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Aquacultural Engineering 6 (1987) 191-208

Predicting Night Time Dissolved Oxygen Loss in Prawn Ponds of Hawaii: Part I. Evaluation of Traditional Methods
Charles R M a d e n j i a n , G a r y L. R o g e r s and A r l o W. Fast Hawaii Institute of Marine Biology,Universityof Hawaii, PO Box 1346, Honolulu, Hawaii 96744, USA

ABSTRACT Several candidates (secchi disc depth, in vivo fluorescence, chlorophyll a concentration, turbidimeter measurement, light absorbance) for an indicator of water coh~mn respiration in prawn (Macrobrachium rosenbergii) ponds of Hawaff were evaluated. The % variation in the water column respiration rate explained by the indicator and water temperature did not exceed 60% for any of the potential indicators. Pond DO and water temperature were monitored overnight at prawn ponds in Kahuku (Oahu). Additionally, water column respiration rate in the ponds was estimated by the dark bottle method. On average, water column (or plankton) respiration comprised 42% of the whole pond respiration. Sediment respiration accounted for 50'2% of the whole pond respiration. The log-linear (or exponential) extrapolation technique yielded substantially more accurate estimates of dawn DO in the Amorient ponds than the linear extrapolation routine.

INTRODUCTION Dissolved oxygen concentration (DO) is one of the most important factors affecting most aquaculture species. When dissolved oxygen levels in aquaculture ponds become low, the culture d organisms may become stressed or even die. During the diurnal cycle, D O is typically lowest at dawn. After sunrise. DO increases due to photosynthesis; but at night, biotic respiration and chemical oxidations result in a net loss of oxygen which can reach critically low concentrations. The farmer can add oxygen to the pond by several methods including: ( 1 ) flushing water into the pond, whereby water is drained from the pond as water with a higher DO is added; and (2) aeration of the pond water 191 ,4quacultural Engineerin~ 0144-8609/87/S03.50-- Elsevier Applied Science Publishers ktd, England. 1987. Printed in Great Britain

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C. P. Madenfian, G. L. Rogers, A. W. Fast

by some mechanical device. Both methods are energy intensive and should not be used unless needed. If early morning DO in the pond was predictable based on a few measurements made at sunset or a few hours thereafter, the farmer would then be able to determine whether emergency aeration was necessary. Such predictions would allow the farmer to aerate only when needed a n d t h u s reduce operating expenses while at the same time achieving intensive levels of crop production. The intent of this study was to describe mathematically the night time dissolved oxygen dynamics of prawn (Macrobrachium rosenbergii) ponds in Hawaii based on a set of a few variables that could be easily measured by the prawn farmer. Accomplishing this objective would furnish the prawn farmer with a useful preductor of early morning DO in the ponds. The night time dynamics of dissolved oxygen in channel catfish (Ictalurus punctatus) ponds in Auburn, Alabama have been successfully modeled by Boyd et al. (1978). Their model accounts for changes in oxygen concentration due to diffusion, oxygen loss from catfish respiration, loss from sediment respiration, and oxygen consumed through plankton respiration. We shall use the term plankton respriation interchangeably with water column respiration. Of the four aforementioned components, plankton respiration was the greatest factor of DO decline in the Auburn catfish ponds (Romaire, 1979). Romaire attributed the success of the model to the reliable prediction of plankton respiration. Romaire was able to predict plankton respiration as a function of water temperature and secchi disc visibility in an equation of the following form:

r=co+cISDD+c2SDD2+c3T+c4T2+csSDD T

(1)

where r=plankton respiration rate (mg O: liter-~ h-~), SDD =secchi disc depth (cm), T-- water temperature (C), c., c~, c 2, c 3, c4, c5 = regression coefficients. The fit of the equation to the Auburn pond data was good (R 2 = 0.82). In Hawaii, Losordo (1980) explained 79% of the variation in plankton respiration rate of freshwater prawn ponds using a regression analysis similar to that of Romaire's, but used relatively few observations. Losordo used only three observations at 28C, and only three observations at 32C. The only other incubation temperature employed in the experiment was 24C, at which 20 observations were recorded. Several researchers have concluded that plankton respiration is usually the major consumer of oxygen at night in freshwater prawn ponds of Hawaii (Losordo, 1980; Costa-Pierce et al., 1984; Costa-

