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Effects of dietary carbohydrate sources on growth and body composition of juvenile abalone (Haliotis discus, Reeve).

ABSTRACT A 16-wk feeding trial was conducted to assess the effects of different types of dietary carbohydrate on growth and body composition of juvenile abalone (Haliotis discus). A total of 1,680 abalone were randomly distributed among 24 containers (70 per container) and fed one of eight diets, including seven experimental diets containing different types of carbohydrates, including dextrin (DT), glucose (GC), corn starch (CS), a-cellulose (CL), maltose (MT), sucrose (SC), and wheat flour (WF), and Undaria to compare effect of experimental diets. Water stability of the diets was measured 12, 24, and 48 h after seawater immersion. Water stability of the diets was different over all periods of time and their significant interaction was also observed. Survival was not significantly affected by diet. Weight gain and specific growth rate were greatest in abalone fed on the CL diet. Shell length, width, height, and soft body weight were greatest in abalone fed the CL diet, followed by the WF. DT. SC. CS, GC, and MT diets and Undaria. Weight gain, specific growth rate, crude protein, and crude lipid contents of the soft body of abalone fed on the experimental diets were greater than those fed on Undaria. This study shows that CL is the most effective carbohydrate source for improving growth of this species of juvenile abalone and practically applicable in formulating abalone feed.

KEY WORDS: abalone, Haliotis discus, carbohydrate source, dextrin, glucose, corn starch, a-cellulose, maltose, sucrose, wheat flour

INTRODUCTION

Annual aquaculture production of abalone (Haliotis spp.) in Korea in 2000 was approximately 20 tons and reached 8,982 tons in 2014 (KOSIS 2015). This upward trend is expected to continue, given the high demand for abalone for human consumption. Several studies involving feeding trials have been conducted to investigate dietary nutrient requirements of abalone (Uki et al. 1985, Uki et al. 1986a, 1986b, Mai et al. 1995a, 1995b, Lee et al. 1998b, Zhang et al. 2009). The development of alternatives for fish meal and macroalgae, which are the most expensive components in abalone diets (Lee et al. 1999, Cho 2010, Kim et al. 2016); dietary inclusion of macro--and microalgae (Lee et al. 1998a, Lee et al. 2000); and improved feeding regimes (Cho et al. 2011) have also been investigated. The optimum dietary protein and lipid requirements for abalone were reported to be 25%-35% and 3%-7%, respectively (Mai et al. 1995a, 1995b, Fleming et al. 1996, Bautista-Teruel et al. 2003).

Abalone are marine gastropods and are slow feeders. Their natural diet consists of 40%-50% carbohydrates, and they have various enzymes capable of hydrolyzing complex carbohydrates (Fleming et al. 1996). Mai et al. (1995a) found that two species of abalone, Haliotis tuberculata and Haliotis discus hannai, have a great capacity to use carbohydrates for energy and perhaps for other nutritional purposes. Britz et al. (1994) demonstrated that the gross maintenance energy metabolism of abalone is carbohydrate-based, like many other gastropods (Emersion 1967), given that seaweeds have low fat content and high storage carbohydrate content. Previous studies have indicated that differences in the complexity of carbohydrates influence their digestion and utilization by aquatic animals (Wilson 1994, Cuzon et al. 2000, Lee et al. 2003, Stone et al. 2003, Tan et al. 2006).

Carbohydrates are thus an important nonprotein energy source for abalone, in contrast to fish and crustaceans, which do not use them as effectively. Abalone have high levels of the digestive enzymes protease, amylase, cellulase, and alginase, but low levels of lipases (Emersoin 1967, Gomez-Pinchetti & Garcia-Reina 1993, Britz et al. 1994, Garcia-Esquivel & Feibeck 2006). The donkey's ear abalone (Haliotis asinine) utilizes high levels of carbohydrates more effectively than lipids to support its growth (Thongrod et al. 2003).

Carbohydrates should, therefore, be included in abalone diets at levels that optimize their use of dietary protein for growth. Research on the effects of dietary carbohydrate sources on the growth and body composition of Haliotis discus is, however, very limited. This species is endemic to the waters off Japan and eastern Asia, including the South Korean Jeju Island (Han 1998). In this study, the effects of various dietary carbohydrate sources on the growth and body composition of juveniles of H. discus were investigated.

