Oceanus

Volume 23, Number 2, Summer 1980

Oceanus

The International Magazine of Marine Science

Volume 23, Number 2, Summer 1980

William H. MacLeish, Editor
Paul R. Ryan, Managing Editor
Deborah Annan, /Ass/sfanf Editor

Editorial Advisory Board

Edward D. Goldberg, Professor of Chemistry, Scripps Institution of Oceanography

Charles D. Hollister, Dean of Graduate Studies, Woods Hole Oceanographic Institution

Holger W. Jannasch, Senior Scientist, Department of Biology, Woods Hole Oceanographic Institution

Judith T. Kildow, Associate Professor of Ocean Policy, Department of Ocean Engineering,

Massachusetts Institute of Technology

John A. Knauss, Dean of the Graduate School of Oceanography, University of Rhode Island

Robert W. Morse, Senior Scientist, Department of Ocean Engineering, and Director of the Marine Policy

and Ocean Management Program, Woods Hole Oceanographic Institution

Ned A. Ostenso, Deputy Assistant Administrator for Research and Development, and Director, Office of Sea Grant,

at the National Oceanic and Atmospheric Administration

Allan R. Robinson, Gordon McKay Professor of Geophysical Fluid Dynamics, Harvard University

David A. Ross, Senior Scientist, Department of Geology and Geophysics, and Sea Grant Coordinator, Woods Hole

Oceanographic Institution

Derek W. Spencer, Associate Director for Research, Woods Hole Oceanographic Institution

Allyn C. Vine, Senior Scientist Emeritus, Department of Geology and Geophysics, Woods Hole Oceanographic Institution

Published by Woods Hole Oceanographic Institut

Charles F. Adams, Chairman, Board of Trustees
Paul M. Fye, President of the Corporation

Townsend Hornor, President of the Associates

<Mk iHPt mtmmmillmSmWJmM A\

John H. Steele, Director of the Institutioft 4MK^HBJSfl^^^H9faaflfl^^

mm

The views expressed in Oceanus are those of the authors and do not
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Oceanus

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Contents

PATTERNS IN PLANKTON

by John H. Steele

Understanding the dynamics of how plankton is distributed in the ocean gives scientists a

clearer picture of its important role in the food chain. 2

EL NINO: LESSONS FOR COASTAL FISHERIES IN AFRICA?

by Michael H. Glantz

Is there a point where too much scientific information about a fishery tends to obscure the

social, political, and economic issues? 9

SUBMARINE HYDROTHERMAL ORE DEPOSITS

by Michael J. Mottl

On the East Pacific Rise at 27 degrees North, hot springs are venting waters at temperatures

as high as 350 degrees Celsius that are precipitating prodigious quantities ofsulfide ore,

minerals rich in copper, zinc, and iron.

!**»

GEORGES BANK: THE LEGAL ISSUES

by Daniel P. Finn

The Georges Bank oil and gas lease sale has raised critical questions about the overlapping

responsibilities among federal agencies and their ability to take effective action where

significant marine resources are at stake. 28

THE DEVELOPMENT OF MARINE SCIENCE IN LATIN AMERICA

by Francisco J. Palacio

Latin American nations have been relying too much on international agencies and

institutions to develop their marine programs. They must now look inward and develop

their own pool of human resources.

CILIA AND CTENOPHORES

by Sidney Tamm

The mechanism of ciliary and flagellar motion is fundamental to our understanding of

various life processes, as well as disturbances in these functions caused by disease or

genetic defects. Ctenophores are the largest known animals that use cilia for

locomotion.

FRONT COVER: Artist E. Kevin King's rendering of th e submersible Alvi n , opera ted by the
Woods Hole Oceanographic Institution. The submarine's latest exploits are described on
page 18. BACK COVER: "Black smoker" on the East Pacific Rise (see page 24). Photo by
Robert Ballard.

Copyright © 1980 by Woods Hole Oceanographic Institution. Oceanus (ISSN 0029-8182)
is published quarterly by Woods Hole Oceanographic Institution, Woods Hole,
Massachusetts 02543. Second-class postage paid at Falmouth, Massachusetts, and
additional mailing points.

1

\

Patterns in Plankton

by John H. Steele

riuman beings are not distributed uniformly over
the surface of the earth. Ourdensity has patternson
many different scales. Populations are generally
sparse in deserts and dense where the soil is
productive; or they tend to concentrate along
coastlines. And, on very small scales, we build
houses in relation to particular terrains or water
supplies. Yet, outside these external constraints
and between the largest and smallest scales, we
ourselves determine much of the pattern of human
activity. In the past, the size of market towns and the
distance between them would depend on the
limited mobility of two or four feet. Now, the
changing distributions within and between cities
are functions of the railroad, the car, and the
airplane. Thus, like all other organisms, we are
partly dependent, partly independent of our
physical and chemical environment; and the
consequences can be seen in the way populations
aggregate in towns, in herds, or on shoals.

The oceans may appear to be much more
uniform than the land. Marine organisms might be
expected to be spread more evenly through the
water than their terrestrial counterparts. Yet the
pelagic terrain can be as rugged as the earth's
surface. The temperature and salinity change with
increasing depth and also can change abruptly at
"fronts" that divide one part of the ocean from
another in the same way that a mountain range can
divide a forest from a desert. So we find that broad
regions of the ocean, such as the Sargasso Sea or the
Antarctic, can have very different levels of basic
productivity. And, at the other extreme, we have to
consider the behavior of individuals or small groups
on scales of meters or millimeters. Between these
limits, in the oceans as on land, we see patterns of
distribution on all scales and we ask the same
questions. Are these patterns determined by the

Unloading catch at Provincetown, Massachusetts. Man's
predation patterns have changed with the advent of new
technology. (Photo by Hank Simmons, PR)

physical environment, or do the organisms, in a
sense, create their own environment within their
food web, and is this seen in the way they are
distributed?

There are certain regularities in size and
composition of open-ocean food chains that we do
notfind on land. The plants living in the upper
layers of the ocean are microscopic, with diameters
usually in the range of 5 to 50 microns. The
herbivores that feed on these are crustaceans
ranging from the size of rice grains to small shrimp.
In turn, the animals that prey on these herbivores
can be small fish — or whales. Thus there is a regular
progression in size as we climb the trophic ladder
(unlike trees, which are grazed on by caterpillars).
Also, there is an increase in longevity from a few
days for the marine plants to several years or
decades for some of the predators. The increase in
length of life from days to months to years roughly
balances the decrease of approximately 90 percent
in productivity as we go one level up in the trophic
system . The result is that the standing stocks at each
trophic level may be about the same order of
magnitude (Sheldon and others, 1972). Butthis level
of biomass can be very different for large areas of
the ocean (Figure 1). It is these estimates of the
productivity of general ocean regions combined
with knowledge of the trophic level at which we
harvest that have provided information on the
maximum probable yield that we can take from the
sea by present conventional methods. The total
predicted yield of around 100 million tons (Ryther,
1969; Bell, 1978) comes almost entirely from coastal
and upwelling areas. This is true partly because
these areas are more productive than the open
ocean, but, more important, because in the former
regions we harvest at trophic levels much closer to
the primary plant production.

These general limitations on our yield from
the sea have been described in Oceanus, Vol. 22,
No. 1 , 1979. Also, there and elsewhere,
consideration has been given to the consequence

I

10-

10-

Phytoplanktor

— — — __ Krill

Whales

Zooplankton ~ — • —

Nekton

10'

10"

EQUIVALENT DIAMETER (m)

Figure 7. Estimates of standing stock in terms of particle
concentration for different trophic groups in two regions;
upper line, Antarctic; lower line, equatorial Pacific
(Sheldon and others, 1972).

of harvesting at lower trophic levels than our
present methods permit. If we accept the
large-scale average picture implicit in Figure 1 , then
there is the same or slightly greater concentration of
biomass at the lower levels, but these also will be
about ten times more productive. On this basis, if
fishing were merely a question of harvesting the
average organic matter in the water, then removing
smaller organisms should be as efficient as
capturing large ones, but since the overall
productivity is so very much higher, the total yield
could be correspondingly greater. It is this potential
that has tested the ingenuity of technologists. The
flaw in this picture is that we do not have an
environment with average concentrations. If we
did, predation would not be ecologically or
economically feasible. This is true not only for
fishermen but also for the natural predators in the
sea. Thus we need to consider the nature of the
variability and the limitations on aggregation for
different groups of organisms.

I have described the very large-scale
limitations on productivity imposed by the
environment. The physical and chemical conditions
present other small-scale limitations on the
concentrations of the primary producers — the
phytoplankton. Since the microscopic plants
depend on the nutrients in the water as well as on
light for their growth, the concentration of these
nutrients in the upper layer will generally determine
the maximum amount of plant material in, say, a
cubic meter of the sea. In coastal regions, the
maximum winter concentration of nitrate (10 to 20
milligram atoms per cubic meter) will give a crop of
about 10 grams biomass* per cubic meter,
equivalent to a concentration of 10~5. Figure 2
demonstrates both the dependence on nutrients
and the small-scale patchiness that can occur during
the early stages of a phytoplankton bloom.

*Assuminga biomass/nitrogen ratio of 70:1.

These plants have a relatively short life span
as individual cells, perhaps Mo 5 days, and have
very little capability of movement through the
water. By sinking, they can increase their
concentration, particularly when the water itself is
rising, and this can, very occasionally, produce
larger densities, such as those found in red tides
(seeOceanus, Vol.21, No. 3, p. 41).

In subtropical regions of the open ocean,
such as the Sargasso Sea, nutrient concentrations
are rarely greater than a tenth of coastal maxima and
usually very much less. Correspondingly, this
imposes a very much smaller upper limit on plant
concentration. In this way, the range of
concentrations for the base of the food chain is
closely defined by the physics and chemistry of the
environment. The small herbivorous Crustacea that
graze on this plant material have a longer lifetime,
on the order of 30 days, and are capable of vertical
motion, usually migrating from deeper water to
feed at night in the surface layers. Since the upper
and deeper layers can have different horizontal
motions, the vertical migration can providea means
for lateral movement and, in turn, permit
aggregations of these small animals in larger
patches. Patches on scales of about 30 to 50
kilometers are observed in the North Sea (Gushing
and Tungate, 1963; Steele, 1974) and can be
contoured by sampling with plankton nets (Figure
3). If we assume that, at night, they are in the upper
few meters, then their concentration can reach
levels of about 100 grams biomass per cubic meter
(Mackas and Boyd, 1979).

Pelagic fish or mammals, such as herring or
whales, feed on these herbivores. These organisms
are capable of directed movement — migrations

10

20

30

kilometres

'•./Nitrate

Spacing of net hauls

Figure 2. Small-scale patchiness of nutrients and
phytoplankton indicated by the fluorescence of the
pigments. These data were collected in early spring in the
surface waters of the North Sea. The inverse relation shows
the dependence of plant growth on a major nutrient,
nitrate. The lines along the bottom indicate the coarser
sampling obtainable with conventional plankton nets,
which will not reveal smaller-scale structure.

SURVEY I

SURVEY

COPEPOD
CARBON
(gC/m2)
2-O

O 2O 4O km

Figure 3. Larger-scale patchiness of phytoplankton
(chlorophyll) and herbivorous zooplankton (copepods) in
summer in the North Sea when the plant distribution is
more influenced by grazing and the general inverse
relations of plants and animals can be seen at scales
between 1 and 100 kilometers (Steele, 1974; Mackas and
Boyd, 1979). Surveys I and II were separated by a 3-day
interval.

that permit them to travel long distances to
aggregate on their food wherever it is at high
concentration. These patterns of migration are not
completely known, nor well understood. Many of
the pathways are along ocean fronts where there are
sharp changes in the physical character of the
environment. Thus the pelagic organ isms use these
signs to guide their movement around the ocean to
regions where food is abundant or to areas where
they aggregate for mating and spawning. In this
way, they make use of features in their
surroundings rather than being controlled by them
as are those organisms in the lower trophic levels
(Figure4).

Thus during their lifetime these predators
can be found over very wide areas of the ocean but
at any instant may be located in a small region.
Within this locality, they will occur in densely
packed schools or shoals so that a cubic meter of
water can contain several kilograms of fish.

For these reasons, we obtain a very different
picture when we consider maximum
concentrations rather than average levels. Perhaps
the best illustration of this difference is found in the
Antarctic when we compare the mean values of
Figure 1 with the maximum values on a cubic meter
basis in Figure 5. There is a relatively small increase
for the phytoplankton dependent, as described, on
the ability to convert nutrients into organic matter.
The herbivores, krill, are known to swarm in
relatively dense aggregations (Omori, 1978). Forthe
baleen whales, which are predators on the krill, if
we choose the appropriate cubic meter, then the
concentration of organic matter in that cubic meter
is unity!

- 60° N

4°W

Figure 4. The migration path of herring around the
northern North Sea, according to Cashing (1955). The
northerly movement in the spring is along the western
edge of less saline water from the Baltic that flows up the
Norwegian coast. The south and eastward fall migration is
along the general axis of Atlantic inflow (Dooley, 1974).
Thus much of the migratory circuit corresponds to current
systems, as well as to food concentrations (Steele, 1961).

rO

E
5 10

Q.

L

-2

o

c
o
O

o
Q.

10

.4

10

'6

/

/

Phyto

!

Whales

Krill

AVERAGE

io-6 io-4 io-2 i"

Particle diameter (m)

Figure 5. A comparison of average concentrations in the
Antarctic with the maximum based on possible
concentrations of organic matter in a cubic meter.

The choice of a cubic meter, which
represents a ton of water or biomass, is appropriate
forthe next predator, man, since this is the unit in
which he measures the "bites" that he takes from
the ocean whether this be herring, whales, or,
potentially, krill.

These features of increasing patchiness with
the steps up the trophic ladder illustrate how the
various populations have adapted to life in the
basically dilute medium of the open sea. It is
unlikely that any group could survive if it had to feed
on the average, so each has evolved to concentrate
where food is most plentiful, in turn supplying
adequate sustenance for the level above. Thus, as
we go from plants to herbivores to predators, we
see less response to the structure of their physical
surroundings and a corresponding ability to use the
larger-scale features of the ocean system such as
fronts or currents. In turn, this ability makes them
less directly dependent on local or short-term
events such as storms which will have marked
effects on phytoplankton production and

Krill is the food of whale and man alike: above, the
Antarctic species Euphausia superba (5 centimeters).

concentration. So the biological "behavior"
social or individual — of the higher trophic levels
plays a more dominant role.

On the other hand, if successful feeding can
take place only on above-average concentrations of
the food, then this can supply an ecological refuge
for the food organisms by removing predation
pressure on the below-average concentrations.
Thus the increasing patchiness with increasing
steps up the trophic ladder can provide both for
efficient energy transferand forpopulation survival
(FigureG).

Baleen whales, for example, feed on the
dense swarms of krill. Thus the growth rate of the
whales and, especially, the excess of food energy
for reproduction depends on the number and
density of these swarms and the whales' ability to
locate them. It is interesting that as the whale
population has declined the time taken to reach
maturity also has decreased, while their fecundity
has increased (Laws, 1977). This would indicate
some inverse relation between whale population
density and availability of the dense concentrations
of their food. Such a relation can help to produce a
natural balance between prey and predator
populations.

If, however, the pelagic predators, such as
herring, are assumed to be gathered in a localized
area and, within that area, concentrated in shoals,
then the low density refuge may not exist for
predation upon them. In this case, it appears that
the relatively large size of the shoals provides for the
population a defense against predators, such as
dolphins which are individually much smaller than
the prey unit if we take that unit to be a shoal. This
convoy theory appears to be a possible explanation
of how a population balance could be achieved with
natural predators operating near the top of the food
chain.

How does this work when we come to
another top predator — man? There are two reasons
why we harvest at or near the top of the natu ral food
chain. The organisms are relatively large and
therefore economically more desirable. Because of
their large size, they are easier to remove from the
water, especially when we do this by straining the
organisms from the water with a net. But, as I have
tried to show, the main reason is the existence of
increasingly dense aggregation of that organic
matter (Figure 6).

How does man himself fit into this pattern
proposed for other predators? How has he adapted
to his role as a top carnivore? Initially, his behavior
was similar to that of land-based predators, such as
seals, with a defined range and thus a localized
impact, spatially and seasonally. Also, the methods
of fishing, with drifting nets capturing individual
fish from a dense shoal, might not be too different
from those of the other natural predators, both

Fish

CONCENTRATION
OF BIOMASS

FRACTION OF TOTAL VOLUME

Figure 6. Schematic, and hypothetical, instantaneous
distributions depicting the relation between
concentration of three trophic levels and the fraction of the
upper layers of the ocean that they occupy. Thus, at any
instant, most of the pelagic fishing will occur in a relatively
small fraction of the total volume, whereas the
phytoplankton are more evenly spread.

capturing a significant fraction without attacking
the "tail" of the distribution (Figure 6).

The development of technology has changed
the nature of this predator not merely quantitatively
but also qualitatively. The fishingvessels have a
much greater range and capacity, but the main
feature is their ability to regard a whole school or
shoal as the unit of prey, searching for these units
with sonar and engulfingthem with a purse seine.
Thus there has been not only an increase in
efficiency but also a change in the nature of this
predator. This evolution, resulting from
technology, has been relatively rapid with no
noticeable adaptation by the individual prey
species. The consequences of the change in
predation pattern have been extreme for these
target species, such as herring or anchovy. Yet in
most cases the whole ecosystem appears to have
adapted by restructuring or rebalancing the food
web so that the energy made available is absorbed
in other pathways. In the North Sea, for example,
there seems to have been a replacement of herring
and mackerel, partly by smaller and less
economically desirable pelagic fish and partly by an
increase in groundfish such as haddock and whiting
which do not have such marked seasonal
concentrations in particular areas. In the Antarctic,
calculations of the present day energy flows within
the food web do not indicate a pronounced surplus
of krill. Rather, itappears likely that other predators
have replaced the whales.

The economic consequences to fisheries of
these changes have been severe, but what is
relevant here is the ecological and behavioral
patterns. The virtual collapse of these pelagic
fisheries is the result not so much of the number of
predators — the fishing boats — as of the ability of
these predators to take their prey as easily at low as
at high densities. This newly created imbalance
results not from changes in the average populations
of prey and predator over large areas but from
interactions on relatively small scales where a
change in "bite" size can switch the system from a
stable to an unstable relation. This switch depends
on the behavior of the prey. The fact that for herring
the catch per unit effort does not decrease
significantly as population levels fall, suggests that
with decreasing stocks, the fish still form the same
size shoals and are thus as easily captured.

Analogies are necessarily incomplete and
may delude ratherthan illuminate. Yet there is a
certain attractive formalism in the comparison of
successively higher trophic levels in the sea with
advancing human structures. There is an increasing
independence of the detailed components of the
physical and chemical environment. A greater
degree of social aggregation provides the basis fora
more efficient utilization of resources. Yet the
economies of scale can create their own problems.
For the seas, at least, it is apparent that our
understanding and our management of these
ecosystems depend on a knowledge not only of
the larger-scale averages but also of the smaller-
scale variability.

For these reasons, much of the interest and
attention in our research is focused on these
intermediate scales where the interactions of plants
and animals are partly but not wholly determined by
the physical conditions. My illustrations have been
taken from the distribution of phytoplankton
because we have methods to measure continuously
the concentration of their photosynthetic pigments
in the upper layers. But we know that part of the
variations we observe must be the result of grazing
on the plants by the herbivorous crustaceans. It has
proved more difficult to obtain comparable data on
this component of the food web, in part because
their vertical movements preclude using
measurements at one depth as an index. We have

Herring

certain natural experiments, such as the study of
zooplankton populations trapped in Gulf Stream
rings, which are eddies detached from the Stream
and moving through the alien environment of the
Sargasso Sea (Wiebe, 1976). These eddies with
scales of about 100 kilometers and lifetimes of 100
days have the correct dimensions for study of
zooplankton populations. The changes in these
populations provide an index of the response to the
changed environment.

