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The Island Mass Effect


Maxwell S. Doty and Mikihiko Oguri,

Department of Botany, University of Hawaii, Honolulu

observers, beginning at least with GRAN (1912) and LOHMANN

(1912), have reported that as the continental margins of oceans are
approached the standing crop of plankton increases. GRAN (1912: 378)
estimated the standing crop to be about 100 times greater near shore than
it is off shore. These increases have been related by GRAN, LOHMANN, and
other authors variously to upwelling, to nutrient-rich run-off water, to the
nature of the variations (e. g., salinity) in the particular waters, and to
attenuation in a shore population carried off shore.
The same increases are presumed to exist with nearness to island shores,
though little attempt has been made heretofore to demonstrate this. For
various reasons some explanation for this phenomenon near small oceanic
islands seems necessary other than the above. Around islands, such as those
of the Hawaiian group, only a relatively small amount of vertical mixing
of the passing waters has been reported. Since the areas studied by MCGARY
(1955) showed little and were the areas where a significant amount of vertical
mixing was anticipated, we are led to say little is expected. While some
run-off water could be expected around high islands there would be little
or none around low islands. A small amount of percolation water could be
expected from any freshwater lens present in the latter case. There would
be expected some, though little, variation in salinity outside Kaneohe Bay
with distance off shore.
One explanation has been offered (DOTY, 1954: 6 ff.) in relation to atolls
which could be expected to apply to reef-surrounded high islands as well.
This hypothesis is, in brief, that the benthic algae, including both the
endozoic species and those which form calcareous reef margins, accumulate
inorganic nutrients from the passing waters, which are relatively poor in
nutrients. They would also concentrate these materials from leaching of
the freshwater lens and from whatever run-off water there might be. These
nutrients would become available to planktonic forms, through for instance,
benthotrophic herbivores.

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Figure 1. The four classes of positions, called stations, from which water samples
were obtained for the present study. The inset shows the area of the large figure in
relation to the whole island of Oahu, one of the Hawaiian Islands.

If there is an accumulation of material by such algal organisms, it follows

that as island masses are approached there may be either an increase in the
size of the standing crop of phytoplankton, or an increase in rate of anabolic
activity, i. e., higher productivity, or both. There seem to be no studies of
productivity and little information to show the quantitative nature of the
reputed or expected increase in standing crop as island shores are approached.
In order to test the hypothesis that there is an increase in productivity as
the shore of an oceanic island is approached, and to determine something
of its magnitude, water samples were obtained from a series of stations
(Figure 1) near the island of Oahu in the Hawaiian chain. These stations
were on the windward side of the island in Kaneohe Bay and along a course
approximately 60 from the south-eastern entrance to this bay. The samples
were obtained by pumping sea water with a cast-iron pump through a
plastic hose into a plastic bucket or by dipping water from the sea with a
plastic bucket. These stations were occupied during 19 cruises spaced over
a period of 14 months.
Salinity, inorganic phosphate-phosphorus, Secchi disk readings, and
temperature information were obtained from the various stations. Plankton

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Plankton Production

1 1


1 1 1 1 1 1

STATION 4 iN'iei


IJ 1 1 1





STATION 2 ( N - 1 6 )



1 1



1 1

1 1



Figure 2. Ability of the waters near the windward shores of Oahu to fix carbon
photosynthetically, plotted as a function of distance from shore.

tows, surface-water oxygen content and other data were gathered less
regularly as well. Various factors prevented our obtaining samples consistently
from below the surface during most of the 14-month study period.
A modification of the dark and light bottle carbon fourteen technique
employed by STEEMANN-NIELSEN (1952) was used to determine photosynthetic
carbon fixing potential in the samples. A known uniform amount of carbon
fourteen, as sodium carbonate, was added to each 276 millilitre pyrex
bottle of the sample water immediately after the water had been dipped or
pumped aboard. This amount was such that with our counting apparatus
the total carbon fourteen count per bottle was at least 2-18 X 106 counts
per minute and in most cases more. The inoculated bottles were exposed
to light from daylight fluorescent tubes, the intensity of which was about
1500 foot-candles in the live well of the University of Hawaii research
vessel, the "Salpa". Sea surface water was circulated through the live
well constantly, and it was determined that operationally no elevation in
temperature took place above that of the sea surface water. A usual incubation
time was 5 hours.
Detailed descriptions of this carbon fourteen technique and the raw data
will be distributed for an indefinite period of time in mimeographed form
upon request.
For the purposes of this paper, which is concerned with comparative
rather than absolute measurement, only the calculated amounts, in
milligrammes, of carbon photosynthetically fixed per cubic metre of surface
water per hour are presented in Figure 2. Since the results were obtained
under operationally uniform conditions, we believe that they reliably reflect

