32 Doty: Mcasurement of water movement
Botanica Marina
Vol. XIV, p. 32—35, 1971
Measurement of Water Movement in Reference
to Benthic Algal Growth*
MAXWELL S. DOTY
Botany Department^ Umvernty of Hawaii; Honolulu^ Hawaii
(Received: 30.9.1970)
Introduction
In the sea it appears that fertility for phytoplankton,
microphytes and macrophytes is rather mutually ex-
clusive. That is to say, a place ideally fertile for one of
these forms is not ideal for the others. RUTTNER (1926)
and FELDMANN (1938), among others, have observed
that benthic populations are reduced äs calm or stagnant
water is reached. On the other hand, CHANDLER feit
(1937) the opposite to be true in respect to phyto-
plankton. WHITFORD (e. g.,1960) has demonstrated ex-
perimentally the tenability of RUTTNER'S hypothesis rela-
tive to lotic and lentic stream plants. RUTTNER's hypo-
thesis was that water movement breaks down boundary
layers of depleted water near the plants, thus increasing
diffusion which in turn results in accelerated growth.
Lack of a method of m e a s u r i n g the effect of water
movement on diffusion has hampered advance in under-
standing the phenomena involved. Thus, little has been
added since to what RUTTNER said, It must be made
clear that this is a process quite different from the mass
action effects associated with stream velocity (GESSNER,
e.g* 5 1950) 5 waveforce(JoNEsandD£METROPOULOS,1968) }
wave mixing influence on temperature (GESSNER and
HAMMER, 1967) or wave-related water surge such äs
NORTH et al. (1969) have reported äs distributional
factors- It is also different from the antagonism between
planktonic and benthic forms which FITZGERALD (1969)
described äs being due primarily to ferdlizer availability
and antibiotic activity.
The mechanism with which we seem to be concerned
here is diffusion; diffusion of materials into a benthic
alga or other organism and the diffusion of metabolic
products outward. MCCONNELL and SIGLER (1959)
actually devised a method which provides integrated
measurements of the diffusion enhancement by current
velocities, turbulence and other factors. For this they
used the weight loss from 2,25 g sodmm chloride tablets
during 1-minute immersions. Just recently the author's
attention has been drawn to the use ot calcium Sulfate
balls on welding rod supports (Muus, 1968) for measur-
ing water movement over a period of a few hours.
Muus did not mention diffusion and, like McCoNNE LL
and SIGLER, related his results to average current
velocities, While effective in non-tidal waters, these
*) This work was supported by research contract AT (04—3)—235
of the U. S. Atomic Energy Commission.
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methods are not suitable, e. g., on reef flats over which
the movement of water varies greatly during the 24.84-
hour tidal or lunar day,
Materials and Methods
The goals of the present work have thus become finding,
testing and putting to use a practical method for measur-
ing the degree to which diffusion is enhanced by water
character and tneasurement This method must fulfill the
conventions o£ minimal cost, mechanical strength, opera-
tional simplicity and usefulness under reef-flat circum-
stances remote from civilization. Following the 5th
International Marine Biological Symposium, where the
problem was discussed informally, over 40 different
techniques or substances were investigated.
The idea of using dissolution of plaster of Paris clods
cemented onto plastic cards was finally adopted. For the
past several years the "clod-cards" resulting have been
used in a wide variety of circumstances across the
Tropical Pacific. To promote further refinement and
use of this technique, the production and an example
of the use of these clod-cards is presented here,
For Held use a hundred or more 30 g clod-cards (Fig, 1)
are made at one titne, The plaster of Paris is dried at
over 60°C for 48 hours before use, mixed with water
and cast in oblong round-bottomed polyethylene ice-
cube trays. With different lots of calcium sulfate, different
proportions of water are used> but this is usually about
45 ml water for each 50 g of calcium sulfate.