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Pierce, pers. comm., Oceanography Dept., Univ. Hawaii, Honolulu, HI). Losordo found that the water column respiration (as measured in the dark bottle) accounted for, on average, about 60% of the overnight DO decline observed in the pond. Based on his simulation model, Romaire (1979) estimated that, on average, about 80% of the overnight DO decrease in the Auburn pond could be attributed to the plankton respiration rate. An average sediment respiration rate of 60 mg 02 m -2 h -~ was reported by Costa-Pierce et al. (1984) for Hawaiian prawn ponds and was the average rate suggested for use in modeling efforts (Costa-Pierce, pers. comm.). However, Fast et al. (1983) and Garza (unpublished manuscript, Hawaii Institute of Marine Biology, Univ. Hawaii, Kaneohe, HI) reported mean sediment respiration rates in Hawaiian prawn ponds of 221 and 281 mg 02 m-2 h-~, respectively. The difference between these estimates and those of Costa-Pierce et al. (1984) may be attributed to differences in technique or location. In an entirely different approach to predicting dawn DO in the Auburn catfish ponds, Boyd et al. (1978) concluded that the DO decline during the night was essentially linear. By measuring DO at dusk and then again 2-3 h after dusk, the morning DO prediction could be extrapolated from the line connecting the two evening DO observations plotted against time (Fig. 1 ). This extrapolation technique performed as well as Romaire's simulation model described above for Alabama fish ponds.

10
8t'N

O
(3)

64-

%` %` %` %` %. %. %.

E """*

O
t'l 2
0

PREDICTED ~ DAWN DO
F

1800

2400

0600

HOUR
Fig. 1. Illustration of the linear extrapolation technique for predicting pond DO at dawn.

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C P. Maden]ian, G. L. Rogers, A. W. Fast

METHODS
Plankton respiration as a function of secchi disc depth

To determine the relationship between plankton respiration, temperature, and secchi." disc depth, the following experiment was conducted. Three prawn ponds (ponds AF1, AF2, and AF3) from a relatively calm (sheltered from wind) site (Aquatic Farms Ltd, Kaneohe) and three prawn ponds (ponds A36, D17, and D18) from a windy site (Amorient Aquafarm Inc., Kahuku) on Oahu were sampled from April to August 1984. During each sampling trip, water was collected from each of the three ponds at the site and secchi disc depths were recorded. Thirty-one trips were undertaken: 15 to Aquatic Farms and 16 to Amorient. Water samples were transported to the University of Hawaii at Manoa for incubation. Pond water was collected at a depth of 10 cm. For each pond sampled, separate incubations were conducted at three different temperatures (21, 26, and 32C). Prior to measuring respiration rates, the water was mixed by bubbling air into it. This allowed the sample to reach a water temperature close to the desired incubation temperature and to bring the water close to the dissolved oxygen saturation point before the incubation began. After approximately 1 h in the bath, the samples were poured into 300-ml BOD bottles (two replicates for each of the three temperature levels) and incubated in the dark for 4-7 h. Respiration rates were calculated by measuring oxygen levels before and after incubation for a known period of time. DO was measured using a YSI model 54 oxygen meter (Yellow Springs Instrument Company Inc., Yellow Springs, Ohio) with a BOD probe equipped with a stirrer. Data were analyzed using the Statistical Analysis System (SAS) package (1982). Regressions were performed on the raw data (without averaging of replicate observations) and on averaged data. Statisticians may argue that by averaging, the analyst eliminates the error contributed by replication and that this error should be included in the regression analyses, but we averaged (averaged among replicated observations)in order to compare our results with Romaire's results. Additionally, from each of the ponds sampled from April to August 1984, two replicate volumes of pond water (50 or 100 ml volumes) were filtered through a GF/C Whatman glassfiber filter (using a low-pressure vacuum) to determine the chlorophyll a concentration in the pond water. Two ml of saturated MgCO 3 solution were added to the water as it was filtered. Filters were stored dry (with desiccant) and then were ground (using tissue grinder) and dissolved in 90% acetone solution (by volume

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in water). Samples were allowed to steep for at least 24 h and then absorbance of the chlorophyll extract was read on a Beckman DU-7 spectrophotometer. The procedure outlined by Jeffrey and Humphrey (1975) was used to estimate chlorophyll a from the extract absorbance data.

Other indicators of plankton respiration rate


In an experiment similar to the one described above, other indicators, including in vivo fluorescence and turbidity measured with a Hach turbidimeter (Hach Company, Loveland, Colorado), were tested. Water samples were collected at a depth of 10 cm from each pond sampled; and two replicate aliquots (between 3 and 5 ml) were pipetted from the water sampling bottle to a cuvette, and in vivo fluorescence was measured using a fluorometer (Turner model 111). Water turbidity was measured using a Hach turbidimeter. Additionally, a 5-6 ml aliquot of pond water was pipetted into a cuvette and absorbance (at 665 nm) of the pond water was measured with a spectrophotometer. In vivo fluorescence and turbidity were measured within 1 h of sample collection. Twenty-two trips were undertaken. During each sampling trip, six ponds were sampled; usually three from Amorient in Kahuku and three from the farm ponds at Brigham Young University-Hawaii Campus (BYU) in Laie. However, during a few trips all six of the ponds sampled were BYU ponds. (Note the BYU site was an intermediate site as far as wind speed was concerned; it was partially sheltered from wind, but not as much as the Aquatic Farms site). Each of the seven prawn ponds at the BYU farm were sampled at least once from February to April 1985 and four different ponds (A31, A36, D 17, and D 18) were sampled at Amorient. Plankton respiration rate was measured as described above. For a particular trip, all water was incubated at one of the three temperatures (21, 26, and 32C).