Our interest in these herbivorous
components of the food web is not only because of
their important role within that web but also
because we can think of harvesting them as a source
of human or animal foodstuff. There have been
some minor attempts at harvesting in the Northern
Hemisphere, in Norwegian and Japanese waters,
and mainly as bait for sport or commercial
fishermen. Butthe best known possibility is the
Antarctic krill where theavailable resource could be
comparable to our present total world yield of fish.
The potential for initial harvesting involves
numerous economic and technical factors but
depends ultimately on the ability to locate and
capture the dense swarms. How do the swarms
aggregate in relation to physical features? Can one
define the behavioral components of the
concentrating mechanisms ? Continued
exploitation will be related to the response of the
population to intensive fishing. Will the swarms
decrease in size and concentration? Or will they be
replenished from peripheral and less dense
communities? Will the controlling mechanisms that
appear to operate with whales also apply to large,
predatory, krill boats? These questions display the
links between the basic ecology and the problems
of management; and the dependence of both on
our understanding of the interactions between
large-scale productivity and small-scale pattern. The
former controls the overall possible yield, the latter
determines how we as well as the natural predators
can take this yield economically and safely.

John H. Steele is Director of the Woods Hole
Oceanographic Institution.

Literature Cited:

Bell, F. W. 1978. Food from the sea: The economics and politics of

ocean fisheries. Colorado: Westview Press.
Gushing, D. H. 1955. On the autumn-spawned herring races of the

North Sea. /. Cons. Int. Explor. Mer. 21 : 44-60.
Gushing, D. H., and D. S. Tungate. 1963. Studies on a Calanus

patch. I. The identification of a Calanus patch./. Mar. Biol.

Assn. U.K. 43:327-337.
Dooley, H. D. 1974. Hypotheses concerning the circulation of the

northern North Sea./. Cons. Int. Explor. Mer. 36(1): 54-61.
Laws, R. M. 1977. Seals and whales in the Southern Ocean. Phil.

Trans. Roy. Soc. Lond. B. 279: 81-%.
Mackas, D. L., and C. M. Boyd. 1979. Spectral analysis of

zooplankton spatial heterogeneity. Science 204: 62-64.
Oceanus. 1979. Vol.22, No. 1,72pp.
Omori, M. 1978. Zooplankton fisheries of the world: a review. Mar.

Biol. 48: 72-76.
Ryther, J. H. 1969. Photosynthesis and fish production in the sea.

Sc/ence166: 72-76.
Sheldon, R. W.,A. Prakash, and W. H. Sutcliffe, Jr. 1972. The size

distribution of particles in the ocean. Limnol. Oceanogr. 17:

327-340.
Steele, ). H. 1961. The environment of a herring fishery. Mar. Res.

Scot. 6: 19pp.
Steele,). H. 1974. The structure of marine ecosystems. Cambridge,

MA: Harvard University Press.
Wiebe, P. 1976. The biology of cold-core rings. Oceanus 19(3):

69-76.

8

•

.V

Lessons for Coastal

L , -

Fisheries in Africa?

by Michael H. Glantz

Ihere is a mystique surrounding scientific research
that attempts to forecast future states of the
environment, probably stemming from the belief
that more information can only be beneficial. The
mystique is often generated by no more than
wishful thinking about the value of such research;
but in some instances that value has not been
determined. A good example is the attempt to
develop an environmental forecast of the
meteorological-oceanographic phenomenon
known as El Nino, which has an adverse effect on
the biological productivity in Peruvian coastal
waters, ultimately affecting Peru's anchoveta fishery
(see Oceanus, Vol. 21, No. 4, p. 40). It is widely
believed that such a reliable forecast would supply
the scientific information needed to manage the
fishery rationally. One might argue, on the other

/Above, part of Peru's anchoveta fleet in harbor.

hand, that considerable scientific information
about the fishery already exists, and that what is
needed is a better use of these data rather than large
quantities of new information.

In addition, the call for more scientific
research often tends to shift attention toward
geophysical processes and away from social,
political, and economic questions. However, it may
be within the social processes that solutions to
many questions about environmentally rational
fisheries management lie. Assessing the societal
value of such a forecast as an exercise in identifying
the role of scientific information in the
development of a healthy fisheries sector is of
interest not only to Peru but also to other
developing countries that border coastal upwelling
regions, such as Mauritania and Somalia, which are
in the process of expanding their commercial
fisheries operations.

f/gure 7. Ma/or coastal upwelling regions of the world and the sea-level atmospheric pressure systems (anti-cyclones)
that influence them.

Peru and El Nino

In terms of biological productivity, the Peruvian
coastal zone is one of the five major coastal
upwelling areas in the world (Figure 1). El Nino
(in Spanish, referringto the Christ child because it
usually occurs at Christmastime) is the name given
to a sporadic invasion of warm, nutrient-poor
surface water into the eastern equatorial Pacific that
tends to overshadowthe naturally occurring coastal
upwelling processes in which deep, nutrient-rich
cold water is brought upward into the sunlit or
euphotic zone. The sudden depletion of nutrients
in the upper sunlit layers of the ocean adversely
affects phytoplankton and zooplankton
production, thereby disrupting the food chain. The
result (of which El Nino is probably only a link in a
series of events*) is that fish populations are also
adversely affected. Fish apparently become widely
dispersed in their search for food, recruitment is
reduced, and a high density of anchoveta appears

"Teleconnections is a word used by meteorologists to
describe apparent correlations between meteorologically

related phenomena in different parts of the world in this

instance between El Nino, cold winters in the United
States and Europe, weakening of the Indian monsoon,
heavy rains and unusual hurricane activity in the Pacific,
and droughts in the Sahel and in northeast Brazil.

near the coast where some pockets of cold,
nutrient-rich deep water upwell to the surface.

Before 1970, Peru's fishing industry
underwent a rapid expansion, with catches
sometimes doublingannually (Figure 2). By the
mid-1960s Peru, registering about 22 percent of the
world's commercial catch by tonnage, was
considered to be one of the world's leading fishing
nations. At that time, physical and economic
conditions appeared favorable for continued
expansion of the anchoveta fishery.

Butoptimism waned in 1972-73 when a major
El Nino event occurred. At its onset, the effect was
hardly noticed. Large numbers of anchoveta were
found close to shore, in high-density pockets where
upwelling to the surface still occurred despite the
general invasion of warm, nutrient-poor surface
water into the area. Record catches were taken in so
short a time that the size and age distribution of the
fish could not be assessed, variables that might have
given insight into the condition of the standing
stock. Later assessments showed that excessive
fishing had already cut heavily into the standing
stock of anchoveta during 1971, when the year-class
recruitment had been a poor one. Thus, at the very
time when (according to many observers) there
should have been less fishing, the opposite
occurred.

10

After the sharp reduction in biological
productivity and anchoveta landings in 1972-73, the
Peruvian government nationalized the fishing
industry, ostensibly to preserve the anchoveta
resource, but also to fulfill economic objectives. In
1976-77, a policy of denationalization was pursued
because the anchoveta had not returned in large
numbers and the government could no longer
subsidize fishermen or workers in fish processing
plants.

It generally has been assumed that with a
reliable El Nihoforecast, thedepletion of anchoveta
stocks could have been avoided, which in turn
would have mitigated, if not averted, the economic,
political, and social dislocations that resulted from
the reduced productivity of the fishery. Hence, the
argument goes, an El Nino forecast ought to be of
great value to decisionmakers in Peru and
elsewhere.* There has been a great interest within
the scientific community to develop such a
capability.

El Nino Forecast Value

Experts knowledgeable about El Nino, Peru, and the
fishmeal industry — in government, fisheries
management, the physical sciences, economics,
and public policy within and outside Peru — were
asked inan open-ended mail survey bytheauthorin
1977 what they thought might be the value of an El
Nino forecast two to four months in advance of the
event. They were asked to suggest what might have
been done in 1972 had there been a reliable
forecast. Responses included such opposing views
as tying up the fishing fleet until a year or so after the
El Nino had passed, thus protectingthe standing
stock of anchoveta; and allowing all-out fishing,
either to thin the stock so the remainder could have
a better chance of survival, or because many of
those caught would die anyway. Both views were
based on the belief that the government wanted to
manage the fishery so as to preserve the resource
for exploitation in future generations; but the lack
of consensus in the scientific community about the
connection between an El Nino and biological
productivity led the respondents to their opposing
views.

Other suggestions included closing the
fishery and subsidizing various sectors of the fish ing
industry, increasing fishmeal storage capacity and
holding stocks in reserve to be sold at elevated
prices during future El Nino events, sharply
reducing the fleet and number of fishmeal plants,
appointing a sole authority to manage the fishery,
and shifting from the northern to the southern zone
on the Peruvian-Chilean border.

*The anchovy is reduced to a higher-value export product,
fishmeal, which in the early 1970s was exported to markets
in the United States and Western Europe as a feed
supplement for poultry, swine, and cattle.

i::

c 10
I 9
1 8

LU 6

O 5

4
3
2

I

CJ
>•

I I I I I I I I I I I I II I I I I I I I I I

5 1955 57 59 61 63 65 67 69 71 73 75 77

YEAR

* Estimated

Figure 2. Before 1970, Peru's fishing industry underwent a
rapid (some say meteoric) expansion with catches in its
early years doubling annually.

Survey respondents also identified some of
the constraints that might diminish the value of an El
Nino forecast, such as the relatively short lead time
(two to six months); forecast unreliability; the
constantly changing characteristics of El Nino -
intensity, duration, and magnitude; and, perhaps
most important, political and economic constraints.
Scientific information is only one aspect that will
figure into decisions made by a fisheries
policymaker; it will be integrated with economic
and political considerations, which often tend to
outweigh the rational scientific arguments for
cautious exploitation of the anchoveta fishery.

Even if there were specific data on the
relationship between an El Nino forecast and
biological productivity, such information would not
necessarily be properly used. There have been
reports on the Peruvian fisheries issued before 1972
that could be considered forecasts; however, the
implicit or explicit warnings in those reports were
not incorporated into the decisionmaking
processes in any significant way. For example, one
report written by R. C. Murphy in 1954 for the
Peruvian Guano Administration noted that unless
management was oriented toward conservation of
the anchoveta fishery, not only would the fishery
collapse, but the guano industry as well.*

In 1969 the Peruvian Marine Institute
(IMARPE) issued a report suggesting the likelihood
that the fishery would collapse from economic
pressures from overcapitalization of the fleet and
the fishmeal processing plants. In 1971, G. J. Paulik
described a scenario that combined economic

*Guano is the droppings from birds such as pelicans,
cormorants, and gannets. These birds feed almost
exclusively on the anchoveta and would likely perish
without them. Guano is rich in nitrogenous and
phosphatic matter and is used as fertilizer.

11

Pelicans, cormorants, andgannets on island off the coast of Peru. The droppings (guano) of the birds are rich in nitrate
for fertilizer. Their sole food is the anchoveta.

pressures similar to those discussed in the 1969
report with the occurrence of a major El Nino,
suggesting the inevitable decline of the anchoveta
fishery. His description approximated what actually
transpired in 1972-73.*

Thus the Peruvian anchoveta situation
suggests that although in theory there might be
many positive uses for a reliable El Nino forecast, in
practice its value may be more limited. A further
limitation is imposed by disagreements within the
scientific community. The survey suggested that
even with a reliable forecast, there would still be no
consensus among scientists about how it might be
linked to biological productivity, and in turn, used
as an effective management tool. Faced with
contradictory views, policymakers might well weigh
all opinions equally, in effect cancelling out the
input of the scientific community to the
decisionmaking process. Despite a constant call for
more research to reduce environmental
uncertainties, there appears to be little likelihood
that such research would lead to unanimity among
scientists.

*After the 1973 low catch of 1 .8 million metric tons
(MMT), the fishery recovered somewhat to a level of 4
MMT, only to col lapse again to less than 1 MMT in 1977. In
1979, catches included about 1 MMT each of anchoveta
and sardine, the latter increasing in number with the
decline in anchoveta population.

Major African Coastal Upwelling Regions

Of the five major upwelling regions, three have
witnessed the collapse or near collapse of a major
commercially exploited pelagic fishery: the
Peruvian anchoveta, the California sardine, and the
South West African pilchard. With respect to the
California sardine industry, the controversy
continues as to whether fluctuating environmental
conditions or heavy fishing were primarily
responsible for its collapse. The literature on the
growth and decline of this fishery is substantial
and still expanding. The South West African
(Namibian*) pilchard decline is also under scrutiny.
It has been suggested recently that scientific
information was available that might have
prevented the collapse, but that industry and
government officials downplayed this scientific
input in favor of economic considerations. The
South West African fisheries is a multispecies one,
making analysis of the situation more complex. Yet
debates similar to those concerning the demise of
the California sardine and the Peruvian anchoveta
are beginning to surface.

What is the situation in the other two major
upwelling areas? Both are in African waters, one off

* Namibia is the name given to South West Africa by those
(including the United Nations) seeking to gain its
independence from the Republic of South Africa.

12

the coast of the Islamic Republic of Mauritania, and
the other off the coast of the Somali Democratic
Republic. Are there lessons that might be learned
from the experiences of the Peruvian, Californian,
and South West African (Namibian) fisheries? To
what level and at what rate should exploitation
occur, given the current environmental
uncertainties?

Africans expressed concern in 1973 and
earlier that there was a high probability that
non-African long-distance fishing vessels would
overfish local waters before the coastal African
countries were technically and economically ready
to join in the fisheries. For example, Bayagabona, a
Nigerian oceanographer, noted in 1973 that there
were already indications "that most of the marine
fish stocks in Western Africa [were] being fished at
maximum sustainable yield and that some [were] in
fact already being overfished." Potential conflicts
also exist among African nations themselves
because the states with rich coastal resources are
among the least populated, such as Mauritania,
Namibia, and Somalia. The nations with larger
populations and demands for fish products such as
the Ivory Coast, Ghana, and Nigeria, are relatively
more affluent and developed, but are poorer with
respect to commercially exploitable marine
resources in their coastal zones.

The establishment of the 200-mile exclusive
economic zone combined with the need for
economic development has focused attention on
the exploitation of coastal marine resources. The
fishing industries in the three African countries that
border major upwellingareas (Somalia, Mauritania,
Namibia) are at different stages of development.
Somalia presently has the least developed
commercial fisheries, Namibia the most developed,
and Mauritania is in between, but considerably
closer to the Somali situation. Although in the past
Mauritania and Somalia have been viewed as among
the poorest countries in Africa with respect to
natural land resources, their coastal waters are
highly productive and must, therefore, be included
in any national inventory of natural resources.

Mauritania

Mauritania is situated in the Sahara desert
with its southern territory extending into the
Sahelian zone, and has a population of 1.4 million.
The drought in the West African Sahel (1968-73)
adversely affected the country's economic
development plans by increasing livestock
mortality, intensifying rural to urban migration, and
accelerating the deterioration of the land's
productivity.

Mauritania is often characterized as a poor,
coastal desert country with few exploitable
resources. Those that it has — iron ore and copper,
for example — are subject to fluctuations in the
international marketplace. Exploitation of its coastal

waters, considered by many researchers to be
among the most biologically productive in the
world, is just now emerging as a high priority in
Mauritanian economic development plans. Until
recently, vessels from more than twenty countries
fished this upwelling region, with little economic
benefit gained by Mauritania. For example,
although the government derived some benefit
through licensing arrangements with foreign
vessels, in several instances the fees were not paid
by vessels that chose instead to "pirate" their
catches or to exploit adjacent waters where fees
were lower.

Mauritania's interest in its coastal fisheries
was heightened by a recent French report that
assessed the value of the annual catch at about $2
billion, notingthat only about4 percent goes to
Mauritania. These conditions may be changing,
however, as Mauritanian officials turn their
attention toward financing and developing their
fisheries. Authorities have requested funds from
the international community (both private and
government) for trawlers, training, fish processing
factories, and storage facilities, and for the
expansion of port facilities at Nouadhibou. Officials
also have increased licensing fees, for example, for
foreign factory ships, and are campaigning for
foreign vessels to process their catches in
Mauritanian factories.

This is a very exciting but crucial time for
those involved in the planning and development of
fisheries-related projects in Mauritania. Foreign
companies recently have begun to join Mauritania
in developing its national fishing industry, and
Mauritania is givingthese firms incentives, such as
the right to expatriate profits, and tax breaks. It is
precisely now, as the demands for heavy capital -
for fleets and factories — are being voiced, that
examples of the growth and development of fishing
industries in other developing countries that border
strong coastal upwelling regions need to be
examined for lessons on what might and might not
be done.

Somalia

Somalia is strategically located on the Horn of
Africa, and has one of the highest percentages of
nomadic populations in Africa. It, too, is a coastal
desert country with biologically productive,
unexploited, coastal upwelling. Its population
depends primarily on resources derived from the
land, such as agriculture and livestock.

In the early 1970s and culminating in 1974-75,
Somalia was plagued by drought; about 20,000
people and more than a million animals died.
Hundreds of thousands of Somalis migrated to
refugee camps. The impact of this natural disaster
prompted the government to take advantage of
these social dislocations to pursue its plans to
convert some of these pastoralists to farmers. In

13

The
Change

from

The town of Baraawe, where the largest fishing cooperative is located

Inset: Women mending nets at Baraawe.

New houses for resettled nomads in Badey with traditional nomadic huts in background.

14

Nomad to Fisherman in Somalia

~» --

Truck belonging to Badey cooperative on
three-day journey to Mogadishu to deliver load
of dried fish.

Fish being dried at processing plant in Baraawe.

Boats used for fishing at Baraawe cooperative.

All Somalia photos by Jan M. Haakonsen

15

Lack of port facilities requires fishermen to anchor their boats away from shore.

addition, about 15,000 of the 120,000 or so destitute
refugees were resettled in four fishing villages along
the southern coast, three of which were established
specifically for the project. This project was the first
phase of a long-term economic and social
development program designed to exploit the
coastal marine resources of Somalia for local
consumption and for export to neighboring
countries. These artisanal fishing efforts would
complement the developing commercial fishing
industry based on the building up of a fleet of
trawlers. As noted recently by Haakonsen, "The
change from nomadic pastoralism to sedentary
fishing is a major one and, on such a large scale,
actually unprecedented in history." In fact, it can be
seen as a major social experiment. A Somali official
noted that:

Until 1972, when this Ministry [of Fisheries] was
formed, Somalia really had no fishing industry,
despite having the longest coastline of any
independent African country. So the very fact that
we've got these nomads to go down to the sea to
fish at all is an enormous achievement. I'm sure our
investment is going to pay off. Then, twenty years
from now fisheries are going to be making a very
significant contribution to Somalia's economy.

The government-sponsored

multidisciplinary team that studied the feasibility of
fishing village projects concluded that:

The problem with the fisheries development in
Somalia is that nobody knows the extent of the

Weighing the catch at Illig, near Badey.

16

resources available for exploitation. ... /As part of
the renewed interest by the Somali government for
sea resource utilization, we proposed that a
thorough study of the seas be undertaken as a
prelude to any developmental ventures in this field.

It these fishing villages are successful, it will
in all probability mean increased exploitation of
their coastal living marine resources as demands
increase in domestic and export markets. As
fisheries management experts have noted for other
fisheries, a major factor that adversely affects
fisheries productivity in the long run is excess
capacity of fleets and factories. As John Gulland, a
fisheries expert, often maintained, it is wiser (and
easier) to prevent the excess capacity from being
developed until more is known about the fisheries
than it is to remove such excess capacity after it has
been determined that it has impaired the biological
productivity of the fishery. This is clearly one of the
messages that Somali and Mauritanian planners can
see from an assessment of the Peruvian situation.

A Look Ahead

The situation in Peru should be of interest not only
to Peruvians, but also to policymakers in countries
seeking to attain higher levels of development
through the exploitation of their own coastal marine
resources, such as Somalia and Mauritania. Viewed
in light of the California sardine and the South West
African pilchard declines, Peru's situation suggests
that over-capitalization of a fishing industry must be
avoided; that lower levels of exploitation must be
made acceptable until environmental uncertainties
can be reduced; that fishing for export should not
be considered a country's primary means of
development; and that more scientific information,
even of better quality, might not ensure that the
right decisions will be made.