At different times clod-cards of other weights have been
used, but it has been found that the larger sizes are ne-
cessary for the calcium sulfatelosstobeacceptably linear
with the time and the water motion conditions found
on reef flats, Other shapes have been less satisfactory,
Since in-use weight loss is found to be sensitive to the
initial size of the clod, the flat bases of the clods are
filed so that each clod within a lot is within less than
0.4 g o£ a selected weight. The nearly-dry weight-
adjusted clods are then cemented to white matte-surfaced
thin plastic cards (Fig. 1) using either epoxy or contact
cemerit. After a 24-hour period for the cement to set,
they are dry weighted, The dry weight is written on an
uppet corner of the card; this number may also serve
äs an identification number.
Throughout, methods suitable to field conditions on
tropical reefs have been employed, For example, the
Botanica Marina / Vol. XIV /1971 / Fase, l
 Doty: Mcasurcment of watcr movement
Fig. l
A clod-card ( I c f t ) ready for use and two (right) after such use that the respective dissolution of about 13.5 and 16.7
material has occurred
clod-cards are dried in the open air and, thus, at tem-
pcratures oftcn between 22 and 29° C Stahle dry weights
are reached more quickly in such air than in a 50-degree
oven where lower weights are obtained, Within two
\veeks after cooling oven-dried clod-cards gain weight
such that they statistically weigh the same äs those dried
in the open air.
Calibration of each lot of clod-cards is necessary to
enable comparison of the results from the use of
difierent clod-card lots- A "number ten" tin can or
cylindrical Container, 15 cm in diarneter and over 11.5cm
high, is obtained or improvised. Five clod-cards are
placed symmetrically on the bottom and 2 liters of the
water in which the in situ measurements are to be made
is added. The water is kept near the in situ temperatures
and changed every day at the same time for at least
four days. This time span is used since a little higher
dissolution rate is obtained at first; that is, the first day
more weight is lost than is lost on each successive day.
On the last day the clod-cards are removed, rinsed and
their dry weight obtained äs before,
The basic calibration value is calculated by the equation:
rn c (tc/tc) = k, in which m c is the mass lost for the
calibration period, tc, and t e is the field exposure period,
The calibration value, k, usually ranges, for different
clod-card lots intended for one 24-hour day exposure,
between 0.25 and 1.00 g per day. It is a measure of the
solution, dissolution and diffusion rates of the calcium
sulfate from the clods under the particular conditions
used. It is the value for still water conditions.
A nurnber of potential sources of the random Variation
in the clod-card calibration results äs well äs in the
results from usage have been found to be relatively
insignificant, A Variation of 10 per cent from the selected
amount of water added to a given amount of calcium
sulfate in making the clods can produce äs much äs an
0.05 g/day difference in their calibration value. A salinity
Variation of plus or minus two parts per thousand may
provide a 0.01 g/day difference. A temperature Variation
of plus or minus 2°C may provide a 0,03 g/day difference.
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33
uf tht» c a l d u m
In the light of the variability between the dissolutiun
rates obtaining for different clod-cards from the same
lot in the same can of water, it appears the temperature
and salinity differences between the calibration circum-
stances and nature are not signiticanu
In field use, to integrate the diffusion enhancement over
the tidal cycle period, the clod-cards are put out at the
most favorable time of day in respect to \vorking
conditions and taken in the next day at the same time,
This exposes the clod-cards to the changes related to
diurnal and tidal factors.
To hold them in place, the clod-cards are taped onto
bricks to obtain measurements on rocky benches or
coral reef flats. Ordinary masking tape is satisfactory if
applied dry. The tape does not remove, but protects,
tbe pencil marks on the cards. The clod-cards can bc
thumb-tacked, nailed or Stapled to stakes driven in mud
flats or taped or held by rubber bands to pegs driven
into cracks between rocks.
After exposure the clod-cards are rinsed in fresh water
and dry weighted once again. The second dry weight
value is recorded under the first äs are the successive
weighings made to check the completeness of drying.
When used in a series across the reef at Waikiki in
Hawaii, the clod-cards lose 1,5 to 10 g in 24 hours.
Elsewhere losses have been obtained of over 22 g from
clod-cards of 32 g initial weight> and the two used clod-
cards in Figure l have lost, respectively, 13.5 and 16.7 g
from their initial weights. Weighing can be done on a
simple trip-balance graduated to tenths of a gram.