Overnight DO monitoring at Amorient ponds


From June to September 1985, 15 overnight monitoring trips were transacted at Amorient farm in Kahuku. Two ponds were monitored during each trip. DO and water temperature were measured at hourly intervals from sunset until after sunrise (until DO started to increase) at two sites (west and east sides of pond) at each pond. Surface and bottom temperature and DO were measured at each site using a YSI model 54 oxygen meter with a field probe with stirrer. Seven different ponds were monitored (AI8. A19, D11, D12, D13, D17, and D18).

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A 2-liter water sample was collected from each pond monitored; the sample was mixed well, allowed to de-gas for 10 min, and a 300-ml BOD dark bottle (lined with aluminum foil and taped with black electrical tape) was filled (allowing to overflow at least one volume of the bottle) and fixed immediately with 2 ml of MgSO 4 solution and 2 ml of alkaliiodide-azide solution. Another dark bottle was then filled and allowed to incubate in the pond overnight by attachment to a wooden stake driven into the pond bottom. About 1 h before dawn, this bottle was fixed as described above. Winkler titrations were performed to determine DO (Environmental Protection Agency, 1983). The respiration rate of the water column (as determined for the dark bottle) was compared with the whole pond respiration, as determined by estimating the respiration coefficient (see Madenjian e t aL, 1987). The whole pond respiration rate was adjusted for the temperature at which the pond was 100% DO saturated (adjustment was based on whole pond respiration rate as an exponential function of water temperature; with exponential constant, k, equal to 0.0542). The respiration rate in the dark bottle suspended in the pond was adjusted for the temperature at which the pond was 100% DO saturated. It was assumed that respiration rate of the plankton was an exponential function of temperature:
r = c e kT

(2)

where r is the respiration rate in mg O: liter- ~h- t, c and k are constants, and T is water temperature (C). k was estimated from the respiration experiments conducted in the lab (where samples were incubated at constant temperature) to be 0.0542. c was estimated by finding the average temperature during incubation in the pond and by calculating the average respiration rate:
e
c = --

kT Y
(3)

The average respiration rate was found simply by dividing the drop in DO by the duration of the incubation in the pond: (initial D O - final DO) (duration of incubation) The average temperature was calculated by fitting an exponential function to the temperature data:
T = a e h'

(4)

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where Tis temperature (C), a and b are constants, and t is time. By integrating over time and dividing by time span of incubation, average temperature was found. Overnight wind speed data coinciding with the sampling trips at Amorient were furnished by the Hawaii Natural Energy Institute, Univ. Hawaii. Wind speed was measured once every 6 min at a height of 24.4 m (Kahuku tower) above the Kahuku fiat approximately 2 km from the Amorient ponds. An average wind speed for each sampling night was calculated. The linear extrapolation technique was applied to the DO monitoring data from Amorient. Additionally, an exponential (or log-linear) extrapolation technique was applied to the same data as follows: the two DO observations (one at dusk and one several hours later) were logtransformed, the logarithmic dawn DO was projected from the straight line connecting the log-transformed observed DO values plotted against time, and the antilogarithm of the logarithmic dawn DO was found. The antilogarithm of the log(dawn DO) was the predicted DO for this loglinear extrapolation technique. For the linear extrapolation technique, if dawn DO was extrapolated to be less than zero, the dawn DO prediction was set equal to zero. Extrapolations were produced based on: (1) observations from the east side of the pond only, (2) observations from the west side of the pond only, and (3) the average of the observations from the west and east sides (an average for the pond; we will designate this set as the pond set). The surface and bottom DO were averaged for each of the three sets of data.

RESULTS
Secchi disc depth as an indicator of plankton respiration rate

The fit of the regression equations for both sites pooled as well as both sites separately were not adequate for prediction purposes (R: between 0-48 and 0.54)(Table 1 ). Variation in plankton respiration rate appeared to increase with increasing turbidity (Fig. 2). For a given day of sampling, pond D 17 at Amorient and pond AF3 at Aquatic Farms usually exhibited a higher respiration rate yet less turbid water than the other two ponds at the site, respectively. The same regression analysis as for all six ponds pooled was performed on data just for these two ponds. The fit for data from ponds D17 and AF3 only was adequate for prediction (R: = 0.81) (Table 1). The regression equation
was:

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TABLE 1 Fraction of Variation (R 2) of Plankton Respiration Rate Explained as a Function of Secchi Disc Depth and Water Temperature in Hawaiian Prawn Ponds (The function was a linear combination of secchi disc depth and water temperature, the quadratic terms of both of these factors, and the interaction term between secchi disc depth and water temperature. Data were for Amorient and Aquatic Farms, Oahu, April to August 1984) Ponds included in analysis Raw data (without averaging) R~ Averaging replicates

Ponds from both sites pooled Amorient ponds Aquatic Farms ponds Ponds AF3 and D17 only

0"536 0"~,80 0"510 0"811

0.539 0.484 0.514 0.813

0.4
o o oo

n-" ~

0.3 -0.2

**

o9
o

o2~] o

lit

o
o

U.I or" 0.0


I

20

40

60

SECCHI DISC DEPTH (cm)


Fig. 2. Respiration rate of plankton (as determined by dark bottle) versus secchi disc depth in Hawaiian prawn ponds for incubation temperature of 26C. Data for both Amorient and Aquatic Farms, Oahu, were pooled. Experiments were conducted from April to August 1984.

r = - 0 - 0 1 - - 0 " 0 0 6 4 S D D + 0 " 0 0 0 1 6 9 S D D 2 + 0-016 T + 0 . 0 0 0 0 8 1 2 T 2 - 0.000344SDD x T w h e r e r - - p l a n k t o n respiration rate (mg 0 2 liter-1 h-1), S D D - - s e c c h i disc d e p t h (cm), T = w a t e r t e m p e r a t u r e (*C). C h l o r o p h y l l a, as d e t e r m i n e d b y extraction and measuring a b s o r b a n c e of the extraction on a s p e c t r o p h o t o m e t e r , was n o t a practical indicator of

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plankton respiration, since the procedure was too tedious and time consuming. However, just to investigate the variation in the same respiration data that was regressed on temperature and secchi disc depth (April to August 1984), respiration rate was regressed as a function of chlorophyll a and temperature using eqn (1) but substituting chlorophyll a for secchi disc depth in the equation. Chlorophyll a was not a suitable indicator of plankton respiration rate; it explained just slightly more variation in the respiration rate than secchi disc depth. The R 2 values ranged between 0.47 and 0.60 (sites pooled and sites separate) (Table 2). Variation in respiration rate increased as chlorophyll a concentration increased. Interestingly, the regression model of respiration on chlorophyll a and temperature based on data for ponds A F 3 and D17 only, explained about 80% of the variation in the respiration rate data. The results for chlorophyll a paralleled the results for secchi disc depth. TABLE 2 Fraction of Variation (R 2) of Plankton Respiration Rate Explained as a Function of Chlorophyll a and WaterTemperature in Hawaiian Prawn Ponds (The function was a linear combination of chlorophyll a and water temperature, the quadratic terms of both of these factors, and the interaction term between chlorophyll a and water temperature. Data were for Amorient and Aquatic Farms, Oahu, April to August 1984)
Ponds included in analysis R2

Ponds from both sites pooled Amorient ponds Aquatic Farms ponds Ponds AF3 and D17 only

0"600 0"473 0"530 0"804

Other indicators of plankton respiration rate


bz vivo fluorescence of a water sample is easily measured; unfortunately fluorescence was also an inadequate predictor of plankton respiration rate, as shown by data for Amorient and B Y U farms, February to April 1985 (Table 3). Fluorescence and temperature accounted for less than 60% of the variation in the respiration rate data. The turbidimeter measurement as well as the spectrophotometer reading (at 665 nm) were inaccurate indices of water column respiration rate in this data set (February to April 1985) (Table 3). Thus, none of the potentially useful candidates for an indicator of plankton respiration rate tested in this study were highly reliable.

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TABLE 3 Fraction of Variation (R 2) of Plankton Respiration Rate Explained as a Function of an Indicator and Water Temperature in Hawaiian Prawn Ponds (The function was a linear combination of the indicator and water temperature, the quadratic terms of both of these factors, and the interaction term between the indicator and water temperature. Indicators included in vivo fluorescence, turbidity (as measured by Hach kit meter), and absorbance (at 665 nm). Data were for Amorient and BYU farm. Oahu, February to April 1985) Ponds included in analysis In vivo fluorescence Ponds from both sites pooled Amorient ponds BYU farm ponds 0-551 0.598 0.594 R2 Turbidity Absorbance