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the carbon fixing ability of the local water masses from which they were
drawn. The ranking of the results of the stations in relation to each other
was consistent on the different cruises. This further induces faith in the
significance of the results obtained.
If one omits the two highest values obtained among the seventeen otherwise
used in calculating the mean and standard deviation presented for Station 3
in Figure 2, one obtains the mean and standard deviation indicated by the
lighter line to the left of the longer vertical bar. The position of each of
these highfigures,the next lowestfigure,and the very lowestfiguresobtained
are indicated by triangles pointing to the left along and above the vertical
bars at Station 3. Between the two lowest left-pointing triangles there are
thus 15 figures (inclusive of the triangles themselves) and it is the values
of 13 of these that we have used as the basis for the lighter line to the left,
indicating their standard deviation; the right-pointing triangle indicates
their mean.
An examination of the differences between the means obtained indicates
a probability that they are reliable as plotted. To this end the T-values
between the means have been entered on the figure. The statistical similarity
between Stations 3 and 2 is interpreted as an indication that the water at
Station 3 is representative of an intermediate condition between reef water
and the hardly-reef-influenced but near-shore water of Station 2. That even
omission of the two highest figures from Station 3 makes little difference in
the picture enhances our confidence that there is a steady elevation in
productivity as Station 3 is approached from off shore.
From other evidence it appears that the two high figures (in Fig. 2) may
have been due to unusually highly productive water masses which had come
out of Kaneohe Bay and still remained undispersed when they were sampled
at Station 3. These numbers were obtained shortly after the close of periods of considerable freshwater run-off into the bay. Although we have
been gathering information on the effects of run-off water, they are too
incomplete to take up here.
From the consistent increase in carbon fixation as shore is approached
we infer continued tenability of, and accept as a working theory, the
hypothesis that productivity is greater as one approaches an island shore.
From our results (Fig. 2) it can be seen that productivity increases at least
two orders of magnitude as shore is approached and one passes into Kaneohe
Bay, for in the shallow water of inshore stations the as yet unassayed
benthic productivity seems to do more than make up for the loss in the
depth of the productive column. We are not yet able to assess clearly the
effects of the bay. Data from a few cruises approaching Pearl Harbor, on
the leeward side of the island, give us results similar to those presented in
Figure 2. It is, of course, desirable to collect more of such data.
We have not yet been able to obtain what we feel we can accept as standingcrop measurements which would enable us to draw a reliable inference as
to the relationship between standing crop and productivity. From the
pigment samples, Secchi disk readings, and the plankton volumes obtained
at the same time as the productivity samples, it is clear, however, that there
is an increase in standing crop as shore is approached.

Plankton Production


This work was supported by the research grant AT(04-3)15 of the U.S.
Atomic Energy Commission and by Pacific Island Research Funds of the
University of Hawaii. The technical assistance of Messrs. Clarence SUZUKI,
Everet C. JONES, Leonard WOLFE, and Dr. Robert GUILLARD is gratefully
To test the hypothesis that as oceanic island shores are approached there
is an increase in phytoplankton productivity, and to determine its magnitude,
an experimental collection of data was obtained by means of the light and
dark bottle technique and carbon fourteen. Over the 14-month period of
sampling a consistent increase of nearly two orders of magnitude was
observed in carbonfixedper hour per cubic metre of surface water, as Oahu
(in the Hawaiian Islands) was approached from a position averaging 15 miles
off shore to windward. The particular experimental conditions yield
calculated raw data figures increasing regularly from 0-096 milligrammes of
carbon photosynthetically fixed per hour per cubic metre at the furthest
off shore station to 4-2 in Kaneohe Bay.
DOTY, M. S., 1954. "Floristic and ecological notes on Raroia". Atoll Res. Bull, 33,
pp. 1-41.
GRAN, H. H., 1912. "Pelagic plant life" in MURRAY and HJORT "Depths of the Ocean",
pp. 307-386.
LOHMANN, H., 1912. "Beitrage zur Charakterisierung des Tier- und Pflanzenlebens
in den von der 'Deutschland' wahrend ihrer Fahrt nach Buenos Ayres durchfahrenen Gebieten des Atlantischen Ozeans". Internat. Rev. d. ges. Hydrobiol.
u. Hydrogr., 5, 2, pp. 185-225; 4, pp. 344-372.
MCGARY, James, 1955. "Mid-Pacific Oceanography. Pt. VI. Hawaiian offshore waters,
December, 1949-November, 1951". U.S. Dept. Int., Fish & Wildl. Serv., Spec.
Sci. Rep. Fish., 152, 138 pp.
STEEMANN NIELSEN, E., 1952. "The use of radio-active carbon (C14) for measuring
organic production in the sea". Journ. du Cons., 18, 1, pp. 117-140.

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From inspection of Figure 2 it appears not unlikely that an asymptotic

value for carbon fixation would be obtained in the waters beyond Station 1
at least at some point along the next 100 miles offshore. Similarly it appears
that some finite value could be reached in the waters nearest the shore and
that perhaps this finite value would be within the next order of magnitude
above that of the fixation obtained at Station 4 in Kaneohe Bay. If this
highest inshore value can be added to that representing the productivity of
the inshore benthic algae one may have a value which, when related to the
energy falling per unit area, will be near a maximum for efficiency in our
marine conditions under climax circumstances.