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Journal of Applied Aquaculture


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Effect of Contrasting Agents on Survival,


Performance, and Condition of Larval
Hybrid Striped Bass Morone chrysops x
M. saxatilis in Tanks
a

Matthew McEntire , Marty Riche , Benjamin H. Beck & Doug Carter


a

United States Department of Agriculture, Agricultural Research


Service, Harry K. Dupree Stuttgart National Aquaculture Research
Center Stuttgart, Arkansas, USA
b

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KaMin, LLC, Macon, Georgia, USA


Published online: 09 Mar 2015.

To cite this article: Matthew McEntire, Marty Riche, Benjamin H. Beck & Doug Carter (2015)
Effect of Contrasting Agents on Survival, Performance, and Condition of Larval Hybrid Striped
Bass Morone chrysops x M. saxatilis in Tanks, Journal of Applied Aquaculture, 27:1, 1-28, DOI:
10.1080/10454438.2014.959814
To link to this article: http://dx.doi.org/10.1080/10454438.2014.959814

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Journal of Applied Aquaculture, 27:128, 2015


Copyright Taylor & Francis Group, LLC
ISSN: 1045-4438 print/1545-0805 online
DOI: 10.1080/10454438.2014.959814

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Effect of Contrasting Agents on Survival,


Performance, and Condition of Larval Hybrid
Striped Bass Morone chrysops x M. saxatilis
in Tanks
MATTHEW MCENTIRE1 , MARTY RICHE1 , BENJAMIN H. BECK1 ,
and DOUG CARTER2
1

United States Department of Agriculture, Agricultural Research Service, Harry K. Dupree


Stuttgart National Aquaculture Research Center Stuttgart, Arkansas, USA
2
KaMin, LLC, Macon, Georgia, USA

Contrasting agents, either algae or inert soil, cause turbidity,


which is important in the tank culture of larval cannibalistic
fish. Optimization of turbidity is critical to successful tank culture of new larval fish, which should include 100 mg/L of sub-5
m particle size in the assessed range. The optimum tested range
of greenwater culture for hybrid striped bass using algae paste
once daily was between 629 and 1127 mg/m3 (350583 cell/mL).
Dispersed kaolin worked as well as greenwater culture. Using contrasting agents improved fish length, condition, taillength ratio
(TLR), and uniformity of the fish at the time of weaning onto prepared diets. Increasing turbidity appears to negatively skew fish
length and TLR. TLR shows promise as an early indicator of cannibalism. Increasing turbidity appears to decrease the size advantage
of the largest fish in the tank, which allows the smallest fish to
catch up.
KEYWORDS Turbidity, larval fish, variation, cannibalism,
standpipe

Address correspondence to Matthew McEntire, United States Department of Agriculture,


Agricultural Research Service, Harry K. Dupree Stuttgart National Aquaculture Research Center,
2955 Highway 130 E, Stuttgart, AR 72160, USA. E-mail: Matthew.McEntire@ARS.USDA.GOV
Color versions of one or more of the figures in the article can be found online at www.
tandfonline.com/wjaa.
1

M. McEntire et al.

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INTRODUCTION
Sunshine bass Morone chrysops x M. saxatilis represent a valuable sector of
the U.S. aquaculture industry valued at nearly $30 million in 2012. Despite
their commercial importance, one factor limiting the growth of the sunshine
bass industry is the lack of refined methods/techniques to rear sunshine bass
fry/fingerlings in tanks. The advantages of tank culture include year-round
availability of fingerlings, ease of harvest, and consistent production. A major
area for improvement in sunshine bass tank culture is the optimization of the
types of live and prepared feeds, including how they are offered to the fry.
Historically, the use of live feeds has progressed from freshwater rotifers
Brachionus calyciflorus paired with salmon starter (Ludwig 1994), to saltwater rotifers Brachionus plicatilis, Artemia nauplii, and prepared feed (Denson
& Smith 1997; Ludwig 2003). More recently recommendations suggest reverting back to just smaller Artemia nauplii with prepared feed to eliminate the
cost associated with rotifer culture (Ludwig & Lochmann 2009). While selecting the appropriate live organism or prepared feed choice is important, other
factors are emerging as important contributors to the overall success of tank
culture, especially the inclusion of turbidity agents or colorants in larval
settings.
The possible benefit of increased turbidity is not a new concept, and
was suggested in 1941 by Doan examining survival of natural larval fish
populations in Lake Erie, but the intentional use of elevated turbidity in the
culture of larval fish dates to the late 1960s with the addition of live algae to
help control metabolic buildup in the tank (Houde 1972). There are a variety
of items that have been used to increase the turbidity of the water:
1. Greenwater: (live algae and algae concentrates) with striped bass, Atlantic
halibut, grouper, turbot, greenback flounder, cobia, pompano, striped
trumpeter (Cavalin & Weirich 2009; Cobcroft et al. 2012; Duray et al. 1996;
Naas et al. 1992; ie et al. 1997; Riley et al. 2009; Schwarz et al. 2008;
Shaw et al. 2006; Snow et al. 1980)
2. Inert soils: bentonite with bluegill (Gardner 1981); fine clay with sunshine
bass (Rees & Cook 1982); settled river sediment fraction and settled volcanic ash fraction with Pacific herring (Boehlert & Morgan 1985); kaolin
with striped bass (Chesney 1989); Kentucky ball clay #4 with walleye
(Bristow & Summerfelt 1994); diatomaceous earth with Atlantic herring
(Utne-Palm 2004); pulverized kaolinite (Kaoline Polsperse 10) with cod
(Meager et al. 2005); ceramic clay (Vingerling K148) with cod (Attramadal
et al. 2012); bentonite with yellowtail amberjack (Stuart et al. 2012)
3. Dye: AquaShade dye with walleye (Bristow et al. 1996).
Numerous reasons have been reported to explain the benefits of the
colorants. Increased turbidity was suggested to reduce the light reflection
and phototactic attraction of larvae to the tank sides, allowing more uniform