0"181 0.588 0.195

0.171 0.413 0.186

Monitoring of DO at Amorient ponds


The log-linear (or exponential) projection technique proved to be a substantially more accurate method of predicting dawn DO than the linear projection technique (Table 4). In the case of the second DO observation made 3 h after the dusk DO observation (east and west observations averaged), the linear extrapolation routine underestimated the dawn DO in all 30 trials. The linear predictions for a lag of 3 h were, on average, 2.21 mg O: liter-~ lower than the observed pond average DO; and the prediction was as much as 4.53 mg liter-~ lower than the observed dawn DO and, at best, was only within 0.9 mg liter-~ of the actual dawn DO. In 27 of the 30 trials, the dawn DO was underestimated by the loglinear extrapolation. For DO measured 3 h after dusk, the average absolute difference between the predicted and observed average pond DO (east and west observations averaged) at dawn was 0.66; the maximum absolute difference (or absolute value of the residual) was 1.42 and at best the prediction was essentially equal to the observed DO. Whole pond respiration rates at the ponds monitored at Amorient ranged from 0.324 to 1-059 mg 02 liter-~ h-1; whereas water column respiration rates (as measured in dark bottles) ranged from 0"035 to 0.478 mg 02 liter-t h-t (Table 5). Water column respiration rate (or plankton respiration rate) accounted for 8-3-52.3% (42-0% on average) of the whole pond respiration rate. The difference between the whole pond respiration rate and the plankton respiration rate ranged from

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TABLE 4 Minimum (Min). Maximum (Max), and Mean Absolute Value of the Residuals from the Linear Extrapolation and the Log-Linear Extrapolation Techniques Applied to Amorient Prawn Pond Data, Oahu, June to September 1985 (The residual was the observed dissolved oxygen concentration (DO) minus the predicted DO (in mg O~ liter-f). Lag referred to the number of hours between the time of the DO measurement at dusk and the time of the second DO measurement. DO was measured from both the east and west sides of the pond; pond DO was the average DO of the west and east side observations)
Lag Min West Max Mean Min East Max Mean Min Pond Max Mean

Linear extrapolation residuals (mg 02 liter- t) 1 0"25 5"27 2"94 0"19 5.95 2 0"74 4'59 2"64 0"02 5.18 3 0-71 4'46 2"27 0"07 5.18 4 0"01 3"75 1'74 0'12 4.69 5 0'40 2"56 1-36 0"05 3.97 6 0'04 1"77 0-93 0"02 2.33 Log-linear extrapolation residuals (mg 02 liter- J) 1 0.28 3"34 1"19 0"21 5'66 2 0-03 2'06 0"86 0.05 2"32 3 0"03 1"83 0"73 0"02 2"17 4 0.19 1"35 0-65 0"05 2"32 5 0.02 0'94 0"48 0.01 1"82 6 0.05 0"60 0'34 0'02 1'15

3.39 2.37 2.04 1.68 1.28 0.79 1'68 0"88 0"78 0"69 0-52 0"34

0.32 0.57 0-90 0.43 0.39 0.01 0'22 0"11 0"00 0"07 0"04 0"02

5.52 5.52 4.53 3.74 3.08 1.91 2'46 1"76 1"42 1"12 1"13 0"70

2.97 2.57 2.21 1.69 1.32 0.85 1'16 0"80 0"66 0"56 0"43 0"27

a b o u t 0.19 to 0"80 mg O , liter- J h - Nof D O . T h e c o n t r i b u t i o n of p r a w n respiration to the whole p o n d respiration rate was minimal; L o s o r d o (1980) estimated the p r a w n respiration rate for A m o r i e n t p o n d s to be b e t w e e n 0.04 and 0.07 mg O , liter-~ h-~. We estimated ,the s e d i m e n t respiration by subtracting 0.05 mg liter-~ h - ] (estimate for p r a w n respiration) f r o m the difference b e t w e e n the whole p o n d respiration a n d the water c o l u m n respiration. S e d i m e n t respiration rate ranged f r o m 0.136 to 0.748 with a m e a n of 0.324 mg O~ liter- ~h - l

DISCUSSION T h e B o y d - R o m a i r e - J o h n s t o n m o d e l described above, in which diffusion, p l a n k t o n respiration, s e d i m e n t respiration, and respiration of the cultured organisms are estimated individually, was i n a p p r o p r i a t e for use with H a w a i i a n p r a w n ponds. R o m a i r e (1979) p o i n t e d out that the bases

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TABLE 5 Whole Pond Respiration Rate (WPR) and Plankton Respiration Rate (PR) at Time of 100% Dissolved Oxygen Saturation for Amorient Prawn Ponds, Oahu, June to September 1985 (Sediment respiration rate (SR) was equal to WPR- P R - 0.05. PR/WPR represented the plankton respiration rate fraction of the whole pond respiration rate. Date was for start of sampling period)
Date Pond Temp. at saturation (C)
WPR

Respiration rate in mg 0,_ litre - ~ hrPR SR

PR WI'R

6 June 85 8June 85 13 June 85 22 June 85 27 June 85 11 July 85 21 July 85 24 July 85 1 Aug 85 6 Aug 85 8 Aug 85 24 Aug 85 31 Aug85 7 Sept 85 21 Sept 85 Mean

D17 DI8 D17 D18 D17 D18 D17 DI8 D17 D18 D12 DI3 D12 DI3 D11 DI2 D12 D13 D 17 D18 D 12 D 13 DI2 DI3 D17 DI8 D 17 D18 A18 AI9

