weddell Sea. WSW, Western Shelf Water; WDW, warm deep water; SWW, surface win ter water; WSBW, Weddell Sea Bottom Water; MDWW, modified deep win ter water. Dotted line indicates shelf edge. See text for further details. (Modified from Warren, 1981.)

.......
"

-,
........

..,

Coats Land

Overlying the AABW and covering most of the abyssal plains of the world ocean is slightly less dense water, uItimately derived from the sinking of surface water in the Norwegian Sea, and termed North Atlantic Deep Water (NADW). Relatively warm (9-12C) and saline (35.3 to 35.5%0)water flows into the Norwegian Sea via the North Atlantic Current. This water is cooled: (i)by lateral mixing with polar waters and (ii) by heat loss to the very cold atmosphere in win ter, weakening the density gradient between the surface and bottom, destabilizing the water column. Vertical, convective cells develop and these generate deep and bottom water in the Norwegian and Greenland Seas. The water produced has a density (0/) of 28.1 resulting in a virtually homogeneous water column below 0.6 km. Sorne of this Norwegian Sea Deep Water flows north into the central Polar Basin via the Lema Trough. The rest spills over the Greenland-lceland-Scotland Ridge (Worthington, 1970; Warren, 1981) into the North Atlantic. ln the North Atlantic most enters via either the 0.8 km deep Faroe Bank Channel or the 0.45 km crest of the Faroelceland Ridge. Cold water also spills over the Wyville Thomson Ridge from the Faroe Bank Channel into the northern RockallTrough, as shown in Fig. 2.16. The route into the N.W. Atlantic basin is by way of the 0.6 km deep sill formed by the Greenland-lceland Rise in the Denmark Strait (Fig. 2.9; see Fig. 2.1 for details of topography). Norwegian Sea Deep

16

I. HISTORY, ENVIRONMENT

AND METHODS

Fig. 2.9. Formation and track of dense, cold deep water (NADW) in the Norwegian Sea and subpolar N. Atlantic from cooling and sinking (denoted by curled ends) of warm (>4 C) surface currents. Other, lighter, cold overflows are also shown. (From McCartney & Talley, 1984.) Arnerican Meteorological Society, 1984.

Surface currents Other light cold overflow

~ ::. 0:) NADW overflow

Water from the first two sources entrains overlying Atlantic water to form Northeast Atlantic Deep Water. The core gradually descends the contours of the continental slope south of lceland and then those of the eastern flank of the Reykjanes Ridge, eventually escaping westwards and northwestwards into the N.W. Atlantic basin at 1.5to 3 km depth through the Charlie Gibbs Fracture Zone at 53N (Fig. 2.1). There it joins the Northwest Atlantic Bottom Water formed from the Denmark Strait overflow, to produce the North Atlantic Deep Water (NADW), which flows south along the eastern continental slope of N. America at 1 to 5 km depth (Rowe & Menzies, 1968)in the Western Boundary Undercurrent. Although the NADW is more saline than AABW owing to the entrained upper waters, it remains less dense because it is warmer. Consequently wherever both water masses penetrate, NADW overlies the AABW. NADW spreads throughout the S.Atlantic and eastwards round S.Africa, and its high-salinity core may be traced into the northern Indian and Pacifie Oceans (Reid & Lynn, 1971). Although these water masses (AABWand NADW) coyer a significant part of the floor of the deep ocean there are other deep waters that bathe restricted parts of the ocean floor. For example, the European Basin of the N.E. Atlantic receives only a little NADW through the Gibbs Fracture