It is apparent that Peru's anchoveta needed
an advocate to counter the challenges of those who
favored all-out commercial exploitation. Given the
environmental and biological uncertainties in
coastal upwelling areas, high levels of exploitation
must be avoided to protect the marine resources in
such regions until those uncertainties are reduced.
From the Peruvian example, it is evident that the
costs associated with social, economic, and political
dislocations caused by the collapse of a fishery
could prove to be greater than the temporary
benefits derived from overexploitation based
primarily on short-term economic interests.

Michael H. Clantz is a political scientist with the
Environmental and Societal Impacts Croup at the National
Center for Atmospheric Research in Boulder, Colorado.

Suggested Readings

Cram, D. 1980. Hidden elements in the development and

implementation of marine resource conservation policy: The
case of the South West African/Namibian fisheries. In Resource
management and environmental uncertainty: lessons from
coastal upwelling fisheries, ed. M. H. Clantz and J. D.
Thompson. New York: Wiley Interscience.

Crutchfield, J.A., and R. Lawson. 1974. West African marine
fisheries — alternatives for management. Washington:
Resources for the Future. 64 pp.

Glantz, M. H. 1979. Science, politics, and economics of the
Peruvian anchoveta fishery. Marine Policy 3(3): 201-10.

Gulland, ]. A. 1978. Fishery management: New strategies for new
conditions. Trans. Am. Fish. Soc. 107(1): 1-11.
— . 1980. Peruvian anchoveta — optimal management. Marine
Policy 4C]): 78-79.

Haakonsen, ). 1979. "Survey on the fishing cooperatives for
resettled nomads in Somalia." Mimeograph. Center for
Developing Area Studies, McGill University, Montreal.

Murphy, R. C. 1954. "Guano and the anchoveta fishery." Official
report to the Peruvian government, 1954. Reprinted in
Resource management and environmental uncertainty: lessons
from coastal upwelling fisheries, ed. M. H. Glantz and ). D.
Thompson. New York: Wiley Interscience, 1980.

Nancock, G. 1977. Nomads settle for the seaside. Geographical
Mag., August 1977, pp. 679-80, 684.

O'Brien, J. I. 1978. El Nino: An example of ocean/atmosphere
interactions. Oceanus 21(4): 40-46.

Paulik, G. J. 1971. Anchovies, birds and fishermen in the Peru
current. In Environment, resources, pollution and society, ed.
W. W. Murdock, Sinnauer Press. Reprinted in Resource
management and environmental uncertainty: lessons from
coastal upwelling fisheries, ed. M. H. Glantz and j. D.
Thompson. New York: Wiley Interscience, 1980.

17

:

by Michael J. Mottl

are rocks and minerals that can be
recovered at a profit," according to a popular
textbook on economic geology — Park and
MacDiarmid'sOreDepos/Ys (1964). Using this
economic criterion, few of the hydrothermal
deposits discovered on the seaf loor in recent years
could be classified as ores because they lie beneath
several thousand meters of seawater, which makes
their recovery prohibitively expensive.
Nevertheless, some deposits are large and rich

H

.

Figure 7. A row of chimneys on the East Pacific Rise at 21
degrees North with multiple vents spouting hot water.
Black or white "smoke" forms on mixing. Black smokers '
are hottest, up to 350 degrees Celsius. Inset: Iron pyrite,
silver sulfide, and other minerals glitter in rock retrieved at
27 degrees North. (Photo by Emory Kristof. ©National
Geographic Society)

enough to be exploitable, and those that are not
promise to be profitable in another way by
increasing our understanding of how hydrothermal
ores form so that we can locate them more easily in
the future.

The study of submarine hydrothermal
deposits was considerably advanced in 1979 with
the discovery of hot springs on the East Pacific Rise
at 21 degrees North. Unlike the warm springs
discovered a few years ago on the Galapagos
Spreading Center (see Oceanus, Vol. 20, No. 3, p.
35), these springs are hot! They are venting water at
temperatures as high as 350 degrees Celsius at
velocities of several meters per second, and are
precipitating prodigious quantities of sulfide ore,
minerals rich in copper, zinc, and iron. The
precipitates form chimneys around the individual
vents that spout black or white smoke composed of
precipitated crystals of sulfides and other minerals
(Figure 1). The discovery is easily the most exciting
and significant in this field si nee the discovery of the
Red Sea hot brines and metal deposits (see
Oceanus, Vol. 22, No. 3, p. 33).

Hydrothermal Ore Deposits

Hydrothermal refers to hot water: hydrothermal
deposits are deposits that have formed by chemical
precipitation from hot solutions, the principal
constituent of which is water. There are many
different types of hydrothermal ore deposits,
formed in a wide variety of environments and under
a wide range of conditions. Together they represent
one of the most economically important classes of
ore deposits, yielding much of the world's supply of
copper, zinc, lead, silver, gold, tin, molybdenum,
and other metals. All hydrothermal ore deposits
have five components that contribute to their
formation: 1) a source for the ore metals; 2) a source
for the water that dissolves the metals and later
precipitates them, concentrating them in the
process; 3) a source of heat; 4) a system of
permeable pathways through which the water flows
from source to the site of deposition; and 5) the site
of deposition. In addition, the ores must be
deposited in a setting that allows them to survive
(for perhaps millions of years) the ravages of
weathering, erosion, and other processes that
would tend to disperse the metals. Variations in
these five components, along with differences in
the physical and chemical conditions experienced
by the hot solution as it travels from its source to the
site of deposition, account for the great variety of
hydrothermal ore deposits exploited today.

The submarine environment provides a
unique combination of components for the
formation of hydrothermal deposits. By far the most
abundant source of heat in this setting is that
associated with the formation of new oceanic
lithosphere along the mid-ocean ridge system,
where the seafloor is spreading apart and basaltic

magma wells up from deep within the earth to form
new ocean crust (see Oceanus, Vol. 17, No. 3). The
new crust becomes highly fractured as a result of
the tensional forces that pull the lithospheric plates
apart. Seawater percolates down through the
fractures, becomes heated through contact with the
hot rock, and begins to react with it, leachingmetals
from the rock. The heated seawater, now
chemically modified and carrying dissolved metals,
is less dense because it is hot, and so tends to rise. It
ascends to the seafloor, exiting as submarine hot
springs whose waters mix with the ocean bottom
water. Thus, for submarine hydrothermal deposits,
the ore-forming solution is usually modified
seawater, the source of metals is the rock of the
ocean crust, and the site of deposition is on or
within the seafloor. This combination of
circumstances has and is producing a variety of
hydrothermal deposits on the seafloor, as well as a
number of economically important analogues that
have found their way onto land, where they are
being mined.

Seawater as an Ore-forming Fluid

The concentrations of ore metals in most natural
waters are extremely low, and in normal seawater
they are especially so. In fact, concentrations in
seawater are so low that simply measuring them
accurately has been one of the most difficult
problems in marine chemistry during the last 30
years. For example, if one makes artificial seawater
in the laboratory from the high-purity "reagent
grade" chemicals used by chemists, it will have 100
to 1 ,000 times the concentration of heavy metals
found in natural seawater!

In order for seawater or any other natural
water to become an ore-forming fluid, some
minimum concentration of ore metals in solution is
required. This is because a metal deposit must be
fairly large before it becomes profitable to mine.
The minimum size for a deposit to be considered
ore is an economic question. The total tonnage of
ore in a given deposit depends on how long it took
the deposit to form and how fast the hydrothermal
solutions were able to transport metals to the site
and deposit them there. The latter depends, in turn,
on the flow rate of the hydrothermal solution, its
total concentration of dissolved metals, and the
efficiency with which the metals were precipitated
as the solution passed through the site of
deposition.

For seawater to become an ore-forming fluid,
its ore metal concentrations must be increased by a
factor of about 20,000 or more. As stated earlier, the
most abundant source for these metals is ocean
crust. The problem is to get the metals out of the
rock and into the seawater, and to keep them there
at high enough concentrations until they can be
precipitated in a restricted locality. Laboratory
experiments on the solubilities of various ore

20

minerals and metals show us that there are three
principal ways to increase the concentration of
metals in an aqueous (water-rich) solution in
contact with rock — increase its temperature (and
pressure), increase its salinity, or decrease its pH,
that is, make it more acidic. The first two of these
methods are known to be important for submarine
hydrothermal deposits; the third is an intriguing
hypothesis awaiting confirmation in a natural
setting.

The temperatures reached by seawater
circulating through newly formed oceanic crust at a
mid-ocean ridge may be estimated in several ways.
Rocks dredged from fault scarps in these submarine
mountain chains indicate from their chemistry,
mineralogy, and oxygen isotopic composition that
they have been altered by seawater at temperatures
ranging from a few degrees to 600 degrees Celsius,
with the bulk of the high-temperature alteration
occurring at 200 to 300 degrees Celsius (see
Oceanus, Vol. 19, No. 4, p. 40).

The first submarine hot springs discovered
along a mid-ocean ridge, those at the Galapagos
Spreading Center, were emitting water at only 20
degrees Celsius, but the chemistry of this water
indicated that it had reacted with basalt at 350 to 400
degrees Celsius (Corliss and others, 1978; Edmond
and others, 1979). Then came the discovery of the 350
degrees Celsius springs on the East Pacific Rise; the
chemistry of this water is presently being analyzed
in John Edmond's laboratory at the Massachusetts
Institute of Technology (see Oceanus, Vol. 23, No.
1,p. 33).

Figures 2 and 3 show the concentration of
iron and manganese in seawater that reacted with
basalt for many months in laboratory experiments at
high temperature. Also shown are data from the
seawater-fed hot spring at Reykjanes, Iceland. Iron
and manganese, while less important as ore, are
much more abundant in both rocks and solution
than are copper, zinc, and other ore metals. All of
these metals can be expected to show the large
increase in concentration with temperature (and
pressure) shown inthefigures. Notethatthe largest
increase occurs above 350 degrees Celsius. For
altered seawater solutions such as these that have
normal salinity and are neither strongly acid nor
strongly alkaline, 350 degrees Celsius represents
{he minimum temperature at which seawater is
likely to become an ore-formingfluid.

Compared to the solutions responsible for
most known hydrothermal ore deposits, seawater is
quite dilute, having only 3.5 percent total dissolved
salts. Typical ore solutions, as deduced from studies
of minute amounts of solution trapped in ore
minerals, range in salinity from 3 to 50 percent. The
second way to turn seawater into an efficient
metal-transporting agent is to increase its salinity.
This mechanism is largely responsible for
producing the only true hydrothermal ore body

2500

1000
500
250

100
50

£ 25

) 10

5

2.5

1
0.5

0.2

t

Bischoff and Dickson (1975),

200°C
time

Reykjanes

A A

CO

O

200

300

T, °C

400

500

Figure 2. Concentration of iron in solutions produced by
reaction of seawater with basalt in laboratory experiments
and in the geothermal system on the Reykjanes Peninsula
of Iceland (from Mottl and others, 1979).

2500

— i — — i — — i — — ;

1000

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500

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250

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Bischoff and Dickson (1975),

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Reykianes L3

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0.5

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200

300

T, °C

400

500

Figure 3. Concentration of manganese in solutions
produced by reaction of seawater with basalt in laboratory
experiments and in the geothermal system at Reykjanes,
Iceland (from Mottl and others, 1979).

21

discovered so far in the oceans: the deposits in the
Atlantis II Deep and in other nearby basins along the
spreadingcenterin the Red Sea. As elsewhere along
the mid-ocean ridge system, the source of the
hydrothermal solutions is seawater, the heat is
derived from the seafloor spreading process, and
the metals are leached from the crustal rocks. What
makes the situation in the Red Sea unique is that the
hot, circulating seawater encounters not only
volcanic rock during its subterranean travels, but
also salt — huge deposits of it, several kilometers
thick. These deposits of predominantly halite
(NaCI, common table salt) formed when the Red Sea
was younger and narrower than it is today. The salt
deposits formed by evaporation of seawater in the
long, narrow new ocean basin when its outlets to
the rest of the oceans were constricted or closed.

Halite is extremely soluble, especially in hot
solutions, so these deposits are readily dissolved
into the circulating seawater, increasing its salinity
to about nine times that of normal seawater. The
resulting hot brine has a much greater capacity for
transportingore metals in solution because most of
these metals readily combine with the chloride ion,
Cl~, in solution to form highly soluble neutral or
charged complex ligands, such as FeCh, CuCI°, and
ZnCI°2; these com pi exes form in greater abundance
at higher temperatures and chloride
concentrations. Thus, the Red Sea hot brines
contain high concentrations of manganese, iron,
lead, zinc, copper, cobalt, silver, and other metals.

The third method for dissolving high
concentrations of metals in seawater is by lowering
its pH, which is the same as raising the hydrogen ion
(H* ) activity in solution. As anyone knows who has
spilled battery acid on his car, metals tend to
dissolve readily in acid (hT-rich) solutions.
Experiments at Stanford University four years ago
led to a startling discovery: ordinary seawater,
when heated to high temperature (greater than 250
degrees Celsius), has a built-in mechanism for
becoming a highly acid hydrothermal solution
(Bischoff and Seyfried, 1978). As temperature
increases, seawater becomes saturated with and
precipitates a previously unknown and as yet
unnamed mineral containing magnesium, sulfate,
and hydroxyl (OH ) ions. Magnesium and sulfate are
two ubiquitous constituents of sea salt, whereas the
hydroxyl ion comes mainly from the water itself:
H2O=H+ + OH . This reaction coupled with
removal of OH from solution into the new mineral
produces free H^ , making the hot seawater very
acid (pH 3.5 at 350 degrees Celsius). Such an acid
solution is very corrosive to the surrounding rocks
and readily attacks them, leaching metals into
solution. This reaction with the rocks tends to
consume the H* and also to convert the
Mg-OH-SO4 mineral into common Mg-OH-silicate
minerals, such as clays. Whether the hot seawater
remains acid depends on the relative rates of the

H+-producing and H*-consuming reactions. These
depend, in turn, on how fast seawater flows
through the submarine hydrothermal system,
compared with how fast it reacts with the rocks it
flows through. If flow rates are more rapid, then
extremely metal-rich hot spring solutions can be
produced (Figure4).

Although such highly acidic hot springs have
yet to be discovered, plans are being formulated to
explore the area where they are most likely to
occur: along the East Pacific Rise south of the
equator. This area of the seafloor contains the most
extensive hydrothermal deposits known to exist in
the oceans. The deposits are predominantly iron
and manganese oxides that have accumulated four
to thirty times faster than the normal rate for the
deep oceans. Mixed in with normal deep-sea
sediment, they are a few meters to a few tens of
meters thick and extend from about 10 to 30 degrees
south, covering an area of 625,000 square
kilometers.

The Seafloor as a Depositional Environment

An ore deposit is by its very natu re a concentration
of metals. Even most ore-forming solutions have
lower concentrations of metals than the rocks from
which the metals were leached. Thus, most of the
real concentrating gets done at the site of
deposition by processes that remove the metals
from solution. If an ore deposit is to be formed,
these processes must operate efficiently and over a
restricted area, removing metals from a very large
volume of solution as it passes through.
Otherwise, in the absence of a depositional
environment with suitable characteristics, the
solution will simply distribute the metals over a
wide area once again.

The common feature of depositional
environments for hydrothermal ore deposits is their
ability to produce a rapid and drastic change in
those properties of the solution which keep the ore
metals dissolved in the first place — temperature,
pressure, salinity, and pH. Another important
property is the degree to which the solution is
oxidized. Rapid and drastic changes in these
properties can cause the metals to be "dumped"
from solution, whereas more gradual changes
cause them to be precipitated slowly over a larger
area. Thus, slowcoolingof a hydrothermal solution
as it ascends toward the surface, by loss of heat to
the surrounding rocks, is less likely to produce an
ore deposit than is the rapid cooling that would
result from a sudden pressure drop, orfrom mixing
with another, cold solution. As for a sudden change
in pH, this is most likely to occur when the
hydrothermal solution encounters either another,
more alkaline solution, or a different type of rock,
such as a calcareous marine sediment, and reacts
with it. A third mechanism for raising pH is boiling,

22

300°C

v/r = 10

/r = 61

160

120

Fe2+

8

ppm

40

pH

50
DAYS

100

50

100

150

DAYS

Figure 4. Concentrations of magnesium and iron and pH with time in solutions from five experiments reacting
sea wafer with basalt. The two experiments on the I eft used low water /rock ratios and simulate flow that is slow relative
to reaction. The three on the right, at high water/rock ratios, simulate flow that is fast relative to reaction (from Mottl
andSeyfried, 1980).

accompanied by loss of acid-forming gases such as
carbon dioxide.

A depositional environment must have two
other properties if it is to produce an ore deposit-
it must be stable and remain in one place long
enough for a sizable amount of ore to accumulate
there, and the ore must be preserved there after
deposition until it can be discovered and exploited.

Most submarine hydrothermal activity is
likelytooccuralongthe mid-ocean ridge system. In
this setting, boiling is unlikely to occur because of
the high pressure from the 2 to 3 kilometers of
overlying seawater. Reaction of hydrothermal
solutions with marine sediments will be the
exception rather than the rule, as sediment is
normally very thin or absent altogether on such very
young crust. One process for precipitating metals
from hydrothermal solutions, however, is virtually
inevitable in any submarine setting: mixing of the
solution with ocean bottom waters. As
hydrothermal solutions are normally hot, slightly
acid, and reducing, whereas normal seawater is
cold, slightly alkaline, and oxygen- rich, mixing of
one with the other produces a drastic and nearly
instantaneous change in the chemical and physical
conditions in solution. The dramatic result can be
seen in Figure 1 , which shows the plume formed by
rapid, turbulent mixing of a 350 degrees Celsius
metal-rich hydrothermal solution with 2 degrees
Celsius bottom water on the East Pacific Rise. The
black smoke consists of iron, copper, and zinc
sulfide minerals that have precipitated on mixing.

Submarine Hydrothermal Deposits

To date, no submarine hydrothermal deposit has
been sufficiently studied that all components
contributing to its formation are known.
Nevertheless, the data at hand suggest some
intriguing relationships among known deposits
along mid-ocean ridges, pointing outthe
importance of special situations in producing and
preserving large deposits. A review of known
deposits illustrates this:

East Pacific Rise, 27 degrees North. The
spectacular discovery in 1979 of hot springs near the
mouth of the Gulf of California was presaged by the
finding the previous year of spire-like chimneys
composed of ore-grade zinc, copper, and iron
sulfide minerals in the same locality (CYAMEX,
1979). These deposits were discovered by the
French submersible Cyana duringthe joint
French-American-Mexican CYAMEX expedition,
and were associated with extinct hydrothermal
vents. When the RISE expedition returned in 1979
with the submersible/A/wn, operated by the Woods
Hole Oceanographic Institution (WHOI), and
foundtheactivevents, it was learned that Cyana had
missed them by at most a few hundred meters, a
fine example of the difficulties of search in the deep
sea!

All of the active vents discovered occuralong
a very narrowzone, 100to200 meters wide, close to
the axis of the spreading center. The RISE
expedition observed 25 temperature anomalies
associated with active vents over a 6-kilometer-long

23

Figure 5. Warm water vents on
the East Pacific Rise at 27 de-
grees North are similar to
those found along the
Galapagos Spreading Center.
Water at about 20 degrees
Celsius flows out from among
the rocks, supporting a di-
verse biological community,
including crabs and foot-long
clams.

segment of the ridge crest (RISE, 1980). The vents
range from the very high-temperature type (Figure
1) inthesouthwestto much cooler, Galapagos-style
vents (Figure 5) in the northeast. The cooler vents
are emitting water at 23 degrees Celsius or less and
are surrounded by diverse colonies of unusual
marine life, including giant clams and tubeworms,
crabs, limpets, and an eel-like fish, which depend
upon the chemically nurtured bacteria that grow in
the vents for their food source. The very
high-temperature vents, by contrast, are too hot for
living creatures, but even they are quickly colonized
in theircooler regions by worms, crabs, and eel-like
fish.

The most distinctive features of the
high-temperature vents are the chimneys. Most of
the chimneys are a few meters tall, but some are as
high as 10 meters. They are composed almost
entirely of sulfide minerals often arranged in
concentric zones (Figure 6), with minor amounts of
sulfate and silicate minerals and silica. The
chimneys are spaced a few meters apart within
elongated vent fields, and typically rise from a
mound composed of debris broken from the
chimneys themselves. These mounds rest directly
on basalt, as there is little or no sediment on such
young crust (less than 20,000 years old). The
chimneys are spouting either "black smoke,"
composed of sulfide minerals, or "white smoke"
composed of pyrite (iron sulfide), barite (barium
sulfate), and silica.