Standard deviation

25"9 25"8 26.6 26.3 26.5 25.4 25.2 24.7 27"9 27.4 27.5 26.9 26.4 25'9 27.7 27.1 27.1 26.9 25.2 25.2 26.5 26.2 27.6 27.6 25.3 25-3 25-7 25.7 24.9 24.8 26.2 1.0

0"745 0.324 0.827 0.375 0.824 0.370 0.677 0.403 0.925 0.426 0.689 0.501 0.830 0.726 0-579 0.698 0.863 0.742 0.820 0-396 0.871 0.685 0-582 0.569 0.878 I).388 0.750 0.411 1.059 0.421 0.645 (5.204

0.278 0.138 0.301 0.157 0.270 0.167 0.295 0-186 0.478 0.190 0.325 0.260 0.361 0.308 0-200 0.351 0-396 0.315 0.359 0.166 (5.372 0.304 0.254 0.234 0-404 (5-181 0.392 0.190 0.261 0.035 0.271 0.097

0.418 0.136 0-476 0-169 0.504 0.154 0.332 0-167 0.397 0.186 0.314 0.191 0.419 0.368 0.330 0.297 0.417 0.377 I).411 I).180 0.449 0"331 (5.277 0-285 0.424 0.157 0.308 0-171 0.748 I).336 [5.324 /). 135

0.372 0.427 0.364 0.417 0-328 0.450 0.435 0.461 0'517 0.446 0.472 0.518 0.435 0.425 0.344 0.503 0.459 0.425 0.437 0.420 15.427 0.443 0.437 15.411 0.460 (5-466 0.523 0-462 I).247 0.083 0.420 0.086

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for success of the model were: (1) a strong relationship between secchi disc depth and plankton respiration rate and (2) plankton respiration rate being the major component contributing to night time DO loss (plankton respiration accounted for, on average, about 80% of the night time drop in DO). It appeared that neither of these two conditions were met in Hawaiian prawn ponds. In general, secchi disc depth was not a reliable indicator of plankton respiration in Hawaiian prawn ponds. There was substantial, unexplained variation in respiration rate in Hawaii, particularly for more turbid water. This incongruity between Alabama and Hawaii could have been due largely to differences in turbidity between the two locations and the nature of the turbidity. Secchi disc depths used in Romaire's regression analysis ranged from 17 to 153 cm; while the range of secchi disc depths for this study was from 6"5 to 56 cm, yet the respiration rate for a given secchi disc depth and temperature was higher for the Auburn ponds than for the Hawaiian prawn ponds. These data suggested that the background turbidity (turbidity due to factors other than plankton) in the Hawaiian ponds is higher than that in Auburn ponds. Apparently, this background turbidity was relatively constant among the different ponds and remains constant with time in Alabama. However, the background turbidity did not appear to be uniform among Hawaiian prawn ponds. The above discussion was based on the assumption that for a given region (Alabama or Hawaii), respiration rate was uniform among ponds for a given unit of plankton turbidity. Of course, there was the possibility that for a given unit of plankton turbidity, the respiration rate was not uniform among the ponds because of differences in the physiological state of the plankton or differences in the plankton species composition between ponds. Another factor to consider when looking at differences between the ponds in Hawaii and those in Alabama was wind speed. Wind velocities at the Auburn site were relatively low, while in Hawaii the tradewinds blow year round with variable intensity. Wind can cause resuspension of settleable material, thus increasing turbidity. Additionally. the phytoplankton species composition was notably different: diatoms were common in Hawaiian prawn ponds (R. York, pers. comm., Hawaii Institute of Marine Biology, Univ. Hawaii, Kaneohe, HI) but were rare in Auburn ponds (C. Boy& pers. comm.. Dept of Fisheries and Allied Aquacultures. Auburn Univ., Auburn, AL). The good fit to data from ponds AF3 and D17 only, suggested that the background turbidity was similar for these two ponds and that it was not subject to the same variability as the other ponds during the study period, for some reason. Since ponds AF3 and D17 showed higher respiration rates for a given secchi disc visibility and temperature than