MATERIALS AND METHODS

Preparation of Abalone und Rearing Conditions of Abalone

Juvenile abalone were purchased from a private hatchery (Deagun Fisheries, Jeju, Korea) and transferred to an abalone farm (Ocean and Fisheries Research Institute, Jeju Special Self-Governing Province, Jeju, Korea). Prior to initiation of the feeding trial, the abalone were acclimated to the experimental conditions for 4 wk and fed on dry Unitaria spp. once daily at a ratio of 2%-3% of total biomass. A total of 1,680 juveniles (mean [+ or -] SE: 1.68 [+ or -] 0.001 g) were randomly distributed into twenty-four 70-L rectangular plastic containers (120 X 36 cm; 70 individuals per container), and 12 containers were randomly placed into each of two 9-ton concrete flow-through tanks with a flow rate of 48.2 L/min/tank. Sand-filtered seawater, at 14.7 - 17.2[degrees]C (mean [+ or -] SD: 16.6[degrees]C [+ or -] 0.52[degrees]C), was supplied throughout the feeding trial. Aeration was supplied in each raceway and the animals were subjected to a natural photoperiod. Abalone were fed, for 16 wk, with the experimental diets once a day at 1,700 h at a quantity to ensure satiation, with a small amount (about 2%-3.5% of their biomass) leftover. Dead individuals were removed daily and the bottoms of the containers were cleaned daily. At the end of the 16-wk feeding trial, abalone were harvested and the group from each container was collectively weighed.

Preparation of the Experimental Diets

Eight diets, including seven experimental ones and the dry Undaria, were prepared in triplicate (Table 1). Each experimental diet included a different carbohydrate source (12%), together with fish meal (28%), corn gluten meal (10%), soybean meal (8%), squid liver oil (0.5%), and soybean oil (0.5%), which served as sources of protein and lipid. The different carbohydrate sources were dextrin (DT). glucose (GC), corn starch (CR), a-cellulose (CL), maltose (MT), sucrose (SC), and wheat flour (WF). Protein and lipid levels in the experimental diets were based on published dietary requirements for abalone (Mai et al. 1995a, 1995b). The dry Undaria was also included as a control.

After the addition of sodium alginate (22%) to each experimental diet, the ingredients were mechanically mixed, and water was added at a ratio of 1:1. A paste was created using an electronic mixer and then shaped into 0.15-cm thick sheets to be hand-cut into 1-[cm.sup.2] flakes. The flakes were dipped in an aqueous solution of 5% Ca[Cl.sub.2] for 1 min, and the excess solution was drained naturally. They were then dried naturally for 2 days and stored at -20[degrees]C until use.

Water stability of the experimental diets was measured 12, 24, and 48 h after seawater immersion and calculated as the percentage of final dry weight to initial dry weight content (Mai et al. 1995a).

Analytical Procedures for Diets and Carcasses

Twenty abalone at the start and 10 abalone from each container at the termination of the feeding trial were sampled and frozen at -40[degrees]C for chemical analysis. Prior to examination, all samples were thawed slightly, and the shell was separated from the soft body tissue. Shell length, shell width, and shell height were measured to a precision of 1.0 mm using a digital caliper (Mitutoyo Corporation, Kawasaki. Japan), and the ratio of soft body weight to total body weight (the soft body weight + the excised shell weight) was calculated to determine a condition index. Specific growth rate (SGR, % body weight gain/day) was calculated using Britz et al. (1996): SGR = [(In (Wf) - ln(Wi))/days of feeding] x 100, where ln(Wf) is final mean weight and ln(Wi) is initial mean weight.

The soft body tissue of the 10 abalone from each container was then homogenized and used for proximate analysis. Crude protein content was determined using the Kjeldahl method (Auto Kjeldahl System, Buchi B-324/435/412, Switzerland); crude lipid content was determined using an ether-extraction method; moisture content was determined by oven-drying at 105[degrees]C for 24 h; and ash content was determined using a muffle furnace at 550[degrees]C for 4 h. All methods were based on AOAC (1990) practices.

Statistical A nalysis

Differences between treatments were tested for significance using one-way analysis of variance (ANOVA) and Duncan's multiple range test (Duncan 1955) in SAS 9.3 (SAS Institute, Cary, NC). Water stability of the experimental diets was tested by ANOVA with repeated measurement designs (Cody & Smith 1991). All percentage data were arcsine-transformed prior to statistical analysis.