While these deposits are certainly ore-grade
(up to 50 percent zinc, 6 percent copper, 0.05
percent silver) and are forming in spectacular
fashion, they are not large or extensive. Forming as
they do on the open seafloor, they are immediately
subjectto oxidation and partial dissolution.
Exposed to oxygen-rich bottom waters, they are
unlikely to survive as sulfides beyond the
sediment-free limits of the axial valley. Thus, their
depositional environment is favorable to formation
of a small deposit, but not to its preservation.

Of greater interest as potential ore bodies are
the metal sulfides that may be deposited within the
seafloor in this setting. Observations made from
Alvin on a return trip in November, 1979, indicate
that vents with temperatures of 350 degrees Celsius
are depositing both copper and zinc, whereas vents
at 273 and 295 degrees Celsius are depositing only
zinc. For the latter vents, copper is probably
precipitating within the crust as the solutions cool
during ascent. Equally intriguing is the possibility
that the Galapagos-style vents in this locality are
much coolerthan the "smokers" becausethey have
mixed with cold seawater within the crust, at
shallow levels, rather than above it in the bottom
waters. This would shift the site of ore deposition to
a more protected location. The chemistry of the
Galapagos warm springs suggests that this is indeed
happening.

Galapagos Spreading Center. No
hydrothermal deposits have been recovered from

24

A

Figure 6. Two views of a chimney
from the East Pacific Rise at 21
degrees North. A: Side view of
15-centimeter high chimney, which
was spouting hot water at 350
degrees Celsius when sampled in
November, 7979. Chimney, at left,
protrudes from a base of massive
sulfides. B: View of same sample
looking upward through chimney.
Center of chimney is hollow and is
lined with chalcopyrite, CuFeS2.
Inner yellow zone is chalcopyrite
and py rite, FeS2. Outer black zone is
wurtzite, ZnS, and pyrite. White
patch near center of photo is
anhydrite, CaSO4, and talc, a
hydrated Mg-silicate. (Photos by
Marjorie Styrt)

the crest of the Galapagos Spreading Center, where
the first warm spring vents were discovered and
sampled in 1977. Features that appear to be extinct
sulfide chimneys have been photographed but not
sampled. Nonetheless, comparison of the
chemistry of the warm spring water with that from
laboratory experiments indicates that the warm
water results from shallow subsurface mixing of
cold seawater with a 350 to 400 degrees Celsius,
metal-rich hydrothermal solution. Thus, one can
imagine the precipitation processes observed on
the East Pacific Rise at 21 degrees North occurring
within the seafloor at the Galapagos, forming a
well-protected ore deposit.

Metal deposits, mostly iron silicates and
manganese oxides, do occur 25 kilometers south of
the Galapagos Spreading Center, where they form
mounds protruding from the sediment.
Precipitation of the metals here is mediated by
reaction of the hydrothermal solution with normal
marine sediments, as the solution emanates from
long fractures in the underlying basalt.

Mid-Atlantic Ridge. The best-known deposit
here is from the TAG hydrothermal area at 26
degrees North, discovered in 1970 by the
Trans-Atlantic Geotraverse project. This deposit is
pure manganese oxide. The TAG area may be similar
to the Galapagos in having extensive shallow
subsurface mixing. The chief unknown in this area is
the temperature, and therefore metal content, of
the hydrothermal end-member solution involvedin
mixing. If high, an ore deposit may be forming
within the crust here as well. Plans have been made
to visit this area with /A/wn in 1982.

East Pacific Rise, 10 to 30 degrees South.
These extensive, rapidly accumulating sediments,
rich in iron and manganese oxides, have already

•

B

25

been mentioned in discussing seawater as an
ore-forming fluid. The remarkable thing about
these deposits istheirgreatvolume, which impliesa
much larger input of hydrothermally-derived metals
than elsewhere along the mid-ocean ridge system.
Whereas the sulfide deposits at 21 degrees North on
the East Pacific Rise are highly localized, these
deposits blanket about half of the total area of new
ocean crust along this part of the ridge, which is
spreading at an abnormally rapid rate. Presumably
the high metal input is related to the fast spreading
rate, but exactly how or why is not known. Perhaps
the hot springs here, none of which have yet been
discovered, are hotter or more acidic than
elsewhere.

Another interesting question concerns the
mineralogy and chemistry of the deposits. Are
sulfides forming here, as at 21 degrees North, which
are then oxidized to oxides and hydroxides by
prolonged contact with ocean bottom waters? If so,
why are these deposits so relatively poor in zinc and
copper? Do rich deposits of zinc and copper lie on
and within the crust in this region? If so, they may be
analogous to the copper deposits on Cyprus. These
deposits are among the earliest known to have been
exploited — they were worked for copper ore as
early as 3000 B.C., and were mined extensively by the
Romans for centuries. They are still yielding
copper today. The deposits are located within a slab
of ocean crust that was thrust onto the land when
Europe collided with Africa millions of years ago.
Similar deposits occur in Newfoundland, Turkey,
and Oman.

Red Sea. As the only genuine hydrothermal
ore deposit found so far in the oceans, the Red Sea
heavy metal deposits deserve special mention. The
unusual metal-transporting properties of the hot
brines have already been discussed. What remains
is to describe the special depositional environment
created by these highly saline hydrothermal
solutions. These solutions exit from the seafloor at
about 100 degrees Celsius, along the floor of deep
fault-bounded basins. Although the solutions are
hot, they are denser than normal seawater because
of their high salinity, and so do not rise and mix
readily with bottom waters. Instead, they remain
within the deep basins, which are thus filled with
dense, metal-rich brines at a temperature of about
56 degrees Celsius. The metals precipitate within
the basins, producing sulfide, silicate, and oxide
minerals, depending on the local chemical
environment within the brines. Because of their
density, the brines are able to create a special and
unusual depositional environment for the ores.
Miningof theseores byaWestGerman firm that will
employ giant submarine "vacuum cleaners" is
about to begin.

Manganese Nodules. The other submarine
ores that promise to be mined eventually are
manganese nodules, potato-size nuggets of

manganese and iron oxides rich in nickel, copper,
cobalt, chromium, and other metals (see Oceanus,
Vol. 21 , No. 1 , p. 60). Manganese nodules are not
hydrothermal deposits. Rather, they accumulate
slowly by chemical precipitation from normal
seawater, in regions of the seafloor with slow
sedimentation rates. A long-standing controversy,
nonetheless, has been whether nodules contain
some amount of metal derived from a hydrothermal
source. Figure 7 shows that manganese derived
from the warm spring vents along the Galapagos
Spreading Center is easily detectable in otherwise
normal ocean bottom waters. This effect can be
seen at least 250 kilometers from the ridge (Bolger,
and others, 1973). This kind of evidence has revived
the controversy over the source of metals in the
nodules, in at least some areas where the nodules

occur.

Total dissolvable
manganese (pg per kg)

0.5 1.0

1,000

2,000

Q.
0)

Q

3,000

4,000

O
O

A

A

0° A

O

Figure 7. Concentration of manganese, including both
dissolved and particulate, in seawater from the open
Pacific (closed circles) and from the Galapagos Rift (open
circles and triangles) (from Klinkhammer and others, 1977).

26

Other Environments. While most
hydrothermal activity in the oceans occurs along the
mid-ocean ridge system, this is not the only
submarine environment where such processes take
place. Two particularly promising non-ridge
environments where ore deposits may be forming
under the sea are the diffuse spreading centers that
create marginal seas behind island arcs, such as the
Sea of Japan, and the bases of the volcanic island
arcs themselves. While hydrothermal processes
here will be similar to those at mid-ocean ridges in
most respects, two factors make them potentially
different. First, the molten magmas that form and
erupt in these environments have a much higher
content of water from the earth's interior dissolved
in them than do the characteristically dry magmas
(less than 1 percent H2O) at mid-ocean ridges. Some
of this water, which would be acid and metal-rich,
may be involved along with seawater in forming
metal deposits in these environments. These water-
and silica-rich magmas also are more likely to erupt
explosively than are mid-ocean ridge lavas,
fracturing the hot young rock and allowing ready
access for seawater. Second, the more silica-rich
rocks formed from these magmas are richer in some
metals, notably lead, than are basalts. Thus, ore
deposits formed here would be lead-zinc-copper
deposits. A noted example of submarine island-arc
ore deposits is the Kuroko ores of Japan. Similar
ores occur in the Philippine Islands, Fiji, Tasmania,
Canada, and possibly Turkey.

What the Future Holds

The discovery of the East Pacific Rise hot springs has
spurred tremendous interest among the
oceanographic community. Plans are now being
made to launch a four-year, multi-institutional
project to explore the East Pacific Rise for additional
areas of hot spring activity and ore deposition. The
major objective of the program will be to examine
the nature of hydrothermal processes along the
mid-ocean ridge system from the slow-spreadingto
the very fast-spreading segments, such as at 10 to 30
degrees South. The project will involve the use of
surface ships; deeply towed instrument packages,
such as Scripps Institution of Oceanography's Deep
Tow and WHOI's Angus sled; new high-precision
multibeam echo soundingfor making highly
accurate topographic maps of the seafloor; and
ultimately manned submersibles, including/\/wn. I
think it is fair to say that this rewarding field of
research is about to experience a veritable
explosion of information, which cannot help but
improve our understanding of how, where, and
why an important class of metallic ore deposits
forms.

Michael j. Mottl is an Assistant Scientist in the Department
of Chemistry at the Woods Hole Oceanographic
Institution.

Acknowledgment

The writing of this article was supported by a grant from
the National Science Foundation — OCE 78-19455 — to the
Woods Hole Oceanographic Institution.

References

Bischoff,). L.,and W. E. Seyfried. 1978. Hydrothermal chemistry of

seawater from 25° to 350°C. Amer. I. Sci. 278:838-860.
Bolger, C. W., P. R. Betzer, and V. V. Cordeev. 1978.

Hydrothermally derived manganese suspended over the

Galapagos Spreading Center. Deep-Sea Res. 25: 721-734.
Corliss, J. B., and others. 1979. Submarine thermal springs on the

Galapagos Rift. Science 203: 1073-1083.
CYAMEX. 1979. Massive deep-sea sulphide ore deposits

discovered on the East Pacific Rise. Nature 277: 523-528.
Edmond, ). M., and others. 1979. On the formation of metal-rich

deposits at ridge crests. Earth Planet. Sci. Lett. 46: 19-30.
Klinkhammer, G., M. Bender, and R. F. Weiss. 1977. Hydrothermal

manganese in the Galapagos Rift. Nature 269: 319-320.
Mottl, M. )., H. D. Holland, and R. F. Corr. 1979. Chemical

exchange during hydrothermal alteration of basalt by seawater

— II. Experimental results for Fe, Mn and sulfur species.

Ceochim. Cosmochim. Acta 43: 869-84.
Mottl, M. ]., and W. E. Seyfried. 1980. Sub-seafloor hydrothermal

systems: rock- vs. seawater-dominated. In Oceanic spreading

centers: Hydrothermal systems, ed. P. A. Rona, and R. P.

Lowell. Stroudsburg, PA: Dowden, Hutchinson, and Ross.
Park, C. F., Jr., and R. A. MacDiarmid. 1964. Ore Deposits. San

Francisco: W. H. Freeman and Company.
RISE. 1980. Hot springs and geophysical experiments on the East

Pacific Rise. Science 207: 1421-1433.

27

69°

Georges Bank

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DEVELOPMENT SCIENCES INC.
SAGAMORE. MASSACHUSETTS

67'

66'

Tracts leased as a result of Lease Sale No. 42, plotted against the spawning areas of major
commercial fisheries. Some scientists believe that spills or discharges from oil operations on the
tracts could affect the commercial fisheries, especially if the eggs or larvae of these species are
exposed to significant pollution by hydrocarbons or other substances associated with drilling.
(Based on A. Rieser and J. Spiller, 1980, work in progress)

The

OIL

LEASE TRACTS

Legal

issues

by Daniel P. Finn

SPAWNING ARIEAS

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WHITING
RED HAKE

41'

67'

On December 18, 1979, the United States
Department of the Interior held Outer Continental
Shelf (OCS) Oil and Gas Lease Sale No. 42 for
Georges Bank, a sale that had been delayed nearly
two years because of legal and political difficulties.
Bids were accepted on 63 of the 116 tracts offered-
each about 22.5 square kilometers in area. In the
spring of 1980, bidders filed exploration plans that
would allow them to begin drilling. However,
further complications will no doubt arise during
subsequent stages of the approval process, because
of continued consultation between Interior and
other federal agencies, review by the states, and
participation by the public. For example, additional
environmental impact statements (EIS) will be
prepared, and further litigation by the states or
outside interest groups is also probable.

The lease sale itself generated considerable
controversy. It was the first formal opportunity to
address the most important issues regarding
whether or how oil and gas activities should be
conducted on Georges Bank. The location of the
tracts offered and the terms and conditions of the
leases are significant in determining the effects of
such activities. Also, the decisionmaking of the
federal agencies and other parties priorto the sale is

a good indication of how future decisions will be

made.

The sale was held after considerable public
disagreement by two federal agencies — the
Department of the Interior, which regulates oil and
gas activities and conducts OCS lease sales through
its Bureau of Land Management, and the National
Oceanic and Atmospheric Administration (NOAA),
which is responsible for fisheries conservation and
management through its National Marine Fisheries
Service (NMFS). NOAA also administers the marine
sanctuaries program, which allows the Department
of Commerce to apply special regulations to protect
especially valuable marine areas. Before the lease
sale could be held, the Carter administration had to

develop a unified position on the terms and
conditions of the sale in order to reconcile the views

of the agencies and to demonstrate to the public
that necessary safeguards had been adopted. As a
result, certain potential lease tracts were deleted
and an interagency committee was formed to advise
Interior on future regulatory measures. Off-site
disposal of drilling effluents was not adopted,
however, meaning that drilling inevitably would
result in a certain degree of pollution. The way this
decision was made raises several interesting legal
issues.

Productivity of Georges Bank

Georges Bank is an extremely productive
commercial fishery area. Annual landings of fish
total about $168 million, which results in a net
economic benefit of more than $1 billion. Although

this represents only a small part of New England's
economy, the fishing industry is concentrated in a
few coastal areas and is an important cultural and
social factor in the life of the region. It is projected
that with proper management Georges Bank could
sustain annual landings of $229 million.
Hydrocarbon reserves, by contrast, are estimated at
123 mil lion barrels of oil and 870 billion cubic feet of
gas, havinga netvalue — less costs of production -
of $588 million. Oil from Georges Bank would
probably supply 5 to 6 days of current national
consumption. Also, a major gas strike could mean
lower prices and greater availability of natural gas
for New England.

The productivity of Georges Bank fisheries,
which include both demersal species (cod,
haddock, pollock, and flounder), and benthic
species (lobster, scallops, and quahogs), appears to
result from a unique combination of oceanographic
conditions in the area, still not fully understood.
The area is characterized by an enclosed or
semienclosed current gyre, strong vertical mixing,
and upwelling. The mixing and upwelling cause
considerable nutrient enrichment and therefore a
high rate of primary productivity. The current gyre
tends to concentrate nutrients and retain plankton,
and causes eggs and larvae of commercial species
generally to remain in the area of high productivity
and develop into strong year-classes.

Several substantive issues have been raised
by groups concerned with the threat drilling poses
to the fisheries. Some scientists fear that oil being
released by routine spills during recovery and
transportation and by discharge of pore waters from
oil-bearing sedimentary strata (formation waters)
during production could lead to chronic pollution
of the water column, resulting in fish egg and larvae
mortality and alteration of planktonic composition.
Some think that because of the intense vertical
mixing, discharged oil and other substances, such
as drilling muds and cuttings, could become
incorporated into sediments or suspended with
sediment particles. Some also believe that the
current gyre could lead to any large spills being
retained within the area, causing damage to
developing year-classes of commercial species,
since their eggs and juveniles are present over large

parts of the Bank during nearly the entire year.
On-site discharge of drilling muds and cuttings
could result in localized toxic effects on benthic
communities, including lobsters, either in areas
adjacent to drilling facilities or in areas subject to
downslope movement of pollutants — for example,
in the canyons that start on the continental shelf
edge. These factors have led some scientists to
conclude that oil and gas development could have
significant effects, both chronic and acute, on the
Bank's fisheries productivity.

These unresolved issues are still undergoing
extensive environmental analysis and will not be

30

46

45-

44-

43H

42-

41-

40-

39

sea le
0 50 100km

71

70 69 68 67 66 65

Direction of surface currents during the spring, illustrating
the current gyre.

discussed here. Ourfocus will be on legal questions
regardingthe sale, including Interior's
responsibility to protect fisheries and to consider
management alternatives that would be more
protective of marine resources, and the effects of
NOAA's actions in considering the designation of
Georges Bank as a marine sanctuary. While these
are not the only legal issues that were raised in
connection with Lease Sale No. 42, they provide an
unusual glimpse into federal decisionmaking and
point up a number of general concerns about the
way important decisions affecting marine resources
are made.

History of the Lease Sale

A lawsuit against the lease sale was brought in
December, 1977, by the State of Massachusetts and
the Conservation Law Foundation of New England,
a public interest group. The First Circuit Court of
Appeals determined that before proceeding with
the sale, the Secretary of the Interior, Cecil D.
Andrus, should ensure that the sale would not
create an unreasonable risk to fisheries. The Court
also commented that the Secretary should consider
the possibility of the area being managed by the
Department of Commerce, through NOAA, as a
marine sanctuary. Under the Marine Sanctuaries
Act of 1972, the Secretary of Commerce, in this case
Acting Secretary Luther Hodges, is empowered to
designate ocean areas as marine sanctuaries to
preserve their resources. Within such areas,
activities may be directly regulated by the

46

45-

44-

43-

42-

41-

40-

39

spill site

scale

0 50 100km

71

70 69 68 67 66 65

Projected area affected by a large spill of 34, 840 metric tons
of oil occurring at the site indicated in early spring after 30
days, plotted against a modeled distribution of fish larvae.
The white circle represents the area in which the ambient
concentration of hydrocarbons in the water column would
exceed 50 parts per billion (ppb). Significant or even
complete mortality of fish eggs and larvae could occur in
areas of 50 ppb concentration. The dark spot represents
the area in which oil would become entrained in
sediments.

Department of Commerce; in addition, actions
proposed by other federal agencies must be
certified by Commerce as consistent with the
purposes of the sanctuary.

The novelty of the Court's conclusions in the
Massachusetts v. Andrus case was that henceforth,
in deciding whether and howto proceed with this
and other lease sales, Interior would have to
consider the goals of a statute administered by the
Department of Commerce — the Fishery
Conservation and Management Act (FCMA) — and
even the possibility that Commerce could manage
the area as a marine sanctuary with objectives that
would likely be differentfrom its own. Interior, after
all, is primarily responsible for conducting the
government's OCS leasing program, whereas
Commerce (through NOAA) is primarily interested
in conserving and managing commercial fisheries
under the FCMA and protecting unique areas
through its marine sanctuaries program.

After the Court's decision, Interior prepared
a supplemental EIS that specifically addressed
marine sanctuary alternatives and considered
fisheries issues in greater detail. Before the draft of
this document was released, the Conservation Law

31

Boundaries of marine sanctuary
proposed by the Conservation
Law Foundation. Jurisdiction
over the area between the lines
marked "Canadian Claim" and
"U.S. Claim" is disputed by the
two countries.