PHYSICAL ENVIRONMENT

17

Zone, where the deep flow is predominantly westwards. Direct evidence of AABW presence in the deepest parts of the basin is restricted to anomalously high silicate content (Mann, Coote & Garner, 1973).Northeast Atlantic Deep Water, derived from the easternmost ScotlandGreenland Ridge overflows, occurs where bottom depths are 2 ta 3.5 km (Ellett & Martin, 1973;Ellett & Roberts, 1973;Ellett et al., 1986)and in the shallower northern areas Labrador Sea Water (LSW) of lesser density covers mu ch of the seabed in the depth range 1.5 to 2 km (Lonsdale & Hollister, 1979).LSWis formed by deep winter mixing south and west of Greenland, and spreads eastwards at intermediate depths, where its density is somewhat greater than the much more saline but warmer Gibraltar, or less precisely 'Mediterranean' water (Cooper, 1952). Gibraltar Outflow Water is formed by the subsurface Mediterranean outflow plunging into the Atlantie from the Gibraltar Sill. It spreads northwards and westwards and blankets much of the continental slope west of Europe. It reaches to depths of 2.45 km in the south (Meincke et al., 1975), but to the west of the British Isles has a more restricted depth range of 0.8 to 1.2 km. There are no major sources of deep water in the northern Pacifie.This is because the surface waters in the N. Pacifie are of such low salinity that even intense wintertime cooling does not increase their density enough for them to sink (Warren, 1981). ln the Mediterranean, the opposite situation creates conditions for deep water formation. The surface water flowing into the Mediterranean through the Straits of Gibraltar is relatively saline, it flows eastwards along the North Afriean coast with branches travelling northwards. Within the enclosed basin, evaporation is high so salinities increase. ln winter, in the northern Aegean, Adriatie and Ligurian Seas, cold offshore winds cool the high salinity surface water, causing massive overturning and the formation of very dense deep and bottom water. However, this is not the water mentioned above as providing a mid-water high salinity core in the Atlantic; this is the Levantine Intermediate Water (Wst, 1961) formed with lesser density off the Turkish south coast in win ter, and which flows west to the Gibraltar Sill at 0.2 to 0.6 km depth. The main ecological significance of these deep water masses lies not in their salinity/temperature characteristies but in that they are weIl oxygenated. AlI the se water masses form at the surface so that their oxygen content is in equilibrium with the atmosphere. After they sink, although the oxygen is slowly used up by metabolic processes, with the exception of limited areas such as the oxygen minimum zone and anoxie basins, e.g. the Black Sea, the supply of oxygen in the deep waters is sufficient to maintain the surficial sediments of the world' s oceans in an oxidized state. ln the open ocean, oxygen concentration near the seabed decreases northward in the Indian and Pacifie Oceans as these areas are the most remote from the supply of oxygenated water at the main site of deep water formation in the North Atlantic (Mantyla & Reid, 1983).Beneath regions of high production (e.g. eastern tropical Pacificand in the Arabian Sea) oxygen minima form whieh can lead to anoxie conditions in bathyal

18

1. HISTORY, ENVIRONMENT

AND METHODS

sediments. ln sorne regions, such as the Mediterranean, black layers in the sediments called sapropels are considered to have been derived during similar anoxie conditions during previous eras. PHYSICAL PROPERTIES OF THE WATERS COVERING THE DEEP OCEAN FLOOR The main feature of the physical properties in the deep sea is that, with the exception of hydrostatic pressure and current energy, these parameters show a very narrow range at any specific site below the permanent thermocline. Unlike coastal waters, solar radiation has no direct ecological significance, as alilight (except bioluminescence), has disappeared by 1 km depth. It does, however, have an indirect effect by being the energy source for surface phytoplankton production, sorne of which enters the deep sea ecosystem via the food chain (see Chapter 11). TEMPERATURE AND SALINITY

The temperature of the waters of the deep sea varies from 4C to -1C (Svedrup et al., 1942).Exceptions are the Mediterranean which is c. 13C between 0.6 and 4 km, the Red Sea where the bottom temperature can be 21.5C at 2 km depth, and the very high temperatures in the immediate vicinity of hydrothermal plumes (see Chapter 15). The lowest temperatures found are -109C in the deep waters of the Antarctic. The salinity is also relatively constant and below 2 km is close to 34.80/00 0.3%0, declining to 34.65%0 at the very deepest levels (Svedrup et al., 1942;Menzies, 1965). OXGYEN CONCENTRATION The oxygen values are near saturation except where the oxygen minimum layer, found at 0.5-0.6 km depth in the open ocean (Fig. 2.7), impinges on the upper continental slope, and in enclosed basins such as the Black Sea which, below 250 m, is anoxie and azoic. However, as the deep water masses progress further from their site of origin, oxygen will be consumed by metabolic processes, and water in the deep N. Pacifie has a relatively low oxygen concentration of 3.6 ml 1-1 (Mantyla & Reid, 1983).There is sorne evidence (Bruun, 1957)that, immediately above the deep-sea bed, there is a slight reduction (0.15 ml 1-1) in oxygen concentration. HYDROSTATIC PRESSURE

The most predictable physical variable is hydrostatic pressure. This increases by 1 atmosphere (1 bar or 105 pascals) per 10 m increase in depth. This increase in pressure, particularly in relation to the low temperature, affects the rates of enzymatic catalysis in deep-sea organisms (Somero et al., 1983).

DEEP-SEA BIOLOGY: A natural history of organisms at the deep-sea floor

John D. Gage
Scottish Marine Biological Association Dunstaffnage Marine Laboratory Oban, UK

Paul A. Tyler
Department of Oceanography University of Southampton Southampton, UK

CAMBRIDGE
UNIVERSITY PRESS