RESULTS

Water stability of the diets was significantly (P < 0.0001) different over all periods of time and their significant (P < 0.0001) interaction (diets X time) was also observed (Fig. 1). After 12 h seawater immersion, the highest water stability was observed in the DT diet, followed by the WF diet, Undaria, GC, CS, CL, SC, and MT diets. The highest water stability was also observed in the DT diet after 24 h seawater immersion, followed by the WF and CS diets.

Undaria, CL, GC, SC, and MT diets. After 48-h seawater immersion, the highest water stability was observed in the DT diet, followed by the WF, CS, CL, and GC diets, Undaria, SC, and MT diets.

Abalone survival was consistently greater than 93.8%, but not significantly different (P > 0.05) among the diets (Table 2). Weight gain and SGR of abalone fed the CL diet were significantly greater 0.05) than that those fed the all other diets. In addition, weight gain and SGR of abalone fed the experimental diets were significantly (P< 0.05) greater than that of abalone fed on Undaria.

Shells were largest (in terms of length, width, and height) and soft body weight was greatest in abalone fed the CL diet, followed by the WF, DT, SC, CS, GC, and MT diets, and Undaria (Table 3). Abalone fed the CL, WF, DT, SC, GC, and MT diets, however, had a significantly (P < 0.05) higher ratio of soft body weight to total body weight than abalone fed the CS diet and Undaria.

The moisture content of the soft body of abalone fed the WF diet was significantly (P < 0.05) greater than that of abalone fed the all other diets (Table 4), with the lowest value observed for those fed the CS diet. The crude protein content was significantly (P< 0.05) greater in abalone fed the MT diet than in those fed the all other diets, and in abalone fed the GC, CS, and SC diets than those fed the DT, CL, and WF diets, and Undaria. The crude lipid content of abalone fed the all experimental diets was significantly (P < 0.05) greater than that fed Undaria. The ash content of the soft body of abalone did not, however, differ significantly (P > 0.05) among the diets.

DISCUSSION

Because abalone are slow eaters, water stability of food is a critical factor in growth of the animals, and deterioration of water quality in abalone farms. Higher water stability in the DT and WF diets in 12, 24, and 48 h seawater immersion compared with Undaria and MT diet partially accounted for the good growth of abalone fed the former in this study. Dry matter content of the all diets tended to decrease sharply within 12 h after seawater immersion, and then stabilized. Similarly, dry matter of the formulated diets sharply decreased within 12 h after seawater immersion, and then stabilized throughout 72-h observation (Lee et al. 2016). Knauer et al. (1993) compared the effect of seven binding agents on 24-h water stability of an artificial diet for abalone (Haliotis midae) and found that the best water stability was obtained with a 1:3 agangelatine mixture, which retained ca. 70% of its dry matter after 24 h.

The greater weight gain and SGR of abalone fed the experimental diets, compared with those of abalone fed on Undaria, was consistent with previous findings that a single macroalgae produces a poorer growth performance in abalone than does a formulated diet (Lee 1998, Lee et al. 1998b, 1998a, 1998c, Cho et al. 2006, Garcia-Esquivel & Felbeck 2006, Cho et al. 2008, Garcia-Esquivei & Feibeck 2009, Dang et al. 2011, Myung et al. 2016).

The fact that weight gain and SGR did not differ significantly among carbohydrate sources (except for the CL diet achieving the best weight gain and SGR) was partially consistent with the study of Lee et al. (1998c) showing that weight gain of juvenile abalone (Haliotis discus hannai) was not affected by the type of carbohydrate included in the diet (they tested diets containing 24.2% WF, 20% DT, 20% SC. 10% each of [alpha]- and [beta]-potato starch, 15% [alpha]-potato starch, 20% [alpha]-potato starch, or 25% [alpha]-potato starch). They concluded that abalone can use carbohydrates with various degrees of polymerization (i.e., mono-, di-, and polysaccharide) relatively well. Erasmus et al. (1997) also detected alginate lyase, carboxymethylcellulase, laminarinase, agarase, carragenase in the hepatopancrease of abalone (Haliotis midae) and proposed that bacteria resident in the digestive system of H. midae assisted in the digestion of alginate, laminarin, agarose, carrageenan, and cellulose in diet. Careful consideration must be given in feeding trials, in which CL was used as filler in abalone feed and considered to have no effect on abalone, because abalone fed the CL diet outgrew those fed the other carbohydrate source in this study. Garcia-Esquivel and Felbeck (2006) observed high cellulase activity in the digestive glands of red abalone (Haliotis rufescens) when they were fed formulated diets and kelp (Macrocystis pyrifera). In earlier work, Nakagawa and Nagayama (1988) showed that high activities of xylanase and carboxymethylcellulase were found in the extracts of abalone (Haliotis discus hannai and Haliotis sieboldii) and demonstrated that distribution of poly-saccharidases might be related to the food habit of marine invertebrates, although the levels of activity is highly variable between species and types of cellulose. Cellulase was also found in the gut of abalone (Haliotis gigantea and Haliotis japonica) (Yokoe & Yasumasu 1964). The high cellulose activity in abalone may explain why the dietary inclusion of CL produced the best growth performance in this study.