Georges\ Bank
Marine Sanctuary
Nomination

Foundation — one of the plaintiffs in the pending
lawsuit — petitioned the Secretary of Commerce to
designate Georges Bank as a marine sanctuary and
manage the area so that the primary federal
objective would be fisheries production and
conservation; oil and gas activities would be subject
to additional regulations to ensure this objective
was achieved. Upon receiving the petition, NOAA
selected the area as an active candidate for a marine
sanctuary. It also provided extremely critical
comments on the treatment of fisheries issues and
the marine sanctuary alternative in Interior's
supplemental EIS, and specifically called Interior's
attention to the petition that it had received. NOAA
then released an "issue paper" on potential
management of Georges Bank as a marine sanctuary
and held a series of public workshops in
Massachusetts and Maine. Among the alternatives
presented in the paper were proposals to limit
future oil and gas activities on Georges Bank to the
proposed Lease Sale No. 42 area, to delete from the
sale those tracts on or near valuable benthic
communities, and to impose additional operational
regulations, including barging away of drill ing muds
and cuttings, reinjection of formation waters, and
on-site location of pollution control and

containment equipment. NOAA also called for
more monitoring of potential biological effects by
lessees. Finally, NOAA argued that even if a marine
sanctuary were not designated, additional
measures would be necessary to make the terms of
the sale protective enough and to ensure that
Interior continued to carry out its responsibilities
properly. These included the formation of a
standing interagency committee to oversee
operations, and definite interagency and public
reviews of various future actions.

Interior reacted very strongly to NOAA's
proposals. Interior claimed that it had primary
authority to regulate oil and gas activities and was
already obliged to ensure that there was no
unreasonable risk to fisheries. Interior indicated
that it perceived no basis in fact for believing that
NOAA could do a better job than it would. Afterthis
exchange, both Interior and NOAA proceeded
toward their own decisions about how to manage
Georges Bank, whiletheCarteradministration tried
to formulate a unified position on the pending
lawsuit and the outstanding issues concerning
fisheries and the possibility of a marine sanctuary.
Finally, on September 21 , 1979, Commerce and
Interior announced that they had reached an

32

agreement. NOAA agreed to withdraw Georges
Bank from its list of active candidates for marine
sanctuary designation, and Interior consented to
theformation of an interagency biological taskforce
that could recommend actions to its operational
official, the Regional Supervisor of the U.S.
Geological Survey (USGS). Interior also agreed to
include certain operational measures either as
stipulations to the leases or as information to
lessees that would give them notice of how their
activities would be regulated. (Including regulatory
provisions in the leases or in the notice of lease sale
is important because the sale of leases creates
private property rights in favor of lessees, who have
paid the government significant sums — in the case
of Georges Bank, in excess of $800 million — and
who will probably invest considerable amounts in
exploratory and other activities with the expectation
of realizing certain levels of economic return on
their leases.)

Regardless of its concessions, the agreement
has been widely viewed as a victory for Interior.
NOAA's issue paper and its comments on Interior's
supplemental EIS presented a strong case that
certain actions were essential to conserve Georges
Bank fisheries. Few of NOAA's recommendations
were followed in the agreement, however. The new
language of the lease stipulations concerning
drilling muds and cuttings and formation waters
called only for undefined "safe" disposal of such
substances; Interior had never previously exercised
its rightto require bargingand reinjection in similar
cases. Lessees were not made subject to extensive
biological monitoring requirements; additional
monitoring would have to be made largely by
federal agencies and others at their own expense.
NOAA's recommendation of on-site location of
cleanup and contingency equipment was adopted,
however, and 12 of the 15 tracts recommended for
deletion by NOAA were removed from the sale.

A key issue was the terms of the charter
developed by Interior and NOAA to establish an
interagency biological taskforcethat was to include
officials of these organizations and the
Environmental Protection Agency. The task force is
not empowered to make binding decisions under
its charter. If the USGS Regional Supervisor
disagrees with recommendations of the task force,
he must state his reasons, however. But three of the
five agencies represented on the task force are
within the Department of the Interior — the Bureau
of Land Management, Fish and Wildlife Service, and
USGS. Although any two agencies in the task force
mayappeal actions of the Regional Supervisor, such
an appeal must be based on the Regional
Supervisor's failure to follow a recommendation of
the task force as a whole — that is, a
recommendation adopted by at least three of its
members.

69 68

67' 66

70 69 68

67 66'

/Areas proposed or actually leased in Lease Sale No. 42: (A)
the 178 tracts originally presented in Interior's final EIS; (B)
the 128 tracts in Interior's supplemental EIS, including the
72 tracts deleted by Interior (open boxes) upon agreement
with NOAA; (Q the 63 tracts actually sold.

33

Reactions to the Agreement

After Interior and Commerce reached this
agreement, the states of Maine and Massachusetts
and the Conservation Law Foundation, which had
petitioned the Secretary of Commerce for a marine
sanctuary, returned to court with additional claims
concerningthe lease sale as it was then proposed by
Interior and the actions that had been taken (or not
taken) by NOAA. They claimed that Interior's
proposed sale was still defective for failing to
protect fisheries and endangered species and to
consider the marine sanctuary alternative in its
supplemental EIS. They also asserted that NOAA
had failed to fulfill its duties by not proceeding with
a formal proposal fora marine sanctuary, a proposal
that it apparently had felt was desirable and perhaps
even essential for adequate protection of the
Georges Bank fisheries. NOAA, they asserted,
should have at least prepared an EIS so that
alternative proposals could be publicly considered.
The plaintiffs also claimed that NOAA's failure to
proceed further affected the validity of the sale,
since had NOAA proposed or designated a
sanctuary, Interior's action might have been
different.

Hasty judicial proceedings followed, since
Interior proposed to hold its sale on November 6,
1979. First, the Federal District Court and then the
First Circuit Court of Appeals, hearing the case as
Conservation Law Foundation v. Andrus, heard
arguments right up to the eve of the sale as then
proposed, and the First Circuit Court issued its
opinion the very morn ing of the proposed sale. The
courts rejected the new contentions of the
plaintiffs. They concluded that Interior had
adequately addressed the fisheries and marine
sanctuaries issues in its supplemental EIS and that
NOAA was apparently under no legal obligation to
consider further any alternative management plan
for the Bank. Although the sale proposed for
November 26, 1979, was delayed by last-minute
proceedings in the United States Supreme Court,
no significant judicial obstacles remained and the
sale was finally held on December 18.

The Marine Sanctuary Issue

The plaintiffs were not successful in pressing their
claims that NOAA was obliged to prepare an EIS on a
marine sanctuary, and that Interior would have to
defer to NOAA's administrative process because it
was required to consider the alternative that
Georges Bank be managed as a marine sanctuary.
The plaintiffs had a difficult case. They were arguing
that NOAA had failed to undertake a required
action. Regarding the lease sale itself, they were
making a complex claim — that Interior was
required to await information generated by NOAA,
and even to respect the autonomy of NOAA's
competing administrative process, since it had to

consider NOAA's actions which could affect how oil
and gas operations would be conducted under
lease.

Ordinarily, a federal agency is free to
proceed with any proposal within its statutory
authority provided that it meets all the legal
requirements in existence at the time of its action.
The original opinion of the First Circuit Court meant
that Interiorwas requiredtoconsiderthesanctuary.
But why should Interior's action be further delayed
or even brought into question simply because
NOAA was tentatively considering a
counter-proposal that could have affected Interior's

34

Working Georges Bank in rough weather. (Photo by NubarAlexanian)

action? NOAA at this stage had not issued a formal
proposal or even a draft EIS presenting definite
options. Indeed, NOAA had begun considering a
marine sanctuary only at the last minute, in
response to a petition submitted by plaintiffs in
pendinglitigation with Interior. Someof the groups
on whose behalf the petition was submitted were
involved in commercial fishing, and thus were in
effect NOAA's "clients" under the FCMA.

Several aspects of the actions taken by both
agencies and the nature of the relationships
between them nevertheless suggest that there was a
legal basis on which the courts could have

prevented the lease sale. First, it is true that it was
Interior which was required to consider the marine
sanctuary as an alternative to its own proposed
action. Nevertheless, NOAA would be required to
proceed in a regular and good-faith manner in
implementing programs committed to its
administration, like the marine sanctuaries
program. Second, although NOAA had not made a
formal proposal for a sanctuary, it was required by
its own regulations to decide within 90 days after
having held public workshops whether to proceed
with a formal proposal. Third, it is indicated in the
public record that NOAA believed the actions

35

CHRONOLOGY OF EVENTS RELATED TO THE GEORGES BANK LEASE SALE

June 17, 1975

Jan. 2, 1976
Oct. 12, 1976
Dec. 7, 1976

Dec. 7-10, 1976

Aug. 29, 1977
Dec. 28, 1977

Jan. 28, 1978

Jan. 31,1978
Feb. 22, 1979

May 10, 1979
May 25, 1979

June 20, 1979
July 16, 1979

July 27, 1979
Aug. 22-24, 1979
Sept. 21,1979

Oct. 5, 1979
Nov. 5, 1979

Nov. 6, 1979

Nov. 9, 1979
Dec. 17, 1979
Dec. 18, 1979

Department of the Interior (DOI) called for nominations of tracts to be included in

proposed Lease Sale No. 42 on Georges Bank. 1,927 tracts were nominated.

DOI proposed a list of 206 lease tracts.

DOI published a draft environmental impact statement (EIS).

DOI deleted 28 tracts because of uncertainty of boundary with Canada, then being

negotiated.

DOI held public hearings on the lease sale and its environmental effects, detailed in

the draft EIS.

DOI issued a final EIS, in five volumes.

DOI deleted 23 tracts to minimize interference with commercial fishing, including

trawling and setting and hauling of lobster pots, and to reduce danger of a spill

reaching shore.

Commonwealth of Massachusetts and Conservation Law Foundation (CLF) and

others obtained a preliminary injunction against proposed sale from District Court.

First proposed sale date.

First Circuit Court dissolved the preliminary injunction but instructed DOI to

consider further fisheries and marine sanctuaries issues.

CLF and others petitioned the Department of Commerce (DOC) to designate

Georges Bank a marine sanctuary.

DOI issued a draft supplemental EIS addressing fisheries and marine sanctuaries

issues.

DOI held public hearings on the draft supplemental EIS.

National Oceanic and Atmospheric Administration (NOAA), a DOC agency,

submitted comments on the supplemental EIS, criticizing DOI's discussion of

fisheries issues and calling attention to CLF's petition.

NOAA released an "issue paper" discussing the possibility that Georges Bank could

be designated a marine sanctuary.

NOAA conducted workshops in Massachusetts and Maine about the marine

sanctuary.

DOI and DOC announced an agreement that would allow the sale to proceed; the

Administrator of NOAA indicated his dissatisfaction with the terms of the sale.

DOI issued a notice of sale for November 6, 1 979.

District Court ruled against plaintiffs' new contentions about the duties of DOI and

NOAA.

First Circuit Court refused to grant a stay delaying the proposed sale; Justice Brennan

of the Supreme Court granted a temporary stay, however.

U.S. Supreme Court vacated Justice Brennan's order.

First Circuit Court rejected appeal from District Court's order of Nov. 5, 1 979.

DOI held Lease Sale No. 42.

proposed by Interior, and even its own agreement
with Interior, were inadequate to protect the
Georges Bank fisheries. The terms of the lease sale
did not address several substantial concerns raised
by NOAA in its comments on Interior's action.
Furthermore, the administrator of NOAA, Richard
A. Frank, made statements at the time the
agreement was announced that indicated he
thought the sale as proposed still presented
significant dangers to the fisheries and should not
have been held.

If NOAA had proceeded with a proposal to
designate Georges Bank as a marine sanctuary, its
action could have significantly affected oil and gas
activities on Georges Bank. It could be argued

therefore that NOAA's failure to proceed with an
action that was within its authority and that it
apparently believed was necessary allowed Interior
to go forward with its lease sale. Thus it appears the
courts had grounds for preventing Interior from
holding the sale because, if the sale was stopped,
NOAA could have fulfilled its statutory
responsibilities to protect the fishery resources and
to give proper consideration to designating a
marine sanctuary. Issuing leases could prevent
NOAA from establishing a sanctuary later.

In its formal notice withdrawing Georges
Bank from consideration, however, NOAA stated
that the agreement it had reached with Interior
significantly increased the ability of the existing

36

Fishermen's wives protesting lease sale outside White House in Washington, D. C. (Photo by Nubar Alexanian)

regulatory framework to protect the fisheries and
made it more likely that sufficient effort would be
devoted to protection, making a marine sanctuary
unnecessary. The courts found nothing in the
public record that indicated NOAA had abused its
administrative discretion, and the First Circuit Court
specifically found that NOAA's action was not
necessarily inconsistent with its earlier position on
the risks of oil and gas operations to the fisheries.

Other aspects of the situation at the time
Interior and NOAA reached their agreement
present additional grounds for concern, however,
loan outside observer, it mayappearthatthere was
strong pressure within the Carter administration for
NOAA to discontinue considering the sanctuary
and to reach an agreement with Interior. NOAA may
have received indications within the executive
branch that the President would not approve
designation of a marine sanctuary. (The statute
requires that all marine sanctuaries must be
approved by the President). If this is true, then
important public interests may have been sacrificed
because of the way in which the interagency

agreement between Interior and NOAA was
concluded.

Although the Marine Sanctuaries Act
expressly provides that the Secretary of Commerce
must consult with otherfederal agenciesand secure
the approval of the President before designating a
marine sanctuary, in this case NOAA stopped well
short of the President's desk. NOAA never issued a
formal proposal or a set of options that would have
permitted full public evaluation of the desirability of
establishing a sanctuary. Whatever process of
accommodation that was accomplished with the
executive branch was carried out informally. The
President did not have the opportunity to evaluate
the conflict between Interior and NOAA in the
context of two well-developed competing
proposals. Similarly, Congress, which has an
interest in the effective implementation of the laws
it has enacted, had no opportunity to oversee the
performance of the Department of Commerce or
the Administration in implementing the Marine
Sanctuaries Act. Finally, because the important
issues were submerged and an internal agreement

37

Gulf Oil executive talks with
State Trooper after being hit in
neck with a bag of oil tossed by
a protester at Georges Bank oil
lease sale on December 18,
7979, in Providence, R.I. (Photo
courtesy Providence Journal-
Bulletin)

was reached, the courts reviewed the final decision
in terms of technical legal issues about the
procedural responsibilities of Interior and NOAA.
For all these reasons, public scrutiny and
participation were limited and the legitimacy of the
ultimate decision was affected. All these factors
could have provided a legal basis for preventingthe
lease sale.

Lessons of the Georges Bank Lease Sale

A wide range of views is held among the general
public and among scientists and government
officials about whether oil and gas activities should
be permitted on Georges Bank and how such
operations should be regulated. Regardless of one's
position, the Georges Bank lease sale raised critical
questions regarding how effective and legitimate
governmental actions can be achieved in situations
where significant marine resources are at stake. It is
important to realize that even the way such
decisions are made has important consequences for
the successful resolution of conflicts between
development and conservation interests in marine
areas.

Numerous statutes have created overlapping
responsibilities among federal agencies often with
conflicting missions and different perspectives on
how marine resources should be managed. If these
agencies fail to conduct their proceedings in an
open and well-coordinated manner, significant

questions will be raised about their decisions
whatever the final outcome. Although important
decisions have yet to be made on how oil and gas
operations should be conducted on Georges Bank,
the history of the lease sale itself shows that we still
have a long way to go in developing institutions that
can bear the strain of conflicting pressures.
Whatever the ultimate decisions, however, they
should be reached in a public and principled
manner so that conflicting opinions of fact and
value about the wisdom of proposed actions are
faced squarely and openly.

Daniel P. Finn, a lawyer, is a Research Fellow with the
Marine Policy and Ocean Management Program at the
Woods Hole Oceanographic Institution. He also has
worked as an attorney for the National Oceanic and
Atmospheric Administration.

38

lopment of

Science

aim

n ffl

menca

Allegorical representation of Magellan's trip around the world. (The Bettmann Archive, Inc.)

by Francisco J. Palacio

I he large contributions to the exploration of the
oceans made by the Spanish and the Portuguese
about five centuries ago have not been matched
subsequently by their descendants. Marine science
in Latin America today is generally underdeveloped,
impoverished. In most cases, there is no strong
national commitment to develop and manage the
coastal zone. Education in the various disciplines is
minimal — this, infact, is the case for most scientific
endeavors in general. And yet — the potential is
tremendous. There is a stirring of interest in the
oceans once again, but Latin nations must look not
to international agencies and institutions to solve

their problems, but inwardly, to themselves, to
theirown pool of human resources. Thetask before
them is monumental, but not impossible. Let us
briefly review the major contributions of the past,
the early events that set the foundations of the
present status of the marine sciences in Latin
America,* and touch on some trends, problems,
and programs.

*The term Latin America is used to include mostly, but not
exclusively, the countries of continental Central and
South America.

39

History

John Murray's superb 1895 historical account of
marine investigations prior to the voyage of the
H.M.S. Challenger recognizes Ferdinand
Magellan's deep-sea soundings in 1521 (in the
present Tuamotu Archipelago) as the first recorded
in the open sea. Magellan's voyage for Spain's
Charles V also was the greatest event in the
thirty-year period from 1492 to 1522, when Spanish
exploration doubled knowledge about the earth,
adding a hemisphere to a world that could, indeed,
be circumnavigated. But the discovery of new lands
to be colonized, natives to be proselytized, species
to be traded, and precious metals to be mined, all
led Spain to change its maritime interests from
exploration to the transport of goods. This was
based upon a land economy supported by a lavish
use of Indian and Negro labor, the character of
which represents an important part of today's
problems in Latin America.

Across the English Channel, Drake, the
defeat of the Armada in 1585, and the Treaty of
London in 1604 opened the doors of the New World
to England and Europe in the 17th century. A
century later, Captain Cook, Harrison's
chronometer, and the sextant opened the Pacific
Ocean to the British Admiralty. Less well known is
the fact that Spain also explored the Pacific. Alvaro
de Mendana (1567), Fernando de Quiros (1595),
Vaez de Torres (1606), and other smaller
expeditions stirred by British and French
discoveries, contributed significantly to the
explorations of the South and North Pacific. Spain's
resentment of European intrusion resulted in a
policy of secrecy of discovery that contrasts with the
numerous publications on the voyages of Cook, La
Perouse, and Vancouver. The diplomatic pressure
exercised by greater military strength forced Spain
in 1790-95 to give up its claims and settlements to
Great Britain. As a result, the world at large did not
realize that at the turn of the 18th century, the
Iberians had made more charts and maps of the
newly discovered regions than any other group of
navigators. By then, not only English vessels, but
also their maps, language, and literature began to
rule the seas.

Although a methodic study of the Peru
current was not made until Humboldt undertook
one in 1802, Pizarro and his followers were well
aware of the force, impact, and vagaries of the
current. We owe the discovery of the Galapagos
Islands, already known to Inca rulers, toan attempt
by Panama's bishop to avoid the current en route to
Peru to settle a dispute between Pizarro and
Almagro. Early sailing directions were compiled by
Cieza de Leon in 1553, and the current was
discussed by Father Acosta in 1608, who identified
the attributes of El Nino (seeOceanus, Vol. 21, No.
4, p. 40). For two centuries, from 1600 to 1800,

Magellan
(The Bettmann Archive, Inc.)

knowledge of the current was derived from short
references in journals and logs. There was a general
feeling that the cool water off the west coast of
South America was related to the height of the
Andes. In the Gulf of Panama, where the Andes are
relatively low, the waters were warm, whereas off
Peru , where they are high, the water was cool. This,
and cloud formations were supposed to have a
cooling effect on the waters. Humboldt made
temperature measurements, concluding that the
water was cooler than the air and thus could not be
cooled by it. He also observed that surface
temperatures rose with increasing distance from
the shore, concluding that the mass of cold water
flowed from higher latitudes. This view was
disputed by Bouganville in 1837; in 1844, de Tessan
suggested an upwelling of the lower layers of water
might account for the low surface temperatures.

The publication of observations by European
visitors and naturalists in the New World reflects a
bias in favor of foreign information over local
knowledge. The first comprehensive report on the
wonders of the New World was Fernandez de
Ov\edo's'\52()Natural History of the West Indies. Its
great popularity in Europe stimulated the
imagination of many future colonizers. Oviedo's
work marked the beginning of a period of reports

40

that continued into the middle of the 19th century,
by which time Linnaeus developed his system of
binomial nomenclature, providingthe mechanism
for classification of living entities.