Such magnitudes and ranges of loss obviate any necessity
for more refined weighing, and variations in individual
clod-card behavior are reduced in practical significance,
The weight lost during an exposure period, a "24-hour
tidal cycle" in our cases, has been treated two ways.
First, it can be used äs CV, a clod-card index value,
The second way uti^es the still-water calibration value
by dividing it into the weight lost by each clod-card
during its period of use in the field so a diffusion index
factor (= DF) is obtained, This DF is accepted äs a
34 Doty: Measufement o£ water movement

Tab. l
Clod-card measurement of diffusion enhancement compared to wet and dry weight of frondose algae sampled in 12 successive 2Q-meter zones
between the shore and reef margin at Waikikl in Honolulu, Hawaii, in December, 1967. The clod-card values are mean loss of weight for clod-card
pairs in terms of grams of calcium sulfate lost over a 24-hour exposure period. For the lOdegrees of frecdom the 5% confJdence levelfor correlation
is 0,576; for 1%, 0.708

Dependent Statistical L Clod-card weight loss for 24-hours exposure on the dates given Mean clod-card
variable äs measure weight loss all
algal sample 21—22 22—23 28—29 29—30 periods

Wet weight b 0.87 1,23 —0.27 0.75 0.60

r 0.70*) 0.68 0.31 0.56 0.62
Dry weiglit b 2.86 1.29 —1.50 1.76 0.93
r 0.51 0.16 0,39 0.29 0,21
*) This value was 0.70322 before rounding.

measure of the degree to which diffusion or dissolution
was enhanced by water movement where the clod-card
was exposed.
As an example of practical usage in the field, clod-card
weight losses (CV's) were measured across the middle of
a study area in Honolulu, Hawaii. This was done
coincident with an experimental harvesting of the algal
Standing crops which provided a mass per square meter
distribution of the seaweeds across the reef in a parallel
fashion. During each of the four days harvesting was
done, clod-card measurements were made.
A high correlation ("r" in Table 1) of CV with Standing
crop size usually obtains in the resulting data» A correla-
tion coefficieni of 0.70, significant to about the l per cent
level, is found among these results. Likewise the
regression slope ( C V in Table 1) is generally consistent
but for one day when the correlation of Standing crop
with wave height is low.
Discussion
One of the major remaining unresolved problems is
variability in weight loss under seemingly uniform
conditions by clod-cards from the same lot, Muus had
problems of this sort, too. Even under our most careful
calibration conditions wild values appear embarrassingly
often« This may mean solid pieces were lost from the
clod-card (greater losses) or that there were bubbles in
the clods (smaller losses). In nature animals sometimes
nibble the Blocks, but this is usually obvious. It is feit
that significant abrasion is detectable under the present
circumstances of use. Muus feit the larger weight loss
from a pair was the more reliable measurement. From
time course plots of the results from clod-card pairs
used in calibration experiments, it appears that the lower
of the two values (i. e.? least weight loss) obtained from
each pair is most often the more reliable, The practical
solution to this has been using the clod-cards in pairs
or in close linear series.
A major technical problem found in applying this method
involves replicating results from successive tidal cycles
and the risks in using results from just one tidal cycle.
If results from one CV measurement period are used,
by chance one can get (Table l, data for Dec. 28—29
in comparison to the rest) a negative regression on one
day whereas the data for the other days and the mean
values of all the times studied show reliable positive
regressions. These particular four CV measurement
periods were during a light weather period and, though
no record was made at the time, it is clear from the wind
record for the Honolulu Airport, about seven rniles
away, that the wind-tide combination on that one day
was different from that on the other three days.
Antecedent events causative o£ total Standing crop size
(i, e., g/m2 for the whole reef area) in such situations
seem to have been (DorY, in press) some few weeks
prior, e. g., three weeks prior in the case of mean wave
height äs judged from the correlations and negative
regressions obtaining between recorded wave height
and biomass. However, from the data in Table 15 it is
rather clearly indicated that the hypothetical relationship
of faster growth where the water is more turbulent i s
sound. Thus, when measured after a period of crop
reduction by storrns, seasonal events or harvesting, the
Standing crop must grow more quickly where the water
moves the more rapidly. That the correlations in Table l
are higher with wet weight, which might be thought
of äs dominated by more recent and more succulent
growth, rather than with the dry weight can be postulated
äs further evidence that growth rate and young Standing
ctop size are related to water motion.