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other ponds, a model employing the regression equation based on these two ponds would tend to underestimate the morning DO in the other four ponds studied. Chlorophyll a determined by extraction and measuring absorbance on a spectrophotometer was also not a highly reliable indicator of water column respiration. Perhaps the respiration rate per unit of chlorophyll a varied between species and between different physiological states of the same species. In vivo fluorescence was not a good predictor of plankton respiration rate. Limitations of in vivo fluorescence as a predictor of chlorophyll a have been noted by Loftus and Seliger (1975) and Strickland (1968). Confounding the correlation between in vivo fluorescence and respiration rate, again, were species differences and different physiological states. Not to be overlooked was the contribution of bacterial respiration to the plankton respiration; bacterial respiration may have contributed substantially to the water column respiration. Chlorophyll a determination by extraction or in vivo fluorescence would not be sensitive to assessing most bacteria populations. It was hoped that measuring turbidity by the Hach meter, and thus avoiding the subjectivity of measurement by secchi disc depth, would lead to a strong relationship between turbidity and respiration rate. However, turbidity, regardless of how it was measured, did not forecast plankton respiration rate well. More detrimental to the usefulness of the Boyd model in Hawaii than not having found an accurate indicator (accurate and easily measured by the farmer) of plankton respiration, was the evidence indicating that the water column respiration was not the major contributor to the whole pond respiration rate. The average benthic respiration rate from the Fast et al. (1983) study of 221 mg 02 mg--" h-~ for a l-acre pond with average depth of 0.75 m corresponded to a rate of 0.299 mg O~ liter-~ h-J; this figure was in fairly close agreement with the average benthic respiration rate estimated from our study at Amorient (0.324 mg O, liter-

h-t).

Losordo's claim that the water column respiration was the major contributor to DO decline in the Amorient ponds was not justified since the diffusion effect was not accounted for. In Losordo's study, when DO dropped below saturation in the ponds, and if wind speeds were substantial (as Lorsodo reported), then oxygen would diffuse into the pond at a substantial rate. Thus, it is likely that more oxygen was consumed overnight by the whole pond than indicated by the difference between dusk and dawn DO measurements. Sediment respiration rate was difficult to study and to the knowledge of the authors, no regression equation using some easily-measured indicator variables and temperature has been

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developed to explain a high percentage of variation in sediment respiration data. Sediment respiration rate estimates from our study at Amorient, as well as the sediment respiration of Hawaiian prawn ponds measured by Fast et al. (1983) and Garza (unpublished manuscript), showed high variability. Boyd used one sediment respiration rate to predict sediment respiration for all the Auburn ponds. This method was adequate for Auburn ponds since, apparently, there was little variation in sediment respiration rate between ponds and the benthic respiration was a relatively unimportant component of whole pond respiration. However, the use of a median or averaged sediment respiration rate to predict sediment respiration for all ponds would yield inaccurate results for Hawaiian prawn ponds at Amorient; mainly due to the high variability and the importance of benthic respiration in Hawaiian ponds. The method of accounting for diffusion in Boyd's model was also inappropriate for Hawaiian prawn ponds, particularly at the Amorient (windy) site. Wind speeds averaged from about 4.5 to 10 m s-~ during the night for the DO monitoring study at Amorient. Boyd used the approach of Schroeder (1975), in which the % saturation of the pond at dusk would be used to forecast the overnight gain or loss of DO by diffusion. The ponds for Schroeder's study were subjected to relatively low wind speed. Suppose a pond was more than 120% DO saturated at dusk (not uncommon for Amorient or BYU ponds). Using Schroeder's approach, the net effect for diffusion overnight would be predicted as a loss of DO from the pond. However, at the Amorient ponds (subjected to strong winds at times), a pond supersaturated in DO at dusk may' drop within a few hours after dusk to below the DO saturation point and thus be undersaturated in DO for most of the night. In this case, the net overnight effect of diffusion on the pond may have been a gain of DO in the pond (see Madenjian et al., 1987). The linear extrapolation method endorsed by Boyd was a poor predictor of dawn DO in Hawaiian prawn ponds. The log-linear projection technique was superior to the linear technique for predicting dawn DO at the Amorient ponds. We may have expected an exponential decline in DO during night time at Amorient (windy site) because'of: (1) diffusion effect and, less importantly, (2) the overnight drop in water temperature. Diffusion of oxygen out of the pond would occur more quickly the higher the DO was above the DO saturation level; the more the DO was below the saturation level, the faster oxygen would diffuse into the pond. Thus, diffusion would act to decrease the rate of decline of DO in the prawn ponds, as the night progressed. Additionally, as temperature dropped (on average about 2-3C for Amorient ponds during the study), so too would the whole pond respiration rate.

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Note that the log-linear extrapolation technique was insensitive to wind speed. Analysis of the residuals (difference between the observed and predicted values) from application of the log-linear method to Amorient ponds revealed that the dawn DO was more accurately estimated (usually underestimated to a lesser degree) as wind speed decreased. As wind speed increased, we would expect the diffusion effect to be stronger. Thus, the degree of curvature of the DO decline would be influenced by wind speed. With increasing wind speed, DO (assuming supersaturation at dusk) would arive at the saturation level more quickly (show a sharper decline to saturation); but when below DO saturation, as wind speed increased, DO would not decrease as quickly. The effect would be more of a bend in the overnight DO curve as wind speed increased. The log-linear extrapolation technique was not tested for still conditions (wind speeds near zero). If we assumed the diffusion effect to be negligible when wind speed is near zero (assuming a negligible decline in temperature), a linear decline in DO during the night would have been predicted. In the case of still conditions, we may have expected the loglinear extrapolation to substantially overestimate the dawn DO (particularly if respiration rate of the pond was high). Prediction of a 'safe' dawn DO level when in fact the dawn DO level was low (say, below 2 mg O2 liter-l) would represent an undesirable situation to the prawn pond manager. A model that incorporated the diffusion effect as a function of wind speed would probably forecast dawn DO more accurately than the log-linear extrapolation method.