Walton and Cowey (1982) reported that the effective utilization of dietary carbohydrates in aquatic animals seems to be closely related to the capacity of their digestive and metabolic systems to adapt to different aquatic environments and different quantities and complexities of dietary carbohydrates. Tan et al. (2006) reported that gibel carp (Carassius auratus gibelio) performed better on a CL diet, followed by soluble starch, SC, DT, and GC diets (each of which included 20% of its respective carbohydrate), on the other hand, Chinese longsnout catfish (Leiocassis longirostris Gunther) responded better to a DT diet, followed by SC, CL, soluble starch, and GC diets (each of which included 6% of its respective carbohydrate). They concluded that omnivorous and carnivorous fishes achieved different abilities to use complex carbohydrates. It is also known that an increased amount of fiber stimulates the activity of cellulase and the growth of microflora in the gut of the Tilapia (Oreochromis mossambicus) (Manju & Dhevendaran 2002). Several researchers, however, have reported reduced weight gain in fish fed the diets containing CL (Leary & Lovell 1975, Hilton et al. 1983, Anderson et al. 1984, Lee et al. 2003). Hilton et al. (1983), in particular, showed that CL is a water-insoluble dietary fiber, and that it increases gastric emptying time in rainbow trout (Salmo gairdneri).

Abalone fed the CL diet in this study had the largest shells and the heaviest soft bodies. All morphometric criteria measured, except the ratio of soft body weight to total body weight, appeared to be relatively well reflected by growth rate, and were also consistent with previous studies showing that morphometric measurements of abalone agreed with growth rate (BautistaTeruel et al. 2003, Cho 2010, Myung et al. 2016).

The chemical composition of the soft bodies was affected by the type of carbohydrate in the diet in this study. The greater crude protein and lipid content of abalone fed the formulated diets, relative to abalone fed on Undaria was partially consistent with previous studies showing that dietary nutrient content affects the proximate composition of abalone (Mai et al. 1995a, 1995b, Thongrod et al. 2003, Cho et al. 2008, Gracia-Esquivel & Felbeck 2009, Cho 2010, Kim et al. 2016).

In conclusion, including any type of carbohydrate in the diet produced a better growth performance than did a diet consisting solely of Undaria. The inclusion of CL in the diet was, therefore, the most effective strategy for increasing the growth of abalone, among the various types of carbohydrates tested. This information is practically helpful in formulating abalone feed.

ACKNOWLEDGMENTS

This research was a part of the projected titled "Development of the granule-type artificial diet for larval fish" funded by the Ministry of Oceans and Fisheries, Korea. In addition, this work was supported by a grant from the National Institute of Fisheries Science (R2017021).

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KI WOOK LEE, (1) HYEON JONG KIM, (2) HEE SUNG KIM, (1) DONG GYU CHOI, (2) BOK IL JANG, (2) SUNG HWOAN CHO, (2) * BYEONG-HEE MIN, (3) KYOUNG-DUK KIM (4) AND YANG-ICK JOO (5)