Duringthe middle of the 19th century, the
interest of maritime nations in exploring marine
resource frontiers (mainly seals and whales), in
charting the waters to reach and exploit the
resources, and in studying the existence of life in
thedeepsea, ledtoseveral expeditions — themost
important for Latin America being the U.S.
Exploring Expedition (1838-42), Darwin's voyage
aboard the Beagle (1831-36), the Challenger
Expedition (1872-76) , the work by Alexander Agassiz
on board the Blake (1877-80), and the work of the
Albatross over a number of years. The majority of
this research fell in the area of marine biology, with
physical, chemical, and geological aspects
developing largely during the 20th century.

During the 19th century, virtually all the
research that was being done in Latin American
waters was conducted by foreign maritime powers;
almost nothing was contributed by Latin countries.
The abundance of land resources combined with
relatively small populations meant there was little
need for ocean exploration. To this day, agriculture
based on inexpensive labor continues to be the
most important aspect of Latin American
economies, with political power generally
concentrated among the landed aristocracy.
Although several capitals and major cities have been
founded along the coasts, an awareness of the
potential of the oceans is only a very recent
phenomenon. Fishing has always been relegated to
the lower social and economic strata and disdained
as a form of servile manual labor.

Spain controlled its immigration to the
colonies so that it could maintain the purity of the
ruling stock, preserve the wealth, and have a strong
voice in church affairs. The historical circumstances
that were adverse for the development of western
science in Spanish America existed well into the
19th century. The settlement of the interior of
southern South America increased the need for
immigrant labor and as political conditions became
more stable, European immigration rose rapidly in
the latter half of the century. This had a significant
influence on the cultural environment and brought
changes in the educational system. The early
advance of science in Argentina, southern Brazil,
and Chile can be traced to European immigrants and
their interest in developing the resources of their
new homelands. It was in these countries where
concern for the oceans developed around the turn
of the century; Mexico, too, showed interest in
developingits marine resources, benefitingfrom its
close proximity to the United States.

While the foundations for marine biological
investigations in the United States were laid at the
end of the 19th century, Latin America's scientific

Pizarro
(The Bettmann Archive, Inc.)

interests were limited mostly to botanical studies.
Prior to World War I , basic work on the propagation
of sound in the water was being investigated in
several countries (see Oceanus, Vol.20, No. 2, p. 8);
Germany's development soon after of submarine
warfare prompted the Allies to speed up their study
of underwater sound as a method of detecting
U-Boats. The war had a fundamental effect on three
events. First, the flow of information from European
universities and the training of scientists in Europe
were reduced. This caused greater self-reliance in
the United States. Second, there wasa mobilization
of scientifictalentamongthe Allies to study all naval
problems related to submarines. And third, the U.S.
government firmly established the precedent of
strong support for marine science.

As World Warl began, Latin America was still
completely dependent on European and, to a lesser
extent, American technology and science. Only in
the area of medicine was there significant Latin
accomplishments. Not being directly involved in
the war, Latin American governments found no
pressing need to develop a naval science or
technology. The educational system, classical in
nature and relatively unaffected by European
scientific pursuits, was scientifically stagnant.
Furthermore, there was no need, indeed no desire,

41

to support the development of indigenous science.
The children of the ruling minorities were generally
educated abroad. National educational programs
found only limited support. The system was geared
toward preparing young aristocrats for traditional
professions and commercial undertakings, which
directly or indirectly perpetuated the oligarchies. A
career in the sciences was an extremely rare choice.
The scientific progress that did take place in
Latin America — and there was some — developed
mainly because of the determination of a few
persistent, outstanding individuals rat her than from
national commitments. The development of
science in Latin America also suffered because of
the isolation from new discoveries as well as from
the neglect of the international scientific
community.

The 20th Century

The spread of western science in Latin America has
occurred in three progressively overlapping stages
-foreign exploration and investigation, the
establishment of a dependent colonial science
(founded on advances made abroad and adopted
locally), and the completion of the process of
transplantation. In the marine sciences, most Latin
American countries a re evolving from the first to the
second stage, with Mexico, Argentina, and Chile
moving into the third stage. The main emphasis in
all countries is overwhelmingly in the area of marine
biology, with concentration on taxonomic studies
stronglyinfluenced by specialists workingin Europe
and the United States. Other disciplines are still
very weak in most countries, with the exception of
Argentina, Brazil, Venezuela, Colombia, and
Mexico. The Lamont-Doherty Geological
Observatory at Columbia University in New York
State and the Woods Hole Oceanographic
Institution have aided these latter countries in the
development of their marine science programs.
The emphasis on biological work can be
traced to the influence of immigrants in southern
South America, who promoted the study of local
faunas and trained a generation of biologists. This is
in marked contrast to the United States where
scientific talents were mobilized to deal with the
oceanographic demands of two world wars, which
greatly expanded the fields of marine geology,
physical and chemical oceanography, and
meteorology.

The period of foreign coastal exploration and
the charting of Latin American coastal regions was
followed by the establishment of hydrographic
offices at the turn of the 20th century; then, there
was a period marked by preliminary observations
and the establishment of natural history collections,
after which came the biological institutes, most
importantly the University of Chile's Montemar
marine biological station at Vina del Mar in 1941 ,
and, in 1946, the University of Sao Paulo's

Oceanographic Institute. The Institute of Marine
Biology at Mar del Plata in Argentina was not
established until 1960 (Table 1).

Latin America's fear of exploitation of its
marine resources by foreign powers led to
jurisdictional pronouncements. In 1952, Peru,
Chile, and Ecuador issued the Declaration of
Santiago, establishing 200-mile territorial waters.
The concern over resources also resulted in the
creation of the Permanent Commission for the
South Pacific, which promoted the development of
fisheries institutes, such as Ecuador's Institute
Nacional de Pesca (1961), Chile's Institute de
Fomento Pesquero (1963), and Peru's Institute del
Mar (1964).

The first Latin American congress on marine
biology took place in Vina del Mar, Chile, in 1949.
But the need for inter-American cooperation in the
field was not recognized until a conference in
Caracas, Venezuela, in 1954. A conference
document recommended the creation of an
Inter-American Oceanographic Institute (to be
located in the Galapagos Islands) under the
authority of a central Oceanographic Commission;
additional regional centers would study biological,
fishery, and ecological problems. At this time, only
Argentina, Brazil, Chile, Cuba, Ecuador, Mexico,
Peru, and Venezuela were actively interested in
marine resources. The first meeting of general
marine specialists took place in Sao Paulo, Brazil, in
1955. Under the auspices of the Panamerican
Institute of Geography and History, a subsequent
meeting of the working group on oceanography
was held in Washington in March of that year. This
meeting produced a report that emphasized the
need for cooperation and exploration in the marine
field and also identified areas of interest. In 1956,
preliminary national reports on the resources of the
continental shelf were presented at a conference in
the Dominican Republic; the coordination
responsibility for these activities was delegated to a
representative oceanographic committee,
headquartered in Mexico City, Mexico. Later that
year, the first meeting of UNESCO's* International
Advisory Committee on Marine Sciences took place
in Lima, Peru. The 1957 International Geophysical
Year had the effect of strengthening scientific
cooperation and, by this time, the United Nations
was playing an important role in scientific efforts. In
1960, UNESCO's semiautonomous coordinating
Intergovernmental Oceanographic Commission
(IOC) was established to advise, promote, and
catalyze ocean-related endeavors. Also, UNESCO's
operational Division of Marine Science, upon
request by member states, was charged with
promoting research programs, disseminating
information, and strengthening national

"United Nations Educational, Scientific, and Cultural
Organization.

42

Table 1 . Distinguished marine institutions in Latin America and date of founding.

Argentina

Museo "Bernardino Rivadavia" Estacion

Hidrobiologica de Puerto Quequen 1 923

Institute Antartico Argentine 1951

Institute de Biologia Marina, Mar del Plata

(INIDEP) 1960

Centro de Investigacion de Biologia Marina 1 961

Comision Nacional de Estudios

Geoheliofisicos Centro Nacional

Patagonico 1968

Institute Argentine de Oceanografia 1 969

Brazil

Institute Oceanografico, Universidade de

Sao Paulo 1946

Departamento de Oceanografia, U. Federal

Pernambuco 1952

Laboratorio de Ciencias do Mar, U. Federal

Ceara 1960

Instituto de Pesquisas da Marinha, Rio de

Janeiro 1962

Base Oceanografica Atlantica, Rio Grande

doSul 1971

Chile

Estacion de Biologia Marina, Montemar,

Vina del Mar, Universidad de Chile 1 941

Departamento de Biologia Marina y

Oceanografia, Universidad de

Concepcion 1957

Instituto de Fomento Pesquero 1963

Colombia

Instituto de Investigaciones Marinas, Punta
deBetm 1966

Cuba

Centro de Investigaciones Pesqueras 1 952

Instituto de Oceanologia 1959

Ecuador

Instituto Nacional de Pesca 1961

Instituto Oceanografico de la Armada 1 972

Mexico

Instituto Nacional de Pesca 1962

Centro de Ciencias del Mar y Limnologia

Universidad Nacional Autonoma de

Mexico 1973

Centro de Investigacion Cientifica y

Educacion Superior de Ensenada 1975

Instituto Oceanografico de la Armada 1 978

Peru

Instituto del Mar del Peru 1964

Puerto Rico

Departamento de Ciencias Marinas,

Universidad de Puerto Rico 1 954

Venezuela

Instituto Oceanografico, Universidad de

Oriente 1959

Estacion de Investigaciones Marinas,

Fundacion La Salle 1960

infrastructures, with emphasis on education and
training. These tasks were performed in
cooperation with the United Nations Development
Program. During the 1960s, the Latin American
countries that had neglected their marine affairs in
the past became actively involved, joint research
investigations were undertaken with
oceanographic institutions, such as the Scripps
Institution of Oceanography, Oregon State
University, the Woods Hole Oceanographic
Institution, Lamont-Doherty, and the University of
Miami; major faunal surveys were conducted in
tropical shelves and in deep-sea waters. Regional
conferences, training courses, and seminars were
held throughout South America. The period of the
mid-1960s also was characterized by the creation of
national oceanographic commissions in Latin
America to coordinate national and international
efforts. The Food and Agriculture Organization
(FAO) of the United Nations was bythistime playing
a major advisory role in developing Latin America's
fisheries. National fishery institutes were created,
some undergraduate marine programs were
initiated, and students began to go abroad for
advanced training, mostly with support from
international scholarships. In 1968, the need for
international cooperation was addressed by efforts
like the Cooperative Investigations for the
Caribbean Sea and Adjacent Regions (CICAR). These
efforts were not too cooperative nor investigative,
but regional governments became more attuned to
the concept of international marine research and
more important, became more realistic about
their own potential for harvesting and protecting
marine resources, and their own scientific and
technical limitations for addressing these efforts.

In 1970, the United States launched the
concept of the International Decade for Ocean
Exploration (IDOE[see Oceanus, Vol. 23, No. l|)
which was approved by the United Nations to
operate within the framework of a comprehensive
long-term program of scientific investigation. Latin
American countries became participants in several
of the IDOE programs (Table 2). In addition, the
Organization of American States (OAS) contributed
funds for fellowships and for equipment needs

43

Table 2. International oceanographic participation by Latin American countries. (Source: UNESCO-IOC)

International Decade for Ocean Exploration

LEPOR

Environmental Environmental Seabed Living

Forecasting Quality Assessment Resources

c/l

i/> "^ c

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~2 .E £ S

E

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Argentina

X

X X XXX

Bahamas

Barbados

Belize

Bolivia (landlocked)

x x x x x

Brazil

X

X XX X X X

Chile

X

x x x x

Colombia

X

XXX X X

Costa Rica

X

X

Cuba

X

X

Ecuador

X

X XX

El Salvador

France (Guadeloupe/Martinique)

X

Guatemala

X

X

Guyana

Haiti

X

X

Honduras

Jamaica

X

X X

Mexico

X

X XX

Netherland Antilles

X

X

Nicaragua

Panama

X

X X

Peru

X

X X

Rep. Dominicana

X

X

Surinam

X

Trinidad & Tobago

X

X

Uruguay

X

x x

Venezuela

X

X X X X X

within its Multinational Marine Science Program. In
the course of the IDOE, it was clear that most Latin
American countries were insufficiently prepared to
participate both scientifically and financially in
oceanographic programs as understood within the
IDOE. Moreover, the promise of initial United
States funding, estimated at $100 million to fully
implement the program, was only fulfilled with $15
million. Thus, although an effective machinery
developed for multinational cooperation, the
original hopes of a truly international scientific
program fell short of expectations, and the growth
of oceanographic competence in developing
countries played a much smaller role than originally
envisioned. Finally, in addition to the important
efforts of the Organization of American States to
promote cooperation and interchange, there have
been development activities promoted by
INTERCIENCIA, the federation of Associations for
the Advancement of Science in the Americas (Brazil,
Canada, Colombia, Costa Rica, Jamaica, Mexico,

the United States, and Venezuela), the Latin
American Association for Biological Oceanography,
and by the Latin American Committee for Marine
and Aquatic Products of SELA, the Economic System
for Latin America.

In brief, the Latin American countries have
emphasized their need for marine science and
technology and have looked to the International
Oceanographic Committee to satisfy those needs,
but the IOC does not have the mandate to
adequately meet these demands. Moreover, the
financial resources available under UNESCO's
regular program are insufficient to meet these
needs. Additional requests for assistance have been
submitted to the Food and Agriculture
Organization, the United Nations Development
Program, and the UN Environmental Program
(UNEP). The latter is particularly active in the area of
environmental quality through its Regional Seas
Program, and within its efforts with ECLA, the
Economic Commission for Latin America. The UN

44

Mussel culture experiment in
Dichato, Chile.

,*•>.-.

.•• -; .;-w sxi 4

Office of Ocean Economics and Technology also
has been active in promoting awareness of the
problems of the coastal zones in Latin America.

A Need to Look Inward

The growth of marine science in Latin America over
the last 30 years, mostly in the area of biology as we
have stated, has been characterized by a weakness
of national commitments, with high expectations
and responsibilities being placed on the efforts of
international bodies. No amount of funds or
technical support from international organizations
can produce effective benefits withouta willingness
on the part of national governments to utilize their
own available means of developing their marine
capability. Support for science, per se, has only
developed in the last few years, and then only in a
few countries. In fact, in view of the difficulties, the
efforts and advances made by some marine
institutions in Latin America have been remarkable.
Government support has been mostly aimed at
developing fisheries, with limited allocations for the
scientific component needed to exploit them
effectively.

The Peruvian experience is a case in point.
Support for basic scientific development relative to
the value of the fisheries was insignificant; ultimate
responsibility fell on FAO specialists and
international panels. Whereas the international
missions were partly successful, mismanagement at
the policy level contributed significantly to the
col lapse of the Peruvian fisheries. Furthermore, the
export of two thirds of Latin America's fish harvest
raises doubts about the sincerity of the concern for
the region's protein deficiencies, lendingcredibility
to claims that, at least in the countries with highest

catches, the hard currency gained by fishery exports
are being used to finance military expenditures.
There also is interest in develop! ngaquaculture, but
the emphasis has been on high-valued crustaceans
for export. The true potential of food for the rural
poor is not beingfully explored.

One of the fundamental barriers to the
growth of marine science in Latin America is the lack
of national science policies that would create an
appropriate environment for scientific
development. Although some progress has been
made, the key factor, and one difficult to assess, is
quality. When one has to labor under adverse
cultural circumstances, deficient basic scientific
training, and a lack of science policies and
government support, it is difficult to produce work
of exceptional quality. The common result is the
production of second-class science. This
regrettable reality is the rule rather than the
exception in the field of marine sciences in Latin
America. The truth is that many of the region's
marine institutions represent fragmented,
disorganized scientific communities, poorly
sustained, intellectually isolated, and generally
directed toward irrelevant goals. Fortunately, the
intellectual potential is quite promising; but the
distribution of intellectual resources is indepen-
dent of the political and economic factors needed to
bring them to fruition. There are now some 50
Ph.D. -level marine scientists in Latin America, all
working arduously to develop the field in their
respective countries. Most are quite young; they
have not yet reached the decisionmaking levels
that would allow them to change and improve their
environments. But there is reason for optimism:
their impact should be felt within the next decade.

45

Institute Oceanographico, Universidade de Sao Paulo,
Sao Paulo, Brazil.

Mariana

The pursuit of oceanographic investigations, as we
have known them duringthis century, will probably
come to an end with adoption of a United
Nations-sponsored Law of the Sea treaty. Indeed,
the way in which these expeditions and
investigations are undertaken has already changed
in the last few years. The new legal, organizational,
administrative, and financial requirements that
must be satisfied will probably impede, discourage,
or foil future scientific oceanic ventures.
Commercial undertakings, too, will probably only
be conducted within strict international legal
conditions. Consequently, three factors will affect
marine research efforts in the coastal areas: the
definition of economic zones, the growing impact
of man's activities on coastal ecosystems and
resources, and the high cost of the operation of
vessels as a result of spiralingfuel prices. Thus, the
outlook for Latin America is one in which there will
be an emphasis on coastal living, mineral, and
nonextractive resources.

The development of an infrastructure for the
study, exploitation, and wise utilization of the
coastal resources poses an interesting challenge for
most Latin American countries. Those countries
without, or with incipient, marine capabilities will
have to adopt effective science policies to create the
circumstances within which marine resources can
be incorporated into the process of development.
And those countries with some capabilities will have
to strengthen or develop the appropriate
mechanisms to exploit their growing expertise.

In addressingtheir common scientific
problems (with a few distinguished exceptions),

Latin American governments and institutions must
forgo the usual practice of "band aid" solutions
(dependence on international specialists or
nominal participation in grandiose, and often
ineffective, international projects in the hope that
somehow their people will be trained for them).
Indeed, the most debilitating characteristic of most
developing countries is not the per capita income,
or their comparatively low gross national product,
but their almost complete dependence upon
external sources (imported science and technology)
to solve their problems.

Thus Latin American countries must make a
determined effort to develop their indigenous
marine scientific competence, with critical
attention to quality rather than quantity. During
most international marine conferences, one hears
with tedious repetition the dire need for trained
scientists, technicians, and managers, as if these
were obstacles that could not be overcome, or that
should be mainly overcome by international
organizations. There is not only a lack of national
commitment to train people but there is also a
failure to use them properly once they are trained.
Indeed, there are many cases in which people that
have received advanced training find it extremely
difficult to do scientific work in their home
countries. As a result, they often choose to accept
slow, comfortable bureaucratization, or leave for
other countries where their scientific pursuits can
be fulfilled under less stifling conditions.

With a few exceptions, the university marine
biological programs in Latin Americaare inadequate
at the undergraduate level, usually requiring early
specialization in fisheries, where, for example,
cookbook population dynamics are taught. The
fundamentals of scientific work are often neglected
and the interdisciplinary nature of oceanography is

••*****»!•-

Institute Nacional de Investigacion y Desarrollo Pesquero,
Mar del Plata, Argentina.

46

disregarded. Indeed, the university is conceived of
only as a transmitter of knowledge, whatever its
value and time lag, and the university's organic
function, the generation of new knowledge
through scientific research, is overlooked. The
university programs — whether called
undergraduate, graduate, or professional — thus
produce an oversupply of deficient graduates that
are neither scientists nor marine or fisheries
biologists. Graduates, often talented people
disabled by their mediocre training, face several
alternatives: a) they fill government research
posts in fisheries departments or institutes; b) they
often take better paying jobs in the fishing industry
where little or no research is done; c) some remain
as instructors, where they propagate a second-class
education; d) others obtain support for advanced
training, eventually returning to the same
circumstances, but with most of the employment
opportunities already filled; and e) a few end up in a
field completely unrelated totheirfour orfive years
of education, representing a significant national
waste.

The institutes themselves are usually the
product of a politically motivated decision rather
than of a science policy program. Adequate funds
are initially approved to construct attractive
buildings that are ceremoniously inaugurated. The
facilities generally are poorly equipped (although in
some cases they are overequipped — unused
instruments being shown to visitors with hazy
predictions as to their eventual use), and usually
extremely deficient in library materials. During
times of economic difficulties, support for scientific
activities is not forthcoming; budgets are
significantly reduced; in a fewyears, this produces a
scientifically isolated and demoralized group of
individuals, strugglingwith administrations to allow
them to accomplish the duties for which they were
hired. Under these conditions, the fine work that is
occasionally produced is quite meritorious.