SUMMARY The Boyd et al. (1978) model, in which the ovemight change in pond DO was modeled as a function of four components: plankton respiration, sediment respiration, diffusion, and respiration of the cultured organisms, was inadequate for Hawaiian prawn ponds. The methods used by Boyd et al. (1978) to predict plankton respiration, sediment respiration, and the diffusion effect were inaccurate when applied to prawn ponds in Hawaii. Furthermore, the data from our study at Amorient suggested that sediment respiration was an important component to the whole pond respiration rate in at least some of the Hawaiian prawn ponds. The night time decline of DO at Amorient (Kahuku, a windy site on Oahu) was better characterized as exponential rather than linear. The log-linear (or exponential) extrapolation technique produced fairly accurate estimates of dawn DO for the Amorient ponds. However, its

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use may be limited since it does not account for the diffusion effect as a function of wind speed.

ACKNOWLEDGEMENTS Gerald Akiyama is acknowledged for his expert advice on laboratory preparation and on maintenance and operation of the equipment. Darwin Bohnet and Eti Eves assisted with field and lab work at the BYU facility. We gratefully acknowledge Roger Fujioka and Henry Gee of the Water Resources Research Center, University of Hawaii for use of their fluorometer and BOD oxygen probe. Robert Young of Oceanic Institute (Waimanalo, Oahu) kindly loaned us several fluorometer lamps. Wind speed data were provided by James Bac, Sueji Yano, and George Curtis of the Hawaii Natural Energy Institute, University of Hawaii. Thanks to Ken Kamiya, manager of the BYU farm, and Ed McSweeny, manager of the Amorient farm, for their cooperation. This work is a result of research ('Development of Pond Oxygen Management Devices and Predictive Procedures for Increasing Prawn Production' project, A/R-19) sponsored, in part, by the University of Hawaii Sea Grant College Program under Institutional Grant No. NA81AA-D-00070 from NOAA, Office of Sea Grant, Department of Commerce; and sponsored, in part, by the Aquacultural Development Program of Hawaii (Contract No. 17663). This is Sea Grant publication UNIHI-SEAGRANT-JC-87-04 and Hawaii Institute of Marine Biology Contribution No. 726.

REFERENCES Boyd, C. E., Romaire, R. E & Johnston, E. (1978). Predicting early morning dissolved oxygen concentrations in channel catfish ponds. Trans. Am. Fish. Soc., 107,484-92. Costa-Pierce, B. A., Craven, D. B., Karl, D. M. & Laws, E. A. (1984). Correlation of in situ respiration rates and microbial biomass in prawn (Macrobrachium rosenbergii) ponds. Aquaculture, 37, 157-68. Environmental Protection Agency (1983). Methods for Chemical Analysis of Water and Wastes, EPA-600/4-79-020. Fast, A. W., Barclay, D. K. & Akiyama, G. (1983). Artificial circulation of Hawaiian prawn ponds~ Univ. Hawaii Sea Grant College Program, UNIHISEAGRANT-CR-84-01. Jeffrey, S. W. & Humphrey, G. E ( 1975 ). New spectrophotometric equations for determining chlorophylls a, b, cj and c~ in higher plants, algae and natural phytoplankton. Biochem. Physiol. Pfl., 167, 191-4.

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Loftus, M. E. & Seliger, H. H. (1975). Some limitations of the in vivo fluorescence technique. Chesapeake Sci., 16, 79-92. Losordo, T. M. (1980). An investigation of the oxyen demand materials of the water column in prawn grow-out ponds, MSc. Thesis, Univ. of California, Davis. Madenjian, C. P., Rogers, G. L. & Fast, A. W. (1987). Predicting night time dissolved oxygen loss in prawn ponds of Hawaii: Part II. A new method. Aquacultural Engineering, 6, 209-25. Romaire, R. P. (1979). Modeling the dynamics of dissolved oxygen in channel catfish production ponds, PhD. Thesis, Auburn Univ. Schroeder, G. L. (1975). Night time material balance for oxygen in fish ponds receiving organic wastes. Bamidgeh, 27, 65-74. Statistical Analysis System (1982). SAS User's Guide: Statistics, SAS Institute Inc., Cary, North Carolina. Strickland, J. D. H. (1968). Continuous measurement of in vivo chlorophyll; a precautionary note. Deep-Sea Res., 15, 225-7.