(1) Department of Convergence Study on the Ocean Science and Technology, Korea Maritime and Ocean University, 727 Taejong-ro, Busan 49112, Korea; (2) Division of Marine Bioscience, Korea Maritime and Ocean University, 727 Taejong-ro, Busan 49112, Korea; (3) Busan Marine Fisheries Resources Research Institute, 12 Myeongji Ocean City 7-ro, Gangseo-gu, Busan 46763, Korea; (4) Ac]itafeed Research Center, National Fisheries Research and Development Institute, 2600 Haean-ro, Cheongha-myeon, Gyeongsangbuk-do 37517, Korea; (5) Division of Electrical and Electronics Engineering, Korea Maritime and Ocean University, 727 Taejong-ro, Busan 49112, Korea

* Corresponding author. E-mail: chosunh@kmou.ac.kr

DOI: 10.2983/036.036.0115

Caption: Figure 1. Water stability of the diets at 12, 24, and 48 h after seawater immersion (means of duplicate [+ or -] SE). [ANOVA with repeated design: time (P < 0.0001) and their interaction (diets x time) (P < 0.0001)]. Different letters in each time point indicates difference between diets within each time point.
TABLE 1.
Feed formulation of the experimental diets (%, DM basis).

                                   Experimental diets

                             DT *   GC ([dagger])   CS ([dagger])

Fish meal                     28        28            28
Corn gluten meal              10        10            10
Soybean meal                  8          8             8
Carbohydrate source           12        12            12
Sea tangle                    13        13            13
Squid liver oil              0.5        0.5           0.5
Soybean oil                  0.5        0.5           0.5
Sodium alginate               22        22            22
Mineral premixi)              4          4             4
Vitamin premix                2          2             2
   ([paragraph])

Nutrients (%. DM)

  Dry matter                 79.6      82.0          86.1
  Crude protein              35.6      35.9          35.7
  Crude lipid                4.0        3.8           3.8
  Carbohydrate               44.3      43.5          44.8
  Ash                        16.1      16.8          15.7

                                       Experimental diets

                            CL ([dagger])   MT ([dagger])    SC
                                                           ([dagger])

Fish meal                        28            28            28
Corn gluten meal                 10            10            10
Soybean meal                      8             8             8
Carbohydrate source              12            12            12
Sea tangle                       13            13            13
Squid liver oil                  0.5           0.5           0.5
Soybean oil                      0.5           0.5           0.5
Sodium alginate                  22            22            22
Mineral premixi)                  4             4             4
Vitamin premix ([paragraph])      2             2             2

Nutrients (%. DM)

  Dry matter                    88.6          84.0          82.9
  Crude protein                 35.0          36.3          36.2
  Crude lipid                    3.8           3.9           3.9
  Carbohydrate                  44.5          43.7          43.7
  Ash                           16.7          16.1          16.2

                                   Experimental diets

                             WF ([double dagger])  Undaria

Fish meal                            28
Corn gluten meal                     10
Soybean meal                         8
Carbohydrate source                  12
Sea tangle                           13
Squid liver oil                     0.5
Soybean oil                         0.5
Sodium alginate                      22
Mineral premixi)                     4
Vitamin premix ([paragraph])         2

Nutrients (%. DM)

  Dry matter                        87.4           79.8
  Crude protein                     35.2           27.2
  Crude lipid                       3.7             1.7
  Carbohydrate                      45.8           45.8
  Ash                               15.3           25.3

* DT was purchased from Duksan Pure Chemicals (Ansan-si, Kyungki-do.
Korea).

([dagger]) GC, CS, CL, MT, and SC were purchased from Sigma-Aldrich
Co. LLC (St. Louis. MO).

([double dagger]) WF was supplied by CJ CheilJedang Corp.
(Seoul. Korea).

([section]) Mineral premix contained the following ingredients (g-kg
mix): NaCl. 10; MgS[0.sub.4].7[H.sub.2]O, 150;
Na[H.sub.2]P[O.sub.4].2[H.sub.2]O, 250; K[H.sub.2]P[O.sub.4], 320;
Ca[H.sub.4](P[O.sub.4])2. [H.sub.2]O,200; Ferriccitrate, 25;
ZnS[O.sub.4] 7[H.sub.2]O.4; Ca-lactate, 38.5; CuCl. 0.3;
Al[Cl.sub.3],.6[H.sub.2]O. 0.15; KI[O.sub.3], 0.03;
[Na.sub.2][Se.sub.2][O.sub.3], 0.01: MnS[O.sub.4].[H.sub.2]O. 2;
Co[Cl.sub.2]. 6H,0. 0.1.