There are significant latent intellectual
resources in Latin America that could be developed
if the right circumstances were provided. Scientists
would produce prodigiously if only given the
appropriate framework and environment. If, in
seeking to wisely manage their marine resources,
Latin American countries do not drastically modify
their approach to scientific education, institutional
development, and the well-beingof theirscientists,
the factors that are negatively affecting the
development of science in general, and the marine
sciences in particular, will be perpetuated. This is a
most somber possibility. Oceanographic
institutions, advanced academic centers, and
international organizations can contribute to bring
about the necessary changes, but the central
decisions to alter the status quo must come from
within the countries themselves.

The Organization of American States is

Departamento de Biologia Marina y Oceanografia,
Universidad de Concepcion, Dichato, Chile.

currently planning a new program for marine
science and resource development in Latin
America. It has subdivided the region into four
areas with, presumably, similar characteristics and
problems. The areas are: Central America, the
Caribbean island countries, Atlantic South America,
and Pacific South America. The program focuses on
the coastal zone. Table 3 summarizes some of the
elements that are being considered. Six targets for
action have been identified: ecosystems research
and resource mapping; renewable and
nonrenewable resources management;
underutilized living and mineral resource
development, socioeconomic analysis of resource
development; recovery of degraded environments;
and identification of alternative marine resource
development possibilities. To work effectively, the
OAS program must operate at government level,
and of course, the implementation of any such
program is subject to national capabilities.

Finally, because of the high priority given in
Latin America to living marine resources, it is
appropriate to briefly review this sector. Data
collected by the Interamerican Development Bank
(IDB) indicate that in 1978 fishery production
amounted to 7.5 million tons, with an approximate
landingvalue of $2 billion. The IDB estimates that
some 2 million people are engaged in fishing
activities, the majority being coastal fisherman. The
theoretical supply is 22 kilos per person, but the
actual average human consumption is only 7 kilos —
that is, two thirds of the region's production is being
exported or processed for animal or industrial use.
These estimates do not include post-harvest losses
that result from lack of an infrastructure, estimated
to reach as high as 30 percent in some countries, or
that result from inappropriate technologies, for

47

example, by-catch losses of up to 70 percent by the
shrimp industry. The IDB further estimates that if it
is assumed that 25 percent of Latin America's
protein deficiency (about 20 million gross tons of
edible meats or 2 million tons of net animal protein
per year) could be satisfied with relatively
inexpensive fishery products, 5 million tons of fish
would have to be produced. The region's potential,
estimated by the FAO, is approximately 7 to 8
million tons per year, excluding krill. A 75 percent
increase over current production would be needed,
requiring an investment of about $3 billion. If the
present rate of human consumption were
maintained through 1990, with an 8.5 kilos per
capita demand, 2 million tons of additional fish
should be caught — that is, a doubling of the
present harvest for direct human use, and an added
investment of about $1.2 billion.

Considerable research will be needed to
continue and expand exploitation of living
resources. In the tropical countries, there has only
been progress in the last decade toward identifying
some of the mechanisms for the production of
organic material; in higher latitudes, the recent
discovery by Victor Gallardo of Chile of a
filamentous sulfide bacteria that may play a
previously unknown trophic role will probably
require a reevaluation of the mechanisms of
primary production. Throughout Latin America,
man's often conflicting demands will undoubtedly
require basic ecological information to provide
alternatives for coastal zone management. In areas
of large fishery harvests, the long-term fluctuations
of fish populations must be understood so that the
economic, social, technical, and industrial
components of the fisheries can be efficiently

Table 3. Summary assessment of elements of a marine resource development program for Latin America.

Resource

Mineral

Living

Non-Extractive

Assessment
Needs

Implications

Regional Diagnosis

Caribbean Sea

Central America

Petroleum

Natural gas

Manganese nodules

Sulphur

Fresh water

Construction materials

Magnesium

Other

Bromine, iodine

Phosphorite

Barite

Heavy metals

Inventories

Extraction technology

Geologists

Engineers

Support needs

Industrialization

Limited

Trinidad-oil

Bahamas-bauxite

Unassessed

Mexico more advanced

Human consumption
Industrial
Botanical
Aquaculture
Biochemical derivates

Ecosystem analysis

Bioproductivity

Stocks

Dynamics

Upwelling

Technology

Support Needs

Limited
Insular species
Diversification needs

Underexploited
Technological limitations

Energy
Recreation
Transportation
Communication
Receptacle for waste

Ocean waves/tides
Thermal gradients
Support activities
Ports/harbors/marinas
Industrial/urban/
agricultural discharges

Recreational high
Thermal gradients

Recreational
Waste receptacle

Atlantic S.A.

Pacific S.A.

Potentially good

Low exploration density

Potentially good

Important
Underexploited

Very important
Diversification needs
Industrialization

Energy, transportation
Waste receptacle,
critical focal areas

Energy/transportation
Waste receptacle

48

managed. Substantial investments based on
inadequate scientific understanding of the various
populations and their interactions with the
environment can lead to disastrous economic
problems, examples of which are already in
evidence.

Much remains to be learned. The only way in
which Latin American nations can successfully
incorporate the oceans into their development
process is to make a determined commitment of
substance rather than of form, generating the
necessary knowledge from their own bank of
human resources.

Francisco J. Palacio is Director of the Tinker Center for
Tropical Marine Coastal Studies in Latin America at the
University of Miami's Rosenstiel School of Marine and
Atmospheric Science.

Non-Resource Dimension

Ecological interrelationships

Economic impact

Analysis of alternatives

Social constraints and impact

Legal aspects, national and international

Institutional aspects

Education and training

Cultural characteristics

Aesthetic considerations

Coastal zone management information

Conflicting uses and pollution

Science policies

Education and technical training

Integral planning and coordination

Balanced development and economic planning

Policy guidelines for resource utilization

Research and institutional infrastructure

Public education and awareness

Scientifically very limited
Problems critical
Weak economics

Incipient awareness of problems
Serious pesticide problems in west coast
Caribbean coast neglected

Improving capabilities, some institutional
difficulties, poor coordination

Socio-political and management problems
Institutional difficulties

Suggested Readings

Allsopp, W. H. L. and F. J. Palacio. 1977. Reflections on interciencia

marine science symposium. Interciencia 2(5): 311-313.
Basalla, C. 1967. The spread of western science. Science 156:

611-622.
Bayer, F. M. 1969. A review of research and exploration in the

Caribbean Sea and adjacent waters. FAO Fish. Rept. No.

71.1.:41-88.
Cervigon, F. 1970. Las ciencias del mar en America Latina. Lagena

25/26: 39-43.
Maldonado-Koerdell, M. 1958. Panorama de los estidios

oceanograficos en algunos paises americanos. Bibl/ogr. Bull, of

Amer. Geography and Oceanography. Vol. 1, Pt. Oceanogr.,

pp. 177-314, Mexico City.
Moravcsik, M. 1975. Science development. The building of science

in less developed countries. Bloomington, Indiana: PASITAM

Publications.
National Adademy of Sciences. 1974. U.S. marine scientific

research assistance to foreign states. Washington :

Proceedings of MAS conference.
National Council on Marine Resources and Engineering

Development. 1968. Marine science activities of the nations of

Latin America. Washington, D.C.: USCPO.
Palacio, F. J. 1977. Towards a marine policy in Latin America.

Woods Hole Oceanographic Institution Technical Report

77-63.
Palacio, F. ]. 1979. Strengthening of marine capabilities in Central

America. Report to the Tinker Foundation, RSMAS, University

of Miami, Florida.
Roche, M. 1976. Early history of science in Spanish America.

Sc/ence194: 806-810.

49

Cilia and

by Sidney Tamm

v^ilia and flagella are threadlike, usually
microscopic projections of cells, specialized to
perform whiplike or undulatory motions. Found
throughout the animal kingdom, their beating
serves to propel small organisms, such as a
paramecium, through a liquid medium, or, when
the ciliated cells are anchored in a tissue, to move
fluids and particles over the surface of the
epithelium. The human body uses cilia extensively
for fluid transport in the respiratory and
reproductive systems; many human sensory cells
contain cilia or ciliary derivatives; and the human
sperm swims by means of its flagellum. In many
invertebrate animals, cilia and flagella perform an
even widervariety of functions, including
maintenance of water currents for feeding and gas
exchange, transport of gametes, food and excretory
products, various kinds of sensory reception, and
locomotion. Like muscle contraction, the activity of
cilia and flagella can be modified by the organism,
resulting in adaptive responses to environmental
stimuli.

The mechanism of ciliary and flagellar
motion, and the means by which their activity is
regulated, are therefore fundamental to our
understanding of various life processes, as well as
disturbances in these functions caused by disease
or genetic defects (such as certain forms of human
sterility). Cilia and flagella also are useful to study
fora more general reason. Their motile machinery is
made of an ordered array of microtubules.
Microtubules are rigid, hollow cylindrical structures
present in nearly all cells. They play an important
role in determining cell shape and form,
distributing the DMA of the chromosomes at cell
division, and aiding or directing the transport of
materials within cells and cell processes, such as
nerve axons. Cilia and flagella provide a model
system for investigating the role of microtubules in
these fundamental forms of cell movement.

Ctenophores, or comb jellies, are among the
most beautiful and voracious of the marine
plankton. They consume enormous quantities of
copepods, fish larvae, and other planktonic forms,

Figure 7. Diversity of form and behavior in Ctenophores.
Small sea-gooseberries, or Pleurobrachia, upper left, catch
copepods with their long tentacles; they are swallowed
whole by large Beroe, upper right. The belt-like Venus'
girdle, or Cestum, lower left, reaches a length of 7 meter,
and can swim like an eel. Leucothea, a lobate ctenophore
with oral lobes, lower right, has muscular fingers that dart
out to paralyze passing plankton.

and thus are an important part of oceanic food
chains. Ctenophores come in many shapes and
sizes (Figure 1), but almost all propel themselves
through the water by eight rows of ciliary paddles,
called comb plates. Each comb plate, or ctene,
consists of a transverse band of thousands of long
cilia, up to 2 millimeters in length, which beat
together as a unit. Ctenophores are the largest
known animals that use cilia for locomotion, and
the longest cilia are found in the comb plates of
Ctenophores. Because the microscopic size of most
other cilia makes them hard to study, the bigger cilia
of Ctenophores are indeed better, and comb plates
are, according to C. A. Horridge, "one of those
examples, occasionally presented in the animal
kingdom, where a giant structure lends itself to the

50

Cte

horcs

analysis of general biophysical problems." Both
locomotory and sensory functions of cilia have been
exploited in many of the behavioral responses of
ctenophores, such as orientation to gravity, escape
reactions, and feeding behavior. Ctenophores also
are among the lowest animals with a nervous
system, and several ciliary responses are under
neural control.

Thus ctenophores provide many unique
advantages for studying mechanisms of ciliary
motion, sensory functions of cilia, and control and
coordination of ciliary activity. Conversely, the
study of cilia in ctenophores reveals much about the
behavior and physiology of this important group of
marineanimals.

Mechanism of Ciliary Coordination

Cilia beat with a rapid, straight-armed effective or
power stroke, which moves the water, followed by a
slower, curling recovery or return stroke (Figure 2).
Adjacent cilia characteristically beat one after
another in a regular sequence, resulting in
metachronal (out-of-phase) waves of ciliary activity
that sweep across the ciliated surface. Since at any
one time some cilia are always performing a power
stroke, metachronal coordination ensures a
uniform and continuous propulsive force.

Ctenophores are the classic material for
experimental investigations on the mechanism of
ciliary coordination, because their ciliary units, or
comb plates, are relatively large and widely spaced.
The comb rows run from the sense organ at the
aboral end of the body to the mouth at the opposite
(oral) end (Figures 1 and 2). The comb plates beat in
an orderly sequence, starting at the aboral ends of
the rows and proceeding orally (Figures 2 and 8).
Metachronal ciliary waves therefore travel down the
comb rowsinanaboral-oral direction. The effective
stroke of the plates is normally in the opposite
direction — toward the aboral pole. Consequently,
the animal swims mouth foremost.

Two opposing theories of ciliary
coordination have been proposed, based largely on
experiments with ctenophores. In 1890, theGerman
physiologist Max Verworn found that preventing
the movement of comb plates in fierce, a pink,
dirigible-shaped ctenophore (Figure 1), stopped
transmission of ciliary waves. Cuttingacrossacomb
row between two plates did not interrupt
coordination, however. Verworn proposed that the
plates are coordinated by mechanical interaction,
-that is, the movement of one plate stimulates the
next plate to beat by hydrodynamic drag forces
transmitted through the seawater.

Working at the Marine Biological Laboratory
in Woods Hole, Massachusetts, the Harvard
physiologist G. H. Parker showed in 1906 that
metachronal waves were propagated past
immobilized plates of Mnemiopsis, a ctenophore
possessing oral lobes (lobate). This led him to
propose a neuroid or epithelial conduction
mechanism, in which a nervelike impulse
conducted through the tissue itself triggers the
beating of successive plates. Subsequently, both
mechanical and neuroid theories have been used to
explain not only metachronism of comb plates, but
also ciliary coordination in other systems.

This controversy has recently been resolved.
More detailed microsurgical experiments on
ctenophores by myself and others have provided
direct evidence for the mechanical theory of
coordination (Figure 3). In fierce and Pleurobrachia
(Figure 1) (but not in Mnemiopsis and other lobate

51

Figure 2. Forward-swimming cydippid larva of
Pleurobrachia. Note the prominent statocyst (s) at the
aboral pole, and mouth (m) at the oral end of the larva.
Rows of ciliary comb plates (cp) are caught in profile by
electronic flash. In this photograph, the plates nearest the
aboral pole have already performed an effective stroke,
and are unrolling toward the mouth in a recovery stroke.
Plates closer to the oral end are "frozen " in late to middle
stages of the effective stroke. The two tentacles (t) are
partially retracted. Magnification 350x.

ctenophores), preventing the movement of comb
plates, amputating plates near the base so that just
the stubs beat, or increasing the distance between
plates, stops transmission of metachronal waves;
however, a narrow cut through the tissue between
plates does not interrupt wave propagation (Figure
3, Table 1). The comb plates are mechanosensitive,
and beat when touched. In fact, Michael Sleigh of
the University of Bristol in England was able to show
that the beat frequency and the angular velocity of
beating can be directly controlled by a mechanical
stimulator vibrating at set frequencies.

Therefore, the comb plates of fierce and
Pleurobrachia are triggered to beat by
hydrodynamic drag forces arising from the
movements of the plates themselves. This group of
ctenophores offers the best demonstration of
mechanical ciliary coordination in any system and,
as such, illustrates a fundamental property of the
ciliary motor. Mechanical triggering or phasing of
beating means that the motile machinery can be
activated and timed by mechanical forces. Current
explanations of bend propagation along cilia and
flagella postulate that force generation is controlled
locally by internal mechanical stress arising from
curvature changes. By demonstrating a built-in
mechanosensitivity of the motile machinery,

ctenophore comb plates provide strong evidence
for such a sensory-motor feedback mechanism in
cilia.

But what is the mechanism of ciliary
coordination in lobate ctenophores? The same
experiments on Pleurobrachia and fierce give
opposite results when done on lobates such as
Mnemiopsis, Bolinopsis, and Leucothea (Figure 3
and Table 1). This is the reason for the earlier
dispute: previous researchers had not recognized
this dichotomy, and had attempted to interpret the
experiments on all ctenophores by one and the
same mechanism. Parker's view that mechanical
interaction between the plates does not coordinate
them in lobates has been confirmed (Table 1). The
key to the problem lies in a narrow tract of short
cilia, the interplate ciliated groove, which runs
between successive plates in lobate ctenophores,
but not in other types. Microsurgical experiments
show that this ciliated pathway is responsible for
coordinating the activity of the comb plates (Figure
3 and Table 1). The question then arises: howare
the cilia of the groove coordinated? According to
recent findings, they are coordinated mechanically.
It is not yet clear how the ciliated groove stimulates
beating of a plate, but this may also be done by
mechanical interaction. If so, the neuroid or
epithelial conduction theory would suffer a
gelatinous death in the very animal that gave rise to
it. This is notto say that epithelial conduction does
not occur; George Mackie of the University of
Victoria and others have demonstrated its
importance in mediatingvarious effector responses
in other invertebrates and in vertebrate embryos.
Epithelial conduction does not seem to coordinate
the beating of cilia, however. In addition, recent
electrophysiological studies have shown that
electrical events do not coordinate ciliary activity in
single-celled organisms, such as Paramecium.

One may then ask, why do lobate
ctenophores use this second type of comb plate
coordination? The reason seems to be that the
interplate ciliated groove allows coordination
between plates that have become widely separated
as a result of damage to the comb rows. In the
laboratory, we have found that removal of a piece of
a comb row results in rapid closing of the wound,
and a dramatic increase in the distance between
comb plates nexttotheoriginal excision (Figure 3j).
In fierce, Pleurobrachia, and other species lacking
an interplate ciliated groove, coordination between
plates is not restored until new plates regenerate in
the gap, a process that takes more than a week. In
lobates, however, coordination reappears as soon
as the wound heals (one to two days), because the
stretched ciliated groove conducts waves across the
gap between the plates. Lobates are known to be
more fragile than other ctenophores, and are
commonly found with widely spaced plates,
indicating previous damage to the rows. Because it

52

Figure 3. Microsurgical experiments to test the mechanism of ciliary coordination. Dotted line represents the interplate
ciliated groove in the comb rows of lobates. (See Table 1) (From Tamm, 1973)

permits the rapid restoration of coordination across
such gaps, the interplate ciliated groove is probably
an adaptation to the more delicate gelatinous body
of this group of ctenophores.

Interestingly, the free-swimming cydippid
larvae of lobates resemble miniature Pleurobrachia,
and likewise do not possess an interplate ciliated
groove. Nevertheless, their comb plates are

coordinated. Thus the type of ciliary coordination
changes during development in this group of
ctenophores!

Mechanosensory Transduction by Motile Cilia

Like many other organisms, ctenophores can sense
the direction of gravity and use it to modify their
behavior. Ctenophores often assume a vertical

Table 1 . Results of ciliary coordination experiments on ctenophores with and without an Interplate Ciliated
Groove (ICG). (From Tamm, 1973)

Experiment

Without ICG
Shown in With ICG (Pleurobrachia,
Figure 3 (Lobata) Beroe)

Prevent movement of comb plate(s) by:
Pressing down one plate
Holding one plate upright
Pressing down several plates
Cut tissue between comb plates
Amputate comb plates
Increase the distance between plates
Prevent movement of ICG cilia
Cut the ICG
Prevent movement of ciliated groove cilia
Cut the ciliated groove

a +(-)*
b +
c +
d +

e^L
•

j +

8
h
i

+ Transmission of metachronal waves

- Interruption of metachronal waves

* When movement of the ICG cilia is also prevented

53

position with their aboral-oral axis parallel to the
direction of gravity. During calm weather they are
found at the surface in a resting or feeding position.
When disturbed, ctenophores swim away from the
surface, and in a container remain at the bottom,
mouth facing downward. If a ctenophore is tilted
from either position of vertical balance, it rights
itself by asymmetrical frequencies of beating on
uppermost and lowermost pairs of comb rows. If
the animal turns to swim up, the lowermost rows
beat faster than the uppermost ones; the converse
occurs during downward steering. The tendency to
swim up or down — that is, the sign of geotaxis — is
called a mood.

Gravity receptors, or statocysts, generally
consist of a movable mass, the statolith, in contact
with ciliated receptor cells. Mechanical stimulation
of the cilia by the weight of the statolith causes
membrane conductance changes in the receptor
cell. These generator or receptor potentials are
transmitted by sensory nerves, and lead to
appropriate behavioral responses. How mechanical
stimuli are transduced into electrical signals is a
major unsolved problem in sensory biology.
Furthermore, because the cilia of some
mechanoreceptors are actively motile, whereas
those of others are not, the role that ciliary motility
plays in mechanosensory transduction also is
unknown.