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Effect of Contrasting Agents on Survival, Performance, and Condition

larval distribution within the tank (Loadman et al. 1986; Bristow et al. 1996).
Better horizontal distribution of larvae within tanks due to more uniform
light intensity at the top of the tank was another explanation (Swenson &
Matson 1976). Particles in the water also act as a contrasting agent, allowing better selection of feed items (Nickum 1978; Boehlert & Morgan 1985).
A light-scattering effect of the particles in the water provides better lighting
of semitransparent food items, thereby increasing food visibility (Boehlert &
Morgan 1985). Another observed benefit was faster larval swimming ability
(Rieger & Summerfelt 1997). Other benefits include a reduced perceptive
field of view of fish (Blaxter & Staines 1971; Moore & Moore 1976) and
the reduced perceived predation pressure outweighing the disadvantage of
reduced ability to see and find food (Ritchie 1972). This also results in
a reduced number of negative interactions between fish, i.e., nipping or
aggression (Hetch & Pienaar 1993).
The observed benefit of turbidity to tank larval culture is likely a
combination of factors, summarized as the physical effects hypothesis that
hinges on improved prey contrast and motivational factors, combined with
reduced perceived predation risk (Gregory & Northcote 1993; Hinshaw
1985). Changes in contrast between the prey and background are the most
frequently demonstrable effects but do not refute the behavioral hypothesis,
as summarized from a good overview paper (Utne-Palm 2002).
Species-specific optimization of turbidity (Shaw et al. 2006) should be
elucidated in the same way that light intensity and tank color have been
found to be species specific (Jirsa et al. 2009). Therefore, the principal goal
of this study was to characterize the utility and feasibility of select turbidity
agents at improving sunshine bass survival and uniformity of size at time
of weaning to an artificial dietan area where a substantial knowledge gap
exists. Specifically, we defined the optimum concentration of algal paste
for greenwater culture based upon fish survival and also compared the
performance of a dispersed formulation of kaolin against greenwater and
clearwater culture of tank-reared hybrid striped bass larvae. In addition to
survival, the influences of the contrasting agents on larval size distribution
(i.e., uniformity) at the time of weaning to artificial feed were examined.

MATERIAL AND METHODS


Two concurrent experiments were conducted. The first evaluated the impact
of algal density, i.e., greenwater, on hybrid striped bass fry growth and
survival. The second evaluated the impact of dispersed kaolin relative to
greenwater and clearwater on fry performance.

In Common Experiments 1 and 2


Hybrid striped bass 4-day post-hatch (dph) larvae from separate spawns were
used across each experiment because there was not a spawn of sufficient

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M. McEntire et al.

size to stock both experiments. Larvae were produced at the Harry K. Dupree
Stuttgart National Aquaculture Research Center, Stuttgart, AR, USA, off season
during the fall of 2012. Larvae were counted and stocked into experimental
tanks by volume. All the water treatments were maintained by exchanging
half the tank water daily for 18 days. The 35-L round fiberglass tanks had a
white flat bottom and black sides and an internal stand pipe with 250 m
nylon mesh (Figure 1). The stand pipes were designed to have sufficient
filtering surface area and to prevent fry impingement on the screen mesh
caused by aeration water currents, while still allowing for a tank overflow.
We constructed a PVC cap filled with epoxy attached to a fixed-depth rigid
tubing (Figure 1) in order to slowly and uniformly drain the tanks halfway in a self-regulating fashion, without the risk of fry impingement on the
standpipe screen.
Regardless of experimental treatment and stocking rate, all fish were
grown under the same feeding regime and a 12:12-h light:dark cycle. Lighting
averaged 1,620 lux at the water surface (photometer Model 401027 Etech
Instruments, Waltham, MA, USA). Salinity was maintained at 5 ppt with
Instant Ocean synthetic sea salt (Aquarium Systems, Mentor, OH, USA).
Dissolved oxygen, temperature, pH, (Hach HQ40d portable meter, (Hach
Company, Loveland, CO, USA) and salinity YSI Model 63 (YSI Inc.,
Yellow Springs, OH, USA) were monitored daily. Ammonia and nitrite
(Hach, Loveland, CO, USA) were measured until 8 dph when the addition
of 75 mg/L ChlorAm-X sodium hydroxymethanesulfonate (AquaScience
Research Group, North Kansas City, MO, USA) required the use of a salicylate method (spectrophotometer DR/4000, Hach, Ames, IA, USA) to observe
the decline in calculated un-ionized ammonia (Emerson et al. 1975). With
the addition of ChlorAm-X to compensate for over 1.5 times the highest
measured ammonia level, ammonia testing was discontinued because of the
expense.
Syphoning of tanks was started on day 3 of the experiment and repeated
every other day after that. The daily half volume water exchange was completed following syphoning to compensate for different amounts of water
removed during syphoning. All tanks were syphoned using a volumetric
pipet with the tapered end removed and attached to a clear 3/8 inch vinyl
tubing. The lack of visibility due to the contrasting agents forced most of
the tanks to be syphoned using a mental grid of the bottom of the tank,
such that an area was considered clean when a lightening of the material
exiting the tank through the clear syphon tubing was observed. In several of
the treatments neither the tank bottom nor the fish were visible until water
treatments were diluted at the end of the study.
The modified feeding regime (Ludwig & Lochmann 2007; Ludwig et al.
2008) consisted of L-type rotifers Brachionus plicatilis (Reed Mariculture
Inc., Campbell, CA, USA) batch cultured on Instant Algae Nanno 3600TM
algae paste (Reed Mariculture Inc., Campbell, CA, USA), micro-cyst Artemia,

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Effect of Contrasting Agents on Survival, Performance, and Condition

FIGURE 1 This internal standpipe design allows for a larger screen surface area to accommodate larger flow rate. The fittings and pipe are schedule 40, except for the internal 3 thin wall
pipe. This allows the fine mesh to be jammed between the pipe and reducers. The overlapping mesh material is centered and fixed to the vertical bar on the thin wall pipe. Especially
in small tanks, this prevents delicate larvae from being impinged on the mesh screen by water
currents generated from forced-air aeration. The smooth half-inch PVC pipe at the water/air
interface decreases the likelihood that larvae will contact or stick to the smooth pipe. In addition, it allows for emergency overflow and also provides the ability to slowly syphon drain all
the tanks to a specific water depth. A half-inch PVC cap with a hole drilled in it to just accommodate a plastic volumetric pipet and attached to a vinyl syphon tubing were assembled. The
pipet is set to the preferred drain level, and then the cap is turned upside down and filled
with epoxy to ensure that the depth does not change. The rate of draining can be adjusted up
and down by the diameter of the syphon tubing. By placing the cap onto the half-inch PVC
pipe at the top of the standpipe, the syphon can be started with the water slowly draining
outside the tank to the specified level.

M. McEntire et al.

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and regular cyst Artemia (INVE Aquaculture Inc., Salt Lake City, UT,
USA). Rotifers were fed three times daily (60 rotifers/mL/tank/day) for
6 days and once on the morning of 11 dph (20 rotifers/mL/tank/day).
Starting 9 dph micro-cyst Artemia (Great Salt Lake, UT, USA) were fed
once daily (6/mL/tank/day) for 3 days and twice daily, starting 12 dph
(16/mL/tank/day) for 2 days. Regular cyst Artemia (Great Salt Lake, UT,
USA) were fed three times daily (12/mL/tank/day) starting 14 dph for 2 days
and four times daily (16/mL/tank/day) for 8 days. Artemia cultures were
enriched with Easy Selco (INVE Aquaculture Inc., Salt Lake City, UT, USA)
for 4 h prior to feeding.

Experiment 1
The first experiment consisted of ten different greenwater levels from Nanno
3600TM algae paste stocked at 65 fry/mL. The highest level was calculated
as a final density of 971 algal cells/mL or 2,326 mg/m3 chlorophyll-a and
was made by mixing 50 mL of paste into 100 L of 5 ppt saltwater prepared with Instant Ocean synthetic sea salt in a mixing tank and then
transferred to the culture tank. Subsequent algae levels were prepared by
serial dilution with 3/5 of the higher algae level, mixed with 5 ppt saltwater
to fill the mix tank up to the 100 L mark. Tank treatment levels were confirmed 0.51 h after the daily water exchange by measuring chlorophyll-a
level (mg/m3 ) (Lloyd & Tucker 1988). Additionally, chlorophyll-a levels
were examined before and after the daily water exchange at 17 dph to
determine the range of daily fluctuation. For the first four days of the
experiment, the half-tank make-up water was made by serially diluting a
constant volume of the previous higher level; subsequently a stock solution of the highest level was appropriately diluted by volume to each
culture tank based upon the tanks treatment level and the culture tank
filled with 5 ppt saltwater that contained ChlorAm-X ammonia binder
(75 mg/L).

Experiment 2
In the second experiment four replicate treatments (Clear = Control, Green =
390 mg/m3 from algae paste, Kao A = 1 g dispersed kaolin/tank, Kao B =
4 g dispersed kaolin/tank) were randomly applied to eight tanks stocked
at 48 fry/mL. The kaolin used in this experiment (KaMin LLC, Macon, GA,
USA) had a particle size that is 100% less than 2 microns. The kaolin particle
would typically be 50 nm thick. Note that the dispersed kaolin powder was
weighed out daily and mixed in 30 mL of fresh water before it was added to
the saltwater in the tanks for good dispersion.

Effect of Contrasting Agents on Survival, Performance, and Condition

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Sampling
Prior to feeding on 19 dph (Experiment 1) or 21 dph (Experiment 2),
subsamples of fish (n 70) were anesthetized in 1 g/L tricane methanesulfonate (MS-222, Tricaine-S, Western Chemical, Inc., Ferndale, WA, USA)
and digital images were taken for morphometric analysis using an Olympus
SZX9 dissecting scope (Olympus Optical Co. Ltd., Tokyo, Japan) at 6.3X
magnification equipped with a Zeiss Axiocam|Cm1 camera (Carl Zeiss
Microscopy GmbH, Munich, Germany). Image analysis was completed
with AxioVison Rel.4.8 software (Carl Zeiss Microscopy GmbH, Munich,
Germany). Fish images were taken from each tank on a Sedgwick Rafter
slide (Graticules Ltd., Tonbridge, England) and compared with a photo of a
micrometer (0.01 mm, Olympus Optical Co. Ltd., Tokyo, Japan). The lengths
of the fish were calculated using the software by comparing the ratio of the
known length of the scale bar on the micrometer to its length in pixels. Given
that the pictures were taken under the same conditions, the ratio permitted
the conversion of pixels into length in mm, which correlated well with the
Sedgwick Rafter slide markings and produced an accuracy and resolution
of more than 0.01 m. Photos of unfed initial fish were taken at 5 dph to
determine initial size and variability of each spawn.
The fish were measured for standard length (SL), fork length (FL), tail
length, and body depth (immediately posterior to the swimbladder). Total
length (TL) was the sum of SL and tail length. Type I cannibalism is typified
by tail-first ingestion of the prey (Hecht & Appelbaum 1988). This suggests
that the occurrence of short ratty tails in larval fish may be a sensitive metric
of aggression. Ineffective attempts at type I cannibalism lead to shortened
tails. Measuring tail length without correlation to SL would be misleading,
however, since fish with a long SL and a short tail could have the same tail
length as a fish with a substantially shorter SL and a long tail. To compensate
for this, tail length was divided by SL of the fish to calculate tail length ratio
(TLR). We hypothesized that increasing algae density would decrease fin
nipping and result in a longer TLR and lower variation in TLR. In a similar
manner we created a body depth ratio (BDR) by taking the SL divided by
the body depth to use as a proxy measurement of fish condition similar
to the index of Smith et al. (2005). We selected the more robust bivariate
morphological index of nutritional status (Smith et al. 2005) instead of using
the body depth divided by head depth (Martin & Malloy 1980), due to their
small sample size.
On day 20 dph (Experiment 1) or 22 dph (Experiment 2), water treatments were diluted out with fresh well water in order to see and harvest
fish. Fish in each tank were bulk weighed to the nearest 0.1 mg and then
four subsamples were collected, weighed, and enumerated to determine
average fish weight per tank as well as survival. Animal care and experimental protocols used in this study were approved by the Harry K. Dupree

M. McEntire et al.

Stuttgart National Aquaculture Research Center Institutional Animal Care and


Use Committee and conformed to Agricultural Research Service Policies and
Procedures 130.4 and 635.1.

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Statistical Analysis
Regression analysis was performed for Experiment 1 to characterize
the relationship of response variables to the logarithm of the mean
chlorophyll-a concentration (mg/m3 ). In addition, the relationship between
the morphometric variables, combinations of morphometric variables, or
the coefficient of variation (CV) of the morphometric variables were plotted against logarithm of the mean chlorophyll-a concentration (mg/m3 ) to
look for definable relationships or trends. The skewness and kurtosis of
these variables was also calculated in Excel (Microsoft Excel 2010 version
14.0, Microsoft Corporation, Redmond, WA, USA). Using a forward model
selection procedure, response data were subjected to polynomial and nonlinear regressions in Excel (Microsoft Excel 2010 version 14.0, Microsoft
Corporation, Redmond, WA, USA) to fit the most parsimonious model
(Ratkowski 1990) of response with respect to the logarithm of the mean
chlorophyll-a concentration (mg/m3 ). Parsimony was empirically defined as
the simplest significant model (P 0.05) with the lowest mean square error
and highest R2 0.25 that explained the data. Morphometric variables, their
combinations, or the CV of morphometric variables were compared among
treatments of the second experiment by using a mixed model ANOVA (Littell
et al. 1996) with repeated measures and compound symmetry covariance
structure, using PROC MIXED (SAS version 9.2 statistical software, SAS
Institute Inc., Cary, NC, USA). Pairwise differences among least squared
means were evaluated with the DIFF option and Tukey-Kramer adjustment of the P value (Kramer 1956; Tukey 1953). Statistical differences were
considered significant at P 0.05.

RESULTS
Water quality was suitable for the culture of hybrid striped bass fry across
both experiments, mean plus or minus standard deviation (20.9 0.4 C,
dissolved oxygen 7.7 0.8 mg/L, pH 8.2 0.1, and salinity 5.0 0.1 ppt).
Measured un-ionized ammonia values peaked at 0.10 mg/L in the highest
greenwater treatment, with the rest of the treatments averaging 0.07 mg/L
prior to water exchange and were less than 0.03 mg/L post exchange, and
nitrite levels did not exceeding 0.3 mg/L but was not a concern given the
salinity and amount of chloride present.

Effect of Contrasting Agents on Survival, Performance, and Condition

The serial dilution study demonstrated a clear linear relationship between


calculated number of cells per mL and mg/m3 chlorophyll-a for both the
initial tank treatment levels (y = 0.3086x 1.619; R2 = 0.991) and the experiment mean treatment levels (y = 0.431x + 20.778; R2 = 0.983). During the
final sampling of the experiment, a large percentage of the fish from the
350 cell/mL treatment level were dropped on the floor during transfer to
the scale for weighing. Without any way to collect the fish, these fry were
considered lost, and that data point was dropped from any analysis; this
didnt affect the morphometric data collected earlier. A third-order polynomial relationship defined by (y = 6E11 x3 6E7 x2 + 0.0011x + 0.4738; R2 =
0.9578) was demonstrated for percent survival versus the log-transformed
chlorophyll-a concentration in mg/m3 (Figure 2). The standard length (mm)
and the CV of SL were plotted for all fish and the top and bottom 25% of fish
sample sorted by SL (Figure 3). Fish SL increased with algae density through
the fifth algae level; a statistically significant deviation of the slope from zero
was indicated (P < 0.001) but resulted in a poor linear fit (R2 = 0.119).
However, the dip at the seventh algae dilution (indicated by the arrow in
Figure 3) defies explanation. A narrowing of the top and bottom quartiles
indicate a decrease in sample range starting at the fifth algae level, which corresponds to a decrease in the CV of SL below the fifth and sixth algae level.
The CV of SL of the largest quartile remains low and fairly uniform except
for the increase at the highest algae level. The tail length (mm) and the CV
of tail length were plotted for all fish and the top and bottom 25% of fish
sampled by SL (Figure 4). Tail length of the fish increased with algae density

100%
80%
% Survival

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Experiment 1

60%
40%
20%
0%

10
100
1000
Log Chlorophyll-a (mg/m3)

10000

FIGURE 2 Percent survival of hybrid striped bass fry at weaning onto a prepared diet in
Experiment 1. Data are plotted against the logarithm of the mean of the chlorophyll-a concentration (mg/m3 ), which resulted in a third-order polynomial relationship (y = 6E11 x3
6E07 x2 + 0.0011x + 0.4738; R2 = 0.9578). Arrow indicates the seventh algae level, which
equates to the level used in Experiment 2.

10

M. McEntire et al.

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FIGURE 3 (A) The mean standard length (SL) of hybrid striped bass fry at the time of weaning
onto a prepared diet in Experiment 1 plotted against the mean logarithm of the chlorophyll-a
concentration (mg/m3 ), partitioned into whole tank SL (solid line), top 25% of SL (large
dashed line) and bottom 25% of SL (dotted line). (B) The coefficient of variation (CV) of SL
plotted against the mean logarithm of the chlorophyll-a concentration (mg/m3 ), partitioned
into whole tank CV (solid line), top 75% CV by SL (large dash/dot line), top 25% CV by SL
(large dashed line), bottom 25% CV by SL (dotted line). Arrow indicates the seventh algae
level, which equates to the level used in Experiment 2.

up to the fifth algae dilution, where it declined to the seventh algae dilution
(indicated by the arrow in Figure 4). A similar tightening to what was found
with standard length occurs between the smallest and largest tail length quartiles at the fifth to seventh algae level. A corresponding decrease in overall
CV of tail length occurs thru the sixth algae level, after which there is a
sharp increase in variation (Figure 4). Tail length CV of the smallest quartile
increased from the sixth algae level through the highest level. Tail length CV
of the largest quartile trends downward, staying low before sharply increasing at the highest algae level. Tail length ratio (Figure 5) exhibited the same
trends and shape as the tail length, with the exception of the exaggerated
convergence of the smallest and largest quartiles toward the mean values
using all the fish (overall) at the highest and lowest algae levels. Fish in the
zero algae level for the most part were observed to have worn rounded tails,

Effect of Contrasting Agents on Survival, Performance, and Condition

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FIGURE 4 (A) The mean tail length (TAIL) of hybrid striped bass fry at the time of weaning
onto a prepared diet in Experiment 1 plotted against the mean logarithm of the chlorophyll-a
concentration (mg/m3 ), partitioned into whole tank TAIL (solid line), top 25% of TAIL (large
dashed line) and bottom 25% of TAIL (dotted line). (B) The coefficient of variation (CV) of
TAIL plotted against the mean logarithm of the chlorophyll-a concentration (mg/m3 ), partitioned into whole tank CV (solid line), top 25% CV by TAIL (large dashed line) and bottom
25% CV by TAIL (dotted line). Arrow indicates the seventh algae level, which equates to the
level used in Experiment 2.

resulting in a more uniform TLR, as indicated by the CV of the TLR. Body


depth ratio (BDR) and the CV of the BDR were plotted for all fish, and the
top and bottom 25% of fish with respect to SL (Figure 6). Body depth ratio
(Figure 6) showed improvement in fish condition with increasing algae levels. There was a convergence of the largest and smallest quartiles toward the
overall BDR, with increasing algae levels, which is striking. At the seventh
algae level (arrow) BDR declined sharply, indicating fry were more uniform
and deeper. The CV of body depth (Figure 6) indicated a decline in variation with increasing algae levels, except for the highest algae level, where
variation spiked. There was a similar convergence of the quartiles toward
the overall CV; however, interestingly from the sixth algae level and higher
there was an increase in variation of the smallest quartile of fry. In addition,
the skewness of the TLR was plotted as a function of the mean natural log
transformed algal cell count (Figure 7). There were no other apparent trends

12

M. McEntire et al.

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FIGURE 5 (A) The mean tail length ratio (TLR) of hybrid striped bass fry at the time of
weaning onto a prepared diet in Experiment 1 plotted against the mean logarithm of the
chlorophyll-a concentration (mg/m3 ), partitioned into whole tank TLR (solid line), top 25% of
TLR (large dashed line) and bottom 25% of TLR (dotted line). (B) The coefficient of variation
(CV) of TLR plotted against the mean logarithm of the chlorophyll-a concentration (mg/m3 ),
partitioned into whole tank CV (solid line), top 25% CV by TLR (large dashed line) and bottom
25% CV by TLR (dotted line). Arrow indicates the seventh algae level, which equates to the
level used in Experiment 2.

in swimbladder inflation rate, mean fish weight, fork or total length, body
depth, or CV for those metrics that were not characterized in the figures.

Experiment 2
The kaolin versus greenwater versus clearwater experiment demonstrated
several differences. The comparison of mean fish weight (Figure 8) distinguished both kaolin treatments to be better than the clearwater treatment,
with greenwater being indistinctive from either (P = 0.050). Survival was
not statistically different (P = 0.178) among treatments (Figure 9). The
morphometric measurements (Table 1) indicated that greenwater and kaolin
treatments were significantly different from the clearwater treatments in SL
(P = 0.012), FL (P = 0.010), TL (P = 0.009), tail length (P = 0.004),
body depth (P = 0.023), and TLR (P = 0.015), but not BDR (P = 0.067).

Effect of Contrasting Agents on Survival, Performance, and Condition

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FIGURE 6 (A) The mean body depth ratio (BDR) of hybrid striped bass fry at the time of
weaning onto a prepared diet in Experiment 1 plotted against the mean logarithm of the
chlorophyll-a concentration (mg/m3 ), partitioned into whole tank BDR (solid line), top 25%
of BDR (large dashed line), and bottom 25% of BDR (dotted line). (B) The coefficient of
variation (CV) of BDR plotted against the mean logarithm of the chlorophyll-a concentration
(mg/m3 ), partitioned into whole tank CV (solid line), top 25% CV by BDR (large dashed line),
and bottom 25% CV by BDR (dotted line). Arrow indicates the seventh algae level, which
equates to the level used in Experiment 2.

FIGURE 7 Skewness of tail length ratio of hybrid striped bass fry at the time of weaning onto
prepared diet plotted against the mean logarithm of the chlorophyll-a concentration (mg/m3 )
in Experiment 1. Skewness, a unitless measure (Zar 1999), typically ranges from 1 for an
extremely left skewed (i.e., right-tailed) distribution through 0 (symmetrical distribution) to
+1 for an extremely right skewed (i.e., left-tailed) distribution. Arrow indicates the seventh
algae level, which equates to the level used in Experiment 2.

14

M. McEntire et al.

B
B
AB

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FIGURE 8 Final mean weights (mg) of hybrid striped bass from Experiment 2 are plotted by
treatment: 29 mg kaolin/L (Kao A), 114 mg kaolin/L (Kao B), greenwater (Grn 7th; 390 mg/m3
chlorophyll-a), or clearwater. Statistical differences are denoted by the use of different letters
(P = 0.050). Error bars represent the standard error of least squared means.

FIGURE 9 Final survival of hybrid striped bass from Experiment 2 are plotted by treatment: 29 mg kaolin/L (Kao A), 114 mg kaolin/L (Kao B), greenwater (Grn 7th; 390 mg/m3
chlorophyll-a), or clearwater and were not statically different (P = 0.178). Error bars represent
the standard error of least squared means.

The overall CV was not significantly different among water treatments. The
pairwise comparison among all combinations of water treatments for each
morphometric measurement or combinations of the morphometric measurements were compared by Tukey-Kramer test resulting in the adjusted
p values (Table 2). The CV for each measurement was denoted under the
heading ALL. Subsequently, the sampled fish in each tank were sorted and
ranked by SL and divided in half rounding down, so there were equal numbers of fish represented above and below the median value. Then from the
smallest and largest half two new CV were calculated to compare among
treatments, to determine if there were segregated differences in variation

15

Effect of Contrasting Agents on Survival, Performance, and Condition

TABLE 1 Least squares means (LSM) and standard error of the mean (SEM) for Experiment 2.
The ANOVA p > F value is of natural log transformed data. The means of coefficient of variation of natural log transformed data were compared to determine if there were differences in
variation among treatments: 29 mg kaolin/L (Kao A), 114 mg kaolin/L (Kao B), greenwater
(Grn 7th; 390 mg/m3 chlorophyll-a) or clearwater control. (The ANOVA p > F is presented
in table as CV p > F.)

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Response
Standard Length
(mm)
Fork Length (mm)
Total Length (mm)
Tail Length (mm)
Body Depth (mm)
Tail Length Ratio
Body Depth Ratio

Clear
Grn7th
KAO-A
LSM SEM LSM SEM LSM SEM

KAO-B
LSM SEM

CV
P

6.67 0.21

7.90 0.21

8.09 0.21

8.78 0.21

0.012

0.940

7.30 0.24
7.50 0.24
0.83 0.05
0.99 0.06
0.12 0.00
6.82 0.15

8.84 0.24
9.12 0.25
1.22 0.05
1.29 0.06
0.15 0.00
6.17 0.15

9.02 0.24
9.31 0.25
1.22 0.05
1.28 0.06
0.15 0.00
6.36 0.15

9.78 0.24
10.19 0.24
1.41 0.05
1.47 0.06
0.16 0.00
6.00 0.15

0.010
0.009
0.004
0.023
0.015
0.067

0.954
0.959
0.695
0.552
0.140
0.064

among the small or large fish within a water treatment. There were differences in variation among the large fish between clear and green, clear
and kaolin B, and kaolin A and kaolin B for SL and TL, which leaves no
difference in variation among clear and kaolin A or green and kaolin B
treatments (Table 2). There was a difference between the CV of the BDR
among the small fish, which distinguished clearwater fish in poorer condition from fry in all the other water treatments as well as distinguishing fry
in the two kaolin levels from one another (Table 2). However, there were
no differences among the CV overall at the alpha = 0.05 level. The mean
kurtosis and skewness plus or minus the standard deviation show positive
kurtosis and negative skewness for both SL and TLR for the water treatments
(Figure 10).

DISCUSSION
There are two explanations for the proposed physical benefit of turbidity.
First, particles can act as contrasting agents, or second, particles can improve
light scattering and lighting of food (Boehlert & Morgan 1985). Examining
the size distribution by weight of volcanic ash turbidity versus sediment
turbidity (Boehlert & Morgan 1985) reveals more support for the contrast theory based upon particle size. Excluding larger particles and summing the less
than 4.8 m particles by weight results in 77 mg/L (7.7% of the 1000 mg/L)
ash optimum feeding level and 85 mg/L (17% of the 500 mg/L) sediment
optimum feeding level. Expression of the continuous suspension data based
upon concentration of sub-4.8 m particles correlates the optimum mean

16

Adj P

0.051
0.038
0.034
0.011
0.070
0.029
0.124

Adj P

0.939
0.960
0.960
1.000
1.000
0.096
0.778

Response

Standard Length (mm)


Fork Length (mm)
Total Length (mm)
Tail Length (mm)
Body Depth (mm)
Tail Length Ratio
Body Depth Ratio

Response

Standard Length (mm)


Fork Length (mm)
Total Length (mm)
Tail Length (mm)
Body Depth (mm)
Tail Length Ratio
Body Depth Ratio

1.000
0.995
1.000
1.000
0.740
1.000
0.163

ALL
Adj P
0.965
0.972
0.972
0.994
0.667
0.999
0.079

Small
Adj P
0.023
0.085
0.029
0.985
0.908
0.996
0.757

Large
Adj P
0.204
0.239
0.187
0.211
0.392
0.776
0.836

Adj P

0.989
0.992
0.999
0.765
0.519
0.263
0.057

0.997
0.992
0.994
0.592
0.696
0.158
0.002

Small
Adj P
0.799
0.490
0.619
0.509
0.982
0.613
0.976

Large
Adj P

0.961
0.987
0.974
1.000
1.000
0.922
0.601

ALL
Adj P

0.892
0.930
0.920
1.000
0.914
0.814
0.285

Small
Adj P

1.000
0.993
0.955
0.979
0.598
0.994
0.660

Large
Adj P

Coefficient of Variation

0.034
0.027
0.024
0.010
0.076
0.041
0.310

ALL
Adj P

Coefficient of Variation

0.013
0.026
0.013
0.373
0.989
0.511
0.926

Adj P

Green/Kao B

0.911
0.896
0.907
0.716
1.000
0.180
0.011

Large
Adj P

Green/Kao A

0.996
1.000
1.000
0.770
0.966
0.237
0.664

Small
Adj P

Coefficient of Variation

Coefficient of Variation
ALL
Adj P

Clear/Kao A

Clear/Green

Treatment Comparisons

0.351
0.379
0.297
0.220
0.356
0.542
0.391

Adj P

0.010
0.009
0.007
0.003
0.019
0.015
0.060

Adj P

0.585
0.618
0.617
0.675
0.894
0.081
0.030

Small
Adj P

0.012
0.031
0.009
0.259
0.749
0.407
0.929

Large
Adj P

0.938
0.941
0.955
1.000
0.713
0.883
0.562

ALL
Adj P

0.681
0.752
0.737
0.998
0.382
0.873
0.023

Small
Adj P

0.021
0.107
0.020
0.886
0.906
0.963
0.997

Large
Adj P

Coefficient of Variation

Kao A/Kao B

0.993
0.991
0.983
0.727
0.977
0.132
0.198

ALL
Adj P

Coefficient of Variation

Clear/Kao B

TABLE 2 Experiment 2: log transformed response variables were used in a mixed model ANOVA with a Tukey test for pairwise comparisons
among treatments (29 mg kaolin/L [Kao A], 114 mg kaolin/L [Kao B], greenwater [Geen; 390 mg/m3 chlorophyll-a] or clearwater control) with
the Adjusted P values (Adj P). Log transformed mean coefficient of variation of response variables were compared by mixed model ANOVA with
multiple range test of Tukey with the adjusted p value. This was done for the whole tank (ALL) and then for the smallest half of the tank by
standard length (Small) and again for the largest half of the tank by standard length (Large). Significance noted alpha < 0.05 is given in bold.

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Effect of Contrasting Agents on Survival, Performance, and Condition

17

FIGURE 10 Final mean kurtosis (A and B) and skewness (C and D) of standard length and
tail length ratio of hybrid striped bass in Experiment 2 are plotted by treatments: 29 mg
kaolin/L (Kao A), 114 mg kaolin/L (Kao B), greenwater (Grn 7th; 390 mg/m3 chlorophyll-a)
or clearwater control. Skewness, a unitless measure (Zar 1999), typically ranges from 1 for an
extremely left skewed (i.e., right-tailed) distribution through 0 (symmetrical distribution) to +1
for an extremely right skewed (i.e., left-tailed) distribution. Platykurtic (domed) distribution
has a negative kurtosis; leptokurtic (peaked) distribution has a positive kurtosis. Error bars
represent the standard deviation of the means.

prey consumption and percent feeding data remarkably well (Boehlert &
Morgan 1985). Given the drastically different reflective properties and particle size mixtures of the volcanic ash and sediment samples of Boehlert
and Morgan (1985), the probability that optimal larval response would correlate based upon small particle size seems improbable. The fact that it does,
demonstrates more support for the contrast theory. Further study is needed
to definitively confirm that particle size predominantly acts as a contrasting
agent. Moreover, particle size should be reported in all future larval turbidity
work, in order to speed discovery in this concept.
An optimum turbidity (either clay or algae) should be determined for
each fish species (Shaw et al. 2006). However, there seems to be an optimum range that is fairly similar across species, likely with slight differences
based upon species-specific differences in eye conformation. Graded 14 m
sediment suspensions at 100 mg/L had no negative impact on the survival
of striped bass larvae, but a significant decrease occurred above 500 mg/L

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18

M. McEntire et al.

(Auld & Schubel 1978). The kaolin treatments in the present study demonstrated that the higher level (114 mg/L) is preferable, which is also similar to
the 100 mg/L of sub-1 m sized ceramic clay used in cod (Attramadal et al.
2012). In pacific herring calculated previously the optimum feeding metrics
were 7785 mg/L of sub-5 m particles (Boehlert & Morgan 1985). All of
these optimal ranges peaked at approximately 100 mg/L, which is the highest turbidity level found in nature and is associated with the bottom 12 m
during tidal water exchanges in the Chesapeake Bay (Auld & Schubel 1978).
A good starting point in the determination of the optimum turbidity level of
a new larval fish species should include 100 mg/L of sub-5 m turbidity.
Based upon prior experience with greenwater, the lower kaolin level was
expected to be optimum. At the highest level tested (4X the lower concentration), we expected to see growth inhibition; instead, we found improved
growth. Further work needs to be completed to refine the optimum amount
and frequency of kaolin used in HSB tank larval culture.
Four main factors (decreased visibility, lower activity, reduced transparency of prey, and an increased preference for smaller food items) are
likely responsible for the gain in performance and survival gain of turbid
over clearwater treatments. Turbidity reduces the field of view of a predator
by reducing light and visibility, thus decreasing the number of successful
encounters with food (Duntley 1943; Vinyard & OBrien 1976), but this also
decreases the effectiveness of large piscivores (Chesney 1989; Fiksen et al.
2002; Giske et al. 1994). The culture of larvae in turbid tanks has been suggested to lower larval activity levels, which results in more energy being
partitioned toward somatic growth (Sirois & Dodson 2000). A reduction
in transparency of transparent food items makes prey easier to visualize
(Boehlert & Morgan 1985). An increased preference for smaller feed items
was observed in bluegill, where bright light and turbidity worked in concert
to favor the selection of smaller zooplankton (Giske et al. 1994; Miner &
Stein 1993).
The last factor is an optical illusion of perspective, which is very similar to apparent size prey selection (OBrien et al. 1976), just applied back
toward the fish. Within the spectrum of food particle sizes, from minimum
detectable up to a maximum allowed by mouth gape (Eggers 1977; Keast &
Webb 1966; Shirota 1970; Wong & Ward 1972), an optimum range of food
particle size exists. From the optimum food particle size range at different
distances, an optimum angle of incidence can be computed, similar to the
apparent size of prey selection (OBrien et al. 1976). This implies that fish in
clearwater will select for particles toward the larger apparent size spectrum,
but under highly turbid conditions fish will select for particles at the smaller
apparent size spectrum. A depiction of the short and long fields of view
of a hypothetical larval fish with a 270 degree field of view helps explain
this illusion of perspective (Figure 11). This does not conflict with the threedimensional approach looking at the area of stimulation of the retina by the

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Effect of Contrasting Agents on Survival, Performance, and Condition

19

FIGURE 11 Hypothetical larval fish (270 field of view) depicted with a short (turbid) or long
(clear) field of view. Prey items are rotifers (circles with slash marks) and artemia (jagged
arrows). Optimum prey size range starts to overlap with cohorts, especially when oriented
going away, which leads to more cannibalistic interactions. Turbidity causes encounters to
occur at such close proximity that cohorts are recognized as not being food. The optimum
percent field of view explains why larger prey are preferred over the smaller prey in the
clearwater, while the reverse occurs in turbid.

prey in proportion to the whole area of possible visual stimulation (Aksnes


& Giske 1993; Eggers 1977). Larger itemsgiven that they dominate such a
relatively large percentage of the short visual field in turbid environments
are probably deemed a possible threat or obstacle and not something to be
investigated as food. This could explain the reduced incidence of cannibalism and better condition of the larvae observed in our turbid treatments, as
indicated by their longer tails.

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20

M. McEntire et al.

The implications of applying apparent size theory (OBrien et al. 1976)


back toward the fish under turbid conditions explains the increased preference for smaller food items, the decreased agonistic behavior, and the
improved performance of the smaller fish in the tank relative to the larger
fish. The preference for food size based on an optimum angle of incidence
shows why smaller items might be selected for under turbid conditions and
selected against under clear conditions (Figure 11). Fewer negative interactions among cohorts are explained by the lower probability of encounter,
and the percentage of that field of view taken up by the cohort. From a
bioenergetics standpoint, negative interactions and the associated stress are
energetically expensive and reduce feed intake (Pankhurst & Van der Kraak
1997). The increased preference for smaller food items should allow smaller
fish to catch up to larger fish because smaller fish are consuming more nutrients per body size from each feed item, relative to larger fish in the same
tank. Turbidity significantly reduces visual field and decreases the advantage
larger fish have over smaller fish in relative terms of search volume. Larger
fish will always have a numerically larger search volume, but given the high
turbidity, search volumes become effectively equivalent among large and
small fish. Therefore larger fish will not be able to eat more because of the
turbidity, which divides the food equally among all fish. Under high turbidity
with a declined food presence post feeding, the relative gain in feed consumed per fish body size becomes more important than the search volume
covered. From a bioenergetics standpoint, smaller feed items mean the animal must expend more energy seeking and capturing prey for an equivalent
caloric intake (Knights 1985). High turbidity should result in a right skewed
and peaked size distribution, as was documented in this study.
To growers, the effect of turbidity on fish size distribution might be
equally important to improved survival. Turbidity might have a structuring
effect on a fish community (Blaber & Blaber 1980; Cyrus & Blaber 1987;
Levy & Northcote 1982). This appears to be true within high-density tank
larvae monoculture. High turbidity coupled with the high illumination of this
study resulted in a peaked and right skewed length distributionan ideal
population to transition onto a prepared diet (Figure 10). The body-depth
ratio corroborates this trend. The best body condition fish were in the high
kaolin treatment followed by the greenwater, the low kaolin, and control
treatments (Tables 1 and 2), even though they are not statistically different.
Turbidity inhibits size variation, which should reduce cannibalism. Size
variation is also a primary cause of agonistic behavior, which in turn can
have the same end effect as cannibalism (Hecht & Pienaar 1993). The elevated turbidity appears to diminish the prevalence of this negative social
interaction, as demonstrated with African catfish, Clarias gariepinus (Hecht
& Pienaar 1993). The fish in highly turbid treatments were in better condition, as evidenced by their longer tail and better TLR (Table 2). This also
suggests that cannibalistic interactions are occurring long before they are

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Effect of Contrasting Agents on Survival, Performance, and Condition

21

apparent to growers with declining survival. Growers could use TLR as an


early indicator of the extent or probability of cannibalism. Tail length ratio
appears to be sensitive toward being negatively skewed by increasing algae
levels (Figure 7). This could allow growers to optimize culture conditions
and predict the severity of cannibalism that will be encountered during the
weaning of fish onto artificial diets.
The algae dose confirmation study and replicated kaolin studies both
support the same conclusions regarding the effects of turbidity on variability and size distribution of the resulting fish population. From Experiment
1, the CV of the smallest quartiles appear to drive most of the CVs overall, as indicated by the close proximity of the smallest quartile CV to the
overall CV number (Figures 36). To highlight this, the CV for standard
length of the biggest 75% of fish were plotted against the smallest quartile fish and were less variable. Even though there were three times more
fish represented, there was less variation, which demonstrates how these
size distributions are peaked and skewed. The smallest fish in the tank are
the most variable in length, whereas the largest fish are more homogenous.
Similarly, in Experiment 2, where the CV overall are not different among
the highest turbidity treatments, but the largest half of fish in the turbid
treatments were statistically less variable in the length indices than the clearwater control (Tables 1 and 2). Conversely, the CV of the smallest half of
fish in the turbidity treatments was less variable in body condition than
the clearwater control. In addition, it distinguished between the two kaolin
treatments with the higher treatment being the least variable in condition as
indicated by body depth ratio. The differences of variability among portions
of the population lend direct support for size structuring of fish based upon
the turbidity treatments. The kurtosis and skewness of the treatments were
highly variable, but there was a positive kurtosis for the water treatments
and a negative skewness for standard length and tail length ratio (Figure 10).
This indicated a tank population that was more peaked and right skewed.
Decreasing size variation is possible in tank hybrid striped bass culture by
using water treatments, at least among portions of the population, which
should have implications to the whole tank population.
Based on the morphological measurements, increasing algae concentrations improved growth and condition of fingerlings in both experiments
(Figures 36; Tables 1 and 2). Total lengths of fish were similar to other
reported studies (Bosworth et al. 1997; Denson & Smith 1997; Ludwig 2003;
Ludwig & Lochmann 2009), given allowances for different durations or temperatures of those experiments. The decrease in tail length ratio at the sixth
algae level (Figure 5) along with the increasing skewness (Figure 7) could
signify a change due to an adaptation of the fish to the growing conditions,
which do not require a large tail in reduced visibility. Examining the body
depth ratio (Figure 6) showed an improvement in fish condition with increasing algae levels above the seventh level. A similar numerical improvement

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22

M. McEntire et al.

of fish condition was present in Experiment 2, though not statistically significant (P = 0.067). As would be expected, the longer and better condition fish
in the high turbidity treatments had mean fish weights that were statistically
different (P = 0.050) between both kaolin treatments and the control, with
greenwater being intermediate (Figure 9). It is also worth noting the inhibition that occurred at the highest algae level in Experiment 1 in the metrics
of length, tail length, and TLR, which coupled with the increased variation
of all measured metrics signifies a maximum algae density for this system.
Both experiments demonstrate the same effects of elevated turbidity on fish
growth and condition, despite their different designs.
The main purpose of the studies was to improve fry survival at time of
weaning to an artificial diet. From Experiment 1 the optimum tested range
of greenwater culture for hybrid striped bass using algae paste once daily
was between 629 and 1127 mg/m3 chlorophyll-a (350583 cell/mL, eighth
to ninth algae level tested). In the 1127 mg/m3 chlorophyll-a treatment, the
survival was 104%, but given that the fry were stocked out by volume, that
is typical (5%) of that method of stock-out (Schwarz et al. 2008) and our
previous experience. Daily fluctuation of chlorophyll-a levels indicated a
settling of algae that equated to approximately the next lower cell count
level. The optimal dose might be lower than suggested here if the treatment
was divided into multiple applications daily for a more uniform and constant
color contrast throughout the day.
Partial budget analysis (Kay et al. 2008) is a way of making minor
changes to the overall production budget and extrapolating the economic
benefit. Phase I fingerlings hybrid striped bass cost $86107 per 1,000 fry
(Eklund et al. 2012). From Denson and Smiths (1997) unreplicated study, a
38% mortality could be expected from this studys ending size 1011 mm
to 3555 mm phase I fingerlings, which allows for the inference of the
benefit of using algae paste and kaolin to be extrapolated. The cost of
using 6291127 mg/m3 rate of Nanno 3600TM algae paste ($70/L) in Eklunds
2,457 L production tank would equate to $557928 for a period of 18 days
per tank. Assuming a 38% loss, the final survival would be 5761%, which
equates to an increase revenue $3,5275,127 per tank. The cost of using
dispersed kaolin clay ($14/kg) at the two rates tested under the same conditions as previous would equate to $1.2520.25 per tank and 41%47%
final survival with an increased revenue of $9262,366 per tank. Given the
relatively low cost of the treatments relative to the increased survival, the
investment should be worth the cost. If algae were already produced for
rotifer culture, as can be inferred from the production scheme of Eklund
et al. (2012), the use of greenwater culture to improve survival should be
worth the investment because it would preclude buying the expensive algae
paste. The annual cost table found in Eklund et al. (2012) is perplexing
because of the use of rotifer cyst with no mention of artemia cyst cost,
which impacts the subsequent economic risk study (Engle & Sapkota 2012).

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Effect of Contrasting Agents on Survival, Performance, and Condition

23

Given that production costs are sensitive to survival rate with a 5%11%
reduction in cost for every 5% improvement in survival (Eklund et al. 2012),
the use of turbidity could likely equate to at least a 25%55% reduction in
tank production cost. Further research is needed to confirm the scalability of
water treatments and to confirm the 38% loss associated with weaning the
fish onto a prepared diet.
The use of kaolin appeared to coat tank surfaces, likely aiding in the
removal of organic matter, similar to a study with Atlantic cod (Attramadal
et al. 2012). The charged property of the kaolin had a noticeable impact
on the strength of the bacteria/fungal mat that grew on the bottom and
sides of the tanks and on the detritus. For example, syphon tubing would
not become occluded when syphoning the kaolin treatments but frequently
would become plugged in the control tanks. Since the interactions between
kaolin and detritus are complex and not well understood (Jarvis et al. 2005),
the quantification of the changes in bacterial growth or the resulting kaolinic
floc properties were not measured or quantified. This anecdotal observation
offers interesting future possibilities, from multiple small syphon tube devices
to clean tanks while excluding fry to automated tank cleaners. There may
be a slope of the bottom of the tank that permits the collection of all the
detritus at a central drain for flushing, with no syphoning needed. Future
work should examine the impact of adding kaolin to the culture of fry during
swimbladder inflation. Given kaolins lipophilic tendencies, further work may
be warranted.

SUMMARY
Turbidity is an important tool in the tank culture of cannibalistic fish.
Optimization of algae level or turbidity level is critical to successful tank
larval culture, which might be much higher than expected. Once daily addition of algae paste for greenwater culture produced the optimum survival
of weaning sized fingerlings between 6291127 mg/m3 (350583 cell/mL).
Kaolin worked as well as greenwater culture. Using greenwater or kaolin
improved the length, larval condition, TLR, and uniformity of the fish at the
time of weaning. Tail length ratio shows promise as an early indicator of
cannibalism and aggressive behavior. Increasing turbidity to an optimal level
appears to inhibit growth or decrease the size advantage of the largest fish
in the tank.

ACKNOWLEDGMENTS
We would like to thank Dr. Sidney Fuller, Troy Bader, and Jason Brown for
providing the fry and helping with feeding and sampling. We would like to

24

M. McEntire et al.

thank Dr. Bart Green and Greg ONeal for their help with chlorophyll-a determination. The authors thank Dr. Steven Rawles for valuable comments on the
manuscript. Mention of trade names or commercial products in this article is
solely for the purpose of providing specific information and does not imply
recommendation or endorsement by the U.S. Department of Agriculture.

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