Use of clod-cards appears to be a practical way of
quantitatively comparing habitats in respect to the effects
of water motion. As long äs one lot of clod-cards is
used, CV (clod-card Index value) is suitäble. McCoNNELL
& SIGLER and Muus used such an index value. However,
since many veiocities are involved, since calibration with
respect to velocity is not practical in the field and since
it is diffusion that is considered important, the DF, the
diffusion factor, is to be preferred over terms thought
to relate to velocity. Also, DF is to be preferred over
CV for, by using DF, comparisons can be made in-
volving use of different clod-card lots at the same place
or äs prepared and used by different people in different
parts of the world. Likewise, it appears that DF is an
integrated measure of the degree to which water
characteristics and water movements of all kinds enhance
diffusion of materials into and out of benthic organisms.
Acknowledgements
The author is appreciative of the considerable technical
assistance provided in the development of this method
and in the carrying out of the numerous laboratory and
neld experiments. In this way, in particular, DR, GERTRU-
DES AGUILAR-SANTOS and MSSRS. MARK M. LITTLER,
GERALD T. KRAFT and CHARLES R. HASTINGS were
most helpfuL

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 ! Meäiurement of watet movement
Referencee
CUANDLER, D. C 1937- Fäte of typical lake pknkton in strcams.
Ecological Monographs 7 (4): 447—479. — Dotr, M, S. (Inprese),
Antecedent event influence ofl benthic algal standing ctops in
Hawaii. Journal of Experiment»! Marine Biology and Ecology. —
FELDMANNT, J. 1938, Rccherches sui la vigitation marine de la
M6ditcrran6e . . . Revue Algologique 10: l—339, (Apparently
issued separately in 1937 äs a thetis and possibly, äs citcd here,
did not appear until 1940.) — FITXGERALD, G. P. 1969, Some
factors in the competition or antagoniam among bacteria, algae,
and aquatic weeds, J. PhycoL 5i 351—359. — GESSNER, R 1950,
Die ökologische Bedeutung der Strömungsgeschwindigkeit
fließender Gewässer und ihre Messung auf kleinstem Raum. Archiv
für Hydrobiologie 43: 159—165. — GESSNER, K and L, HAMMER.
1967, Die litotale Algenvegetation an den Küsten von Ost-Vene-
zuela. Int. Revue Ges. HydrobioL 52 (5): 657—692, — JONES,W, E.
and A. DEMETROPOULOS, 1968, Exposure to wave action: measure-
ments of an important ecological parameter on rocky shores on
Botanica Marina/ . XIV/1971 /Fstoe. l
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35
Anglesey. Journal of Experimental Marine Biology andEcology2:
46—63. — MCCONNELL, W. J., and W. F. SIGLER, 1959, Chloro-
phyll and productivity in a mountain river. Limnology and Oceano-
graphy 4: 335—35L — Muus, B. J. 1968, A field method for
measuring "exposure** by means of plaster balls. A preliminary
account. Sarsia 34: 61—68, ~ NORTH, W. J,, C T. MITCHELL and
L. G. JONES, 1969, Mass cultures of Afacrotysiis, Chapter 5 in
Wheeler J. North, Kelp Habitat Improvement Project Annual
Report l July 1968—30 June 1969- Frontmatter plus 142 pages.
(Duplicatedanddistributedby W, M. KeckLaboratoryof Environ-
mental Health Engineering, California Institute of Technology,
Pasadena, California.) — RUTTNER, F. 1926, Bemerkungen über
den Sauerstoffgehalt der Gewässer und dessen Respiratorischen
Wert. Naturwissenschaften 14: 1237—1239, — WHITFORD, L, A.
1960, The current effect and growth of fresh-water algae. Trans-
acüons of the American Microscopical Society 79: 302—309,