The ctenophore statocyst has several unique
advantages for our investigation of the
mechanoreceptor function of motile cilia. The
statocyst is located at the su rface of the aboral pole,
and is readily accessible for observation and
experimental manipulations (Figures 2 and 4). The
cilia themselves are arranged into four conspicuous
groups, called balancers, which are more than 50
micrometers long and support a round statolith
(Figure 4). The balancers are motile, and act as
pacemakers for the comb rows. When a balancer
beats, a single wave of beating is propagated by
mechanical interaction along the two ciliated
grooves connected to its base, and thence to the
pair of comb rows in that quadrant of the body.

At the turn of the century, several
researchers realized that geotaxis must somehow
depend upon the motile responses of the balancers
to deformation by the statolith. Recently these
responses have been directly observed, and their
effects on beat frequency of the comb rows
quantitated in my laboratory at the Marine
Biological Laboratory. By using a new device, the
cteno-tilter, we have been able to precisely vary the
orientation of the animal with respect to gravity
(FigureS). Films of the beat frequency of comb rows
versus tilt angle of the ctenophore demonstrate that
the geotactic response is caused by inhibitory and
excitatory effects on beat frequency. For example,
rotating a negatively geotactic animal from 180
degrees (all rows inactive) to a horizontal position

V

Figure4. Statocystofa Pleurobrachia/arva. Twoof the four
sickle-shaped balancers (b) that support the cellular
statolith (s) are visible. In adult statocysts, each balancer
consists of a group of 150 to 200 cilia. A transparent dome
of non motile cilia (d) encloses the statocyst, and prevents
the escape of newly released statolith cells from the
epithelial floor (e). Magnification l,300x.

excites activity of the lowermost rows; tiltingfromO
degrees (all rows active) to a horizontal position
inhibits the beating of the uppermost rows (Figure
5, right). The same changes in orientation have the
opposite effects in positively geotactic animals
(Figure 5, left). Whether a given position will excite
or inhibit beating thus depends on the mood. The
ctenophore statocyst is primarily a static receptor,
not an acceleration detector, since the frequency
responses at a given position do not change with
time.

The mechanical and motile responses of the
balancers themselves have been visualized in
cydippid larvae of ctenophores, which have a
prominent statocyst and are particularly suitablefor
high-resolution microscopy (Figures 2 and 4). By
using a micro-cydippid-tilter (Figure 6), we
observed that the weight of the statolith deforms
the distal part of the balancers, and increases or
decreases their rate of beating, depending on the
direction of deformation and the mood. In a
negatively geotactic animal, for example, when a
balancer is pulled away from the center of the
statocyst, its beat frequency increases; when it is
pulled toward the center, its rate of beating
decreases.

Mechanosensitivity of the balancers may also
be demonstrated by artificial mechanical stimuli.
Bending a balancer with a fine tungsten needle

54

270°-

270°-

180C

geotaxis

-90e

- geotaxis

Figures. Ceotactic responses of Pleurobrachia, as shown with thecteno-tilter. Thectenophoreis pinned to a transparent
disk that can be rotated in a vertical plane underwater. Relative beat frequencies of the comb rows, indicated by number
of waves, during positive and negative geotaxis are shown for the two vertical and horizontal positions.

operated by a micromanipulator increases or
decreases its beat frequency, depending on the
direction of deformation and the mood. This
change in frequency is transmitted to the
appropriate pair of comb rows, thus mimicking a
geotactic response.

Directional sensitivity is also typical of
mechanotransduction in vertebrate hair cells, such
as found in the lateral-line organs, vestibular
organs, and auditory organs. Instead of a motile
response, however, be ndingthe hair bundle in one
direction causes a depolarizing receptor potential,
whereas displacement in the opposite direction
results in a hyperpolarizing response. In addition,
all other known mechanoreceptors do not reverse
the sign of their directional sensitivity.

In hair cells and other statocysts that give
electrical signals as their primary response, the
receptor potentials are graded with amplitude of
the displacement stimulus. In the ctenophore
statocyst, where only motile responses are
functionally expressed, one may ask: is the beat
frequency of a balancer graded with the extent of its
deformation? The cteno-tilter shows that it is (Figure
7). Both increases and decreases in the beat
frequency of comb rows are approximately linear
functions of tilt angle up to 40 to 50 degrees
deviation from vertical. Thereafter, the maximum
frequency response is maintained to the horizontal
position.

As previously mentioned, the ability to
reverse the sign of geotaxis is a unique feature of
ctenophore mechanoreceptors. What is the
mechanism of mood? Reversal in the frequency

response of a balancer to the same mechanical
stimulus points to a change in the receptor
(balancer) cell itself. In this regard, electrical
stimulation of the whole statocyst causes rapid
beating of all balancers and comb rows. This
indicates that membrane conductance changes of
the balancer cells can control the motility of the
balancers. In addition, it has been known fora long
time that changes in mood can be caused by a
variety of stimuli, such as water turbulence, light
intensity, and hydrostatic pressure.

Taken together, these findings suggest that
reversal in directional sensitivity of the balancers is
caused by electrical changes in the balancer cells,
mediated by input from the nervous system. The
effects of anesthetics, which block nervous
responses, provide support for this view: without a
functional nervous system, all animals become
positively geotactic and do not change moods!
Finally, electron microscopy reveals nerves in the
statocyst that make numerous synaptic contacts
onto the balancer cells.

Geotaxis in ctenophores thus appears to
involve a novel combination of mechanosensory
transduction and motility: a mechanoelectrical
process, similar to that in other known
mechanoreceptors; and an electrical-motile step,
perhaps analogous to the recently discovered
relationship between beat frequency and
membrane potential in Paramecium. Nervous
stimulation of the balancer cells, via input from
various sensory receptors, apparently acts at the
second step to cause reversals in the sign of
geotaxis.

55

Figure 6. Deformations and motile responses of balancers
(b) induced by the gravitational load of the statolith (s). A
micro-cydippid-tilter was made by placing a microscope
slide of immobilized Pleurobrachia larvae on a vertical
rotating stage of a horizontally oriented microscope. The
responses were filmed at 200 frames per second to slow
down the movements. Tracings 0-2 show the effective
stroke of balancers at 5 millisecond intervals: 0 indicates
resting position, 2 represents the maximum excursion.
Top, ctenophore oriented in 0 degrees position (see also
FigureS). All four balancers beat at 9 hertz. Bottom left and
right, positive geotactic response at 90 degrees. The lower
balancer beats at 7 to 2 hertz (left); the upper balancer
beats at 13 hertz (right).

Control of Ciliary Beat Direction

Besides the regulation of beat frequency, as
manifested by the geotactic response, other
parameters of ciliary and flagellar activity can be
controlled by the organism. These modifications in
the beat pattern are transient responses to various
environmental stimuli, and result in adaptive
changes in behavior. Examples include changes in
shape and symmetry of flagellar bending waves,
during chemotactic turning of sperm; the ciliary
arrest response which interrupts water flow and
particle transport in filter feeders, such as mussels
andascidians; and changes in the direction of ciliary
effective stroke, as in the avoiding reaction of
Paramecium and ctenophores (Figure 8).

These diverse motor responses share a
common basis: all are mediated by a transient
increase in intracellular calcium concentration.
However, the means by which calcium modifies the
pattern of ciliary and flagellar motility is not
understood, nor are the specific ciliary structures
involved in the responses identified.

The problem of regulatingciliary and flagellar
activity is closely related to the question of how they
beat. Acilium orflagellum consists of a cylinder of
nine doublet microtubules surroundinga central

pair of single microtubules, the entire 9 + 2
structure being held together by radial and
circumferential linkages (Figure 9). A series of
paired projections, called arms, are attached along
one side of each outer doublet microtubule. The
arms consist of the enzyme dynein, which can split
adenosine triphosphate (ATP), the energy source
for ciliary and flagellar movement.

From the work of Ian and Barbara Gibbons of
the University of Hawaii's Kewalo Marine
Laboratory, and Peter Satir of the Albert Einstein
College of Medicine in the Bronx, New York, it is
clear that the force for ciliary motion is generated by
the relative sliding of doublet microtubules past
one another, driven by the ATP-powered dynein
arms which act as cyclic cross-bridges. Other
structures linkingthe peripheral doublets to the
central pair are thought to confine or restrain this
local sliding, thereby converting translational
movement into bending movement. Since all nine
pairsof dynein arms are arranged uniformly around
the periphery (Figure 9), and generate force in only
one direction, arms on opposite sides of the cilium
will act antagonistically with respect to bending.
Most investigators therefore believe that bending in
a given direction is caused by activation of the
dynein arms on only one side of the cilium, whereas
bending in the opposite direction is caused by
active sliding of doublet microtubules on the other
side.

How does this switching mechanism in cilia
work?Thatis, howdoes calcium selectively activate
sliding between different sets of doublet
microtubules around the cilium? An attractive
theory, based on studies showing a correlation
between orientation of the central pair and
direction of beating in protozoan cilia, is that
calcium causes a rotation of the central pair, which
acts as a distributor to signal sliding of doublet
microtubules in sequence.

A critical test of this theory has been carried
out recently with ctenophores. We discovered that
stimuli which increase the flow of calcium into cells
cause a 180-degree reversal in beat direction of the
comb plates of cydippid larvae (Figure 8). The larvae
swim backwards for a short time, as in an avoiding
reaction of Paramecium! Although ciliary reversal is
common in protozoa, this is the first direct
demonstration of it in multicellularanimals.

Ctenophores have distinct advantages for
analyzing the structural basis of beat direction. The
comb plate cilia have unique morphological
markers, not present in protozoan cilia, for
identifying sidedness of thecentral pairand specific
doublet microtubules (Figure 9). In addition, a
single comb plate contains thousands of long cilia
which beat synchronously, thereby facilitating
transmission electron microscopy.

To do this work, we "froze" ciliary activity on
forward and backward swimming larvae by a rapid

56

Beroe ( + geotaxis)

Pleurobrachia (-geotaxis)

90<

Tilt angle ( from vertical )

Figure 7. Quantitative analysis ofgeotactic responses using the cteno-tilter. Beat frequency and tilt angle were
determined from films.

a

Figured. Pleurobrachia larvae swimming forward in seawater (top row, drawn at 60 millisecond intervals from film at 200
frames per second), and backward in seawater containing a high concentration of potassium chloride (bottom row,
drawn at 8 millisecond intervals from film at 400 frames per second). During forward swimming (a-f), the effective stroke
of the comb plates is directed aborally (arrows in c), and the plates unroll toward the mouth in the recovery stroke. During
backward swimming (g-l), the beat is reversed 180 degrees toward the mouth (arrows in j, k). Successive plates are
triggered to beat in an aboral-oral sequence during normal swimming, top row, and in the reverse order during backward
locomotion, bottom row.

57

Figure 9. Transverse section through comb plate cilia of
ctenophore larvae. In addition to the 9 + 2 arrangement of
microtubules (shown in cross-section), comb plate cilia
have compartmenting lamellae (cl) that connect doublet
microtubules No. 3 and No. 8 of adjacent cilia into rows
running normal to the plane of bending. Also, a dense
midfilament (mf) is located on only one side of the central
pair. During forward swimming, the effective stroke (ES) is
directed aborally, toward doublets Nos. 5 and 6 and away
from the midfilament. During backward locomotion, the
effective stroke is directed orally, toward doublet No. 7
and the midfilament. Thus neither the central pair nor the
peripheral doublets rotate when beat direction reverses.

chemical fixation technique. The orientation of the
ciliary markers with respect to the direction of
beating was determined by electron microscopy of
transverse thin sections through the comb plates. If
theorientation of thecentral pair really controlsthe
direction of bending, then during a 180-degree
ciliary reversal, it should also rotate 180 degrees. We
found, however, that reversal of beat direction is

not accompanied by 180-degree rotation of the
central pair or of the nine doublet microtubules
(Figure 9).

This clear-cut result shows that the
orientation of the central pair does not control the
direction of ciliary bending — that is, which
doublets actively slide and which do not. Ca* ' may
therefore act directly on the sliding process itself by
differentially activating dynein ATPase activity
around the cilium. Although the central pair may
still determine thep/ane of bending, it now seems
likely that its orientation is only a passive
consequence'of the bending.

We also discovered a second type of ciliary
reversal response specific to the feeding behavior
of adult Pleurobrachia (Figure 10). When a
Pleurobrachia catches prey (such as copepods) with
one of its long tentacles, the tentacle immediately
contracts. The two comb rows on either side of the
food-carrying tentacle then beat in the reversed
direction at high frequency , sweeping it toward the
mouth. The other six comb rows continue to beat in
the normal direction; as a result, the ctenophore
spins toward the food-capturing tentacle, further
wrapping the prey into the mouth. Reversed
beating only lasts a short time, after which the
animal resumes forward swimming.

Thus when copepods are fed to pinned
Pleurobrachia, which are unable to rotate, the
unilateral ciliary reversal response can be analyzed
by high-speed cinemicrography, microsurgery, and
electrophysiology. Together with the anatomical
studies by Mari-Luz Hernandez-Nicaise of the
Universite Claude Bernard, Villeurbanne, France,

Figure 70. Unilateral ciliary reversal during the feeding response of
adult Pleurobrachia. The direction of effective stroke (ES) reverses in
the two comb rows next to the contracted tentacle that bears the
copepod. Since the other six rows (only three shown here without
plates drawn) beat in the normal direction, the animal spins in the
direction shown (large arrow), further wrapping the prey into the
mouth, which also bends toward it.

58

the results so far indicate that unilateral ciliary
reversal is mediated by a local nervous pathway that
leads from either tentacle to the two adjacent
subtentacular comb rows.

Although inhibitory and excitatory responses
of cilia have been shown to be under nervous
control in various animals, including ctenophores,
this would be the first clear example of ciliary
reversal controlled by nerves (whether or not the
nervous system plays a role in triggering the global
ciliary reversal response in ctenophore larvae is not
yet known).

Thus ctenophores have told us a great deal
about how cilia work. We also have seen some of
the marvelous behaviors that cilia can bestow upon
these animals. In the words of Dr. Sebastian Beroe
of the Stazione Zoologica in Naples, "cilia and
ctenophores, like love and marriage, go together
like a horse and carriage."

Sidney Tamm is Associate Professor of Biology in the
Boston University Marine Program at the Marine Biological
Laboratory, Woods Hole, Massachusetts.

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Back Issues

Limited quantities of back issues are available at $4. 00 each; a 25 percent discount is
offered on orders of five or more. PLEASE select alternatives for those i n very limited
supply. We accept only prepaid orders. Checks should be made payable to Woods Hole
Oceanographic Institution; checks accompanying foreign orders must be payable in U.S.
currency and drawn on a U.S. bank. Address orders to: Oceanus Back Issues,
1172 Commonwealth Ave., Boston, MA 02134.

DEEP-SEA PHOTOGRAPHY, Vol. 18:3, Spring 1975 — Very limited supply.

ESTUARIES, Vol. 19:5, Fall 1976— Of great societal importance, estuaries are complex environments
increasingly subject to stress. The issue deals with their hydrodynamics, nutrient flows, and pollution
patterns, as well as plant and animal life — and the constitutional issues posed by estuarine management.

SOUND IN THE SEA, Vol. 20:2, Spring 1977 — Beginning with a chronicle of man's use of ocean acoustics, this
issue covers the use of acoustics in navigation, probing the ocean, penetrating the bottom, studying the
behavior of whales, and in marine fisheries. In addition, there is an article on the military uses of acoustics in
the era of nuclear submarines.

GENERAL ISSUE, Vol. 20:3, Summer 1977 — The controversial 200-mile limit constitutes a mini-theme in this
issue, including its effect on U.S. fisheries, management plans within regional councils, and the complex
boundary disputes between the U.S. and Canada. Other articles deal with the electric and magnetic sense of
sharks, the effects of tritium on ocean dynamics, nitrogen fixation in salt marshes, and the discovery during a
recent Galapagos Rift expedition of marine animal colonies existing on what was thought to be a barren ocean
floor.

THE DEEP SEA, Vol. 21 : 1 , Winter 1978 — Over the last decade, scientists have become increasingly interested
in the deep waters and sediments of the abyss. Articles in this issue discuss manganese nodules, the rain of
particles from surface waters, sediment transport, population dynamics, mixing of sediments by organisms,
deep-sea microbiology — and the possible threat to freedom of this kind of research posed by international
negotiations.

MARINE MAMMALS, Vol. 21:2, Spring 1978 — Attitudes toward marine mammals are changing worldwide.
This phenomenon is appraised in the issue along with articles on the bowhead whale, the sea otter's
interaction with man, behavioral aspects of the tuna/porpoise problem, strandings,a radio tag for big
whales, and strategies for protecting habitats.

GENERAL ISSUE, Vol. 21 :3, Summer 1978 — The lead article looks at the future of
deep-ocean drilling, which is at a critical juncture in its development. Another piece —
heavily illustrated with sharp, clear micrographs — describes the role of the scanning
electron microscope in marine science. Rounding out the issue are articles on helium
isotopes, seagrasses, red tide and paralytic shellfish poisoning, and the green sea
turtle of the Cayman Islands.

OCEANS AND CLIMATE, Vol. 21 :4, Fall 1978 — This issue examines how the oceans
interact with the atmosphere to affect our climate. Articles deal with the numerous
problems involved in climate research, the El Nino phenomenon, past ice ages, how
the ocean heat balance is determined, and the roles of carbon dioxide, ocean
temperatures, and sea ice.

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HARVESTING THE SEA, Vol. 22:1 , Spring 1979— Although there will be two billion more mouths to feed in the
year 2000, it is doubtful that the global fish harvest will increase much beyond present yields. Nevertheless,
third world countries are looking to more accessible vessel and fishery technology to meet their protein
needs. These topics and others — the effects of the new law of the sea regime, postharvest fish losses,
long-range fisheries, and krill harvesting — are discussed in this issue. Also included are articles on
aquaculture in China, the dangers of introducing exotic species into aquatic ecosystems, and cultural
deterrents to eating fish.

GENERAL ISSUE, Vol. 22:2, Summer 1979 — This issue features a report by a group of eminent marine
biologists on their recent deep-sea discoveries of hitherto unknown forms of life in the Galapagos Rift area.
Another article discusses how scuba diving is revolutionizing the world of plankton biology. Also included are
pieces on fish schooling, coastal mixing processes, chlorine in the marine environment, drugs from the sea,
and Mexico's shrimp industry.

OCEAN/CONTINENT BOUNDARIES, Vol. 22:3, Fall 1979— Continental margins are no longer being studied for
plate tectonics data alone, but are being analyzed in terms of oil and gas prospects. Articles deal with present
hydrocarbon assessments, ancient sea-level changes that bear on petroleum formations, and a close-up of the
geology of the North Atlantic, a currentfrontierof hydrocarbon exploration. Othertopics include ophiolites,
subduction zones, earthquakes, and the formation of a new ocean, the Red Sea.

OCEAN ENERGY, Vol. 22:4, Winter 1979/80 — How much new energy can the oceans supply as conventional
resources diminish? The authors in this issue say a great deal, but that most options — thermal and salinity
gradients, currents, wi nd, waves, biomass, and tides — are long-term prospects with important social
ramifications.

A DECADE OF BIG OCEAN SCIENCE, Vol 23 : 1 , Spring 1980 — As it has i n other major branches of research, big
science has become a powerful force in oceanography. The I nternational Decade of Ocean Exploration is the
case study. Eight articles examine scientific advances, management problems, political negotiations, and the
attitudes of oceanographers toward the team approach.

OUT OF PRINT

SEA-FLOOR SPREADING, Vol. 17:3, Winter 1974

AIR-SEA INTERACTION, Vol. 17:4, Spring 1974

ENERGY AND THE SEA, Vol. 17:5, Summer 1974

MARINE POLLUTION, Vol. 18:1, Fall, 1974

FOOD FROM THE SEA, Vol. 18:2, Winter 1975

THE SOUTHERN OCEAN, Vol. 18:4, Summer 1975

SEAWARD EXPANSION, Vol. 19:1, Fall 1975

MARINE BIOME DICINE, Vol. 19:2, Winter 1976

OCEAN EDDIES, Vol. 19:3, Spring 1976

GENERAL ISSUE, Vol. 19:4, Summer 1976

HIGH-LEVEL NUCLEAR WASTES IN THE SEABED? Vol. 20: 1 , Winter 1977

OIL IN COASTAL WATERS, Vol. 20:4, Fall 1977

Oceanus

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