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Growth and energy utilization of juvenile pink abalone Haliotis corrugata fed diets containing different levels of protein and two starch:lipid ratios.

ABSTRACT Juvenile pink abalone abalone (ăbəlō`nē), popular name in the United States for a univalve gastropod mollusk of the genus Haliotis, members of which are also called ear shells, or sea ears, as their shape resembles the human ear.  Haliotis corrugata (initial mean length = 10.7 [+ or -] 0.3 mm; initial mean weight = 0.15 g) were grown under laboratory conditions in an aerated aer·ate  
tr.v. aer·at·ed, aer·at·ing, aer·ates
1. To supply with air or expose to the circulation of air: aerate soil.

 flow-through seawater seawater

Water that makes up the oceans and seas. Seawater is a complex mixture of 96.5% water, 2.5% salts, and small amounts of other substances. Much of the world's magnesium is recovered from seawater, as are large quantities of bromine.
 system at 21 [+ or -] 1[degrees]C. For 131 days, abalone were fed diets containing three different levels of protein (42%, 36% and 32%), each level containing two ranges of starch starch, white, odorless, tasteless, carbohydrate powder. It plays a vital role in the biochemistry of both plants and animals and has important commercial uses. :lipid lipid

Any of a diverse class of organic compounds, found in all living things, that are greasy and insoluble in water. One of the three large classes of substances in foods and living cells, lipids contain more than twice as much energy (calories) per unit of weight as the
 ratios (1.5-1.8 or 3.2-3.6). The gross energy content of the diets ranged from 4.26-3.68 kcal/g. Within a particular protein level, growth did not differ significantly. When dietary protein levels exceeded 32%, a trend of higher growth of abalone was observed for dietary treatments that contained lower levels of lipid (higher starch to lipid ratios). The amount of protein consumed daily for the 32% dietary protein is apparently insufficient to meet requirements for energy and maximum growth. No notable diet-dependent differences in the proportional composition (dry weight) of the shell and the soft tissue, and the energy content of the soft tissue were observed. Daily food intake and consumed energy were significantly different among treatments, being inversely in·verse  
1. Reversed in order, nature, or effect.

2. Mathematics Of or relating to an inverse or an inverse function.

3. Archaic Turned upside down; inverted.

 related to the calculated P:E ratios. The increased consumption may have been somewhat overestimated because of loss arising from lower water stability of the diets. Ammonia ammonia, chemical compound, NH3, colorless gas that is about one half as dense as air at ordinary temperatures and pressures. It has a characteristic pungent, penetrating odor.  excretion excretion, process of eliminating from an organism waste products of metabolism and other materials that are of no use. It is an essential process in all forms of life. In one-celled organisms wastes are discharged through the surface of the cell.  ranged from 7.9-4.8 [micro]g N[H.sub.4.sup.+]/h/g abalone but was not significantly different among treatments except for the treatment containing 32% crude protein and a low carbohydrate carbohydrate, any member of a large class of chemical compounds that includes sugars, starches, cellulose, and related compounds. These compounds are produced naturally by green plants from carbon dioxide and water (see photosynthesis). :lipid ratio. Specific Dynamic Action (SDA SDA
specific dynamic action

Serotonin dopamine antagonist (SDA)
The newer second-generation antipsychotic drugs, also called atypical antipsychotics.
) comprised nearly 50% of measured oxygen consumption and did not significantly differ among treatments. Respiration respiration, process by which an organism exchanges gases with its environment. The term now refers to the overall process by which oxygen is abstracted from air and is transported to the cells for the oxidation of organic molecules while carbon dioxide (CO  increased during the night showing a typical circadian circadian /cir·ca·di·an/ (ser-ka´de-an) denoting a 24-hour period; see under rhythm.

Relating to biological variations or rhythms with a cycle of about 24 hours.
 pattern reported in Haliotis genera genera, in taxonomy: see classification. . Measured mucus mucus /mu·cus/ (mu´kus) the free slime of the mucous membranes, composed of secretion of the glands, various salts, desquamated cells, and leukocytes.

 production did not vary among treatments. Carbohydrate preferentially serves as the principal energy source and a consistent amount of dietary protein is also used as an energy source, regardless of dietary protein content. Within a developed energy budget, approximately 70% of the ingested in·gest  
tr.v. in·gest·ed, in·gest·ing, in·gests
1. To take into the body by the mouth for digestion or absorption. See Synonyms at eat.

 energy was lost through feces feces
 or excrement or stools

Solid bodily waste discharged from the colon through the anus during defecation. Normal feces are 75% water. The rest is about 30% dead bacteria, 30% indigestible food matter, 10–20% cholesterol and other fats,
, and approximately 25% of the ingested energy was metabolized, with 7% to 10% channeled to growth. A direct determination of available digestible digestible

having the quality of being able to be digested.

digestible energy
the