([paragraph]) Vitamin premix contained the following amount, which
were diluted in cellulose (g-kg mix): L-ascorbic acid, 200:
[alpha]-tocopheryl acetate, 20; thiamin hydrochloride, 5:
riboflavin. 8:pyridoxine, 2; niacin. 40; Ca-D-pantothenate.
12; myo-inositol, 200; D-biotin, 0.4; folic acid. 1.5: p-amino
benzoic acid, 20; [K.sub.3]. 4; A, 1.5; [D.sub.3], 0.003; choline
chloride, 200: cyanocobalamin, 0.003.

TABLE 2.
Survival (%), weight gain (g/abalone), and SGR (%/dav) of juvenile
abalone fed the experimental diets containing the various sources of
carbohydrate for 16 wk.

Experimental diets   Initial weight (g/abalone)

DT                      1.67 [+ or -] 0.002
GC                      1.68 [+ or -] 0.003
CS                      1.68 [+ or -] 0.003
CL                      1.68 [+ or -] 0.004
MT                      1.68 [+ or -] 0.004
SC                      1.67 [+ or -] 0.001
WF                      1.67 [+ or -] 0.001
Unitaria                 1.68 [+ or -] 0.02

Experimental diets   Final weight (g/abalone)

DT                   4.33 [+ or -] 0.063 (b)
GC                   4.17 [+ or -] 0.126 (bc)
CS                    4.26 [+ or -] 0.044(b)
CL                   4.80 [+ or -] 0.800 (a)
MT                   4.12 [+ or -] 0.061 (bc)
SC                   4.29 [+ or -] 0.059 (b)
WF                   4.35 [+ or -] 0.158 (b)
Unitaria             3.95 [+ or -] 0.031 (c)

Experimental diets        Survival (%)

DT                   94.3 [+ or -] 0.82 (a)
GC                   93.8 [+ or -] 2.65 (a)
CS                   94.3 [+ or -] 1.65 (a)
CL                   94.3 [+ or -] 1.43 (a)
MT                   94.3 [+ or -] 1.43 (a)
SC                   96.2 [+ or -] 1.72 (a)
WF                   95.7 [+ or -] 1.43 (a)
Unitaria             95.7 [+ or -] 0.82 (a)

Experimental diets    Weight gain (g/abalone)

DT                    2.65 [+ or -] 0.062 (b)
GC                   2.49 [+ or -] 0.124 (bc)
CS                    2.58 [+ or -] 0.044 (b)
CL                    3.12 [+ or -] 0.098 (a)
MT                   2.44 [+ or -] 0.058 (bc)
SC                    2.61 [+ or -] 0.059 (b)
WF                    2.68 [+ or -] 0.159 (b)
Unitaria              2.27 [+ or -] 0.033 (c)

Experimental diets         SGR * (%/day)

DT                    0.83 [+ or -] 0.012 (b)
GC                   0.79 [+ or -] 0.025 (bc)
CS                    0.81 [+ or -] 0.009 (b)
CL                    0.91 [+ or -] 0.017 (a)
MT                   0.78 [+ or -] 0.011 (bc)
SC                    0.82 [+ or -] 0.012 (b)
WF                    0.83 [+ or -] 0.032 (b)
Unitaria              0.74 [+ or -] 0.008 (c)

Values (mean [+ or -] SE: n = 3) in the same column sharing the same
superscript letter are not significantly different (P > 0.05).

* SG R (%/day) = [(ln(Wf /ln(Wi))/days of feeding] x 100. where
ln(Wf = natural log of the final mean weight of abalone and ln(Wi) =
natural log of the initial mean weight of abalone.

TABLE 3.
Shell length (mm), shell width (mm), shell height (mm), soft body
weight (g/individual), and the ratio of soft body weight to total
weight of abalone fed the experimental diets containing the various
sources of carbohydrate for 16 wk.

Experimental diets      Shell length (mm)         Shell width (mm)

DT                   34.5 [+ or -] 0.06 (b)    23.7 [+ or -] 0.07 (b)
GC                   33.5 [+ or -] 0.05 (de)   23.2 [+ or -] 0.05 (c)
CS                   33.8 [+ or -] 0.04 (cd)   23.3 [+ or -] 0.04 (c)
CL                   35.3 [+ or -] 0.11 (a)    24.7 [+ or -] 0.10 (a)
MT                   33.3 [+ or -] 0.03 (e)    23.2 [+ or -] 0.08 (c)
SC                   34.0 [+ or -] 0.21 (c)    23.5 [+ or -] 0.03 (b)
WF                   34.6 [+ or -] 0.21 (b)    23.8 [+ or -] 0.12 (b)
Undaria              32.5 [+ or -] 0.17 (f)    22.3 [+ or -] 0.15 (d)

Experimental diets     Shell height (mm)       Soft body weight (g)

DT                   6.8 [+ or -] 0.07 (b)    2.6 [+ or -] 0.07 (b)
GC                   6.4 [+ or -] 0.05 (cd)   2.3 [+ or -] 0.02 (de)
CS                   6.5 [+ or -] 0.05 (c)    2.4 [+ or -] 0.03 (cd)
CL                   7.1 [+ or -] 0.07 (a)    3.0 [+ or -] 0.05 (a)
MT                   6.3 [+ or -] 0.04 (d)    2.2 [+ or -] 0.01 (e)
SC                   6.6 [+ or -] 0.06 (c)    2.5 [+ or -] 0.10 (c)
WF                   6.9 [+ or -] 0.09 (ab)   2.7 [+ or -] 0.06 (b)
Undaria              6.1 [+ or -] 0.08 (e)    2.0 [+ or -] 0.04 (f)

                     Soft body weight/total
Experimental diets           weight

DT                   0.67 [+ or -] 0.007 (a)
GC                   0.65 [+ or -] 0.003 (a)
CS                   0.63 [+ or -] 0.008 (b)
CL                   0.67 [+ or -] 0.004 (a)
MT                   0.65 [+ or -] 0.009 (a)
SC                   0.67 [+ or -] 0.015 (a)
WF                   0.67 [+ or -] 0.004 (a)
Undaria              0.62 [+ or -] 0.005 (b)

Values (mean [+ or -] SE; n = 3) in the same column sharing the same
superscript letter are not significantly different (P > 0.05).

TABLE 4.
Chemical composition (%) of the soft body of abalone fed the
experimental diets containing the various sources of carbohydrate for
16 wk.

Experimental diets          Moisture               Crude protein

DT                   28.4 [+ or -] 0.07 (bc)   23.8 [+ or -] 0.05 (c)
GC                   28.5 [+ or -] 0.06 (b)    24.3 [+ or -] 0.04 (b)
CS                   28.3 [+ or -] 0.02 (c)    24.2 [+ or -] 0.02 (b)
CL                   28.5 [+ or -] 0.04 (b)    24.0 [+ or -] 0.07 (c)
MT                   28.5 [+ or -] 0.04 (b)    24.6 [+ or -] 0.06 (a)
SC                   28.5 [+ or -] 0.05 (b)    24.3 [+ or -] 0.05 (b)
WF                   28.7 [+ or -] 0.05 (a)    23.9 [+ or -] 0.07 (c)
Undaria              28.4 [+ or -] 0.06 (bc)   22.9 [+ or -] 0.08 (d)

Experimental diets        Crude lipid                 Ash

DT                   1.6 [+ or -] 0.07 (a)   3.7 [+ or -] 0.07 (a)
GC                   1.6 [+ or -] 0.04 (a)   3.7 [+ or -] 0.04 (a)
CS                   1.7 [+ or -] 0.03 (a)   3.7 [+ or -] 0.03 (a)
CL                   1.6 [+ or -] 0.05 (a)   3.6 [+ or -] 0.06 (a)
MT                   1.6 [+ or -] 0.06 (a)   3.7 [+ or -] 0.02 (a)
SC                   1.5 [+ or -] 0.04 (a)   3.7 [+ or -] 0.08 (a)
WF                   1.5 [+ or -] 0.05 (a)   3.7 [+ or -] 0.03 (a)
Undaria              1.0 [+ or -] 0.02 (b)   3.7 [+ or -] 0.06 (a)

Values (mean [+ or -] SE; n = 3) in the same column sharing the same
superscript letter are not significantly different (P > 0.05).
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Article Details
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Author:Lee, Ki Wook; Kim, Hyeon Jong; Kim, Hee Sung; Choi, Dong Gyu; Jang, Bok Il; Cho, Sung Hwoan; Min, By
Publication:Journal of Shellfish Research
Article Type:Report
Date:Apr 1, 2017
Words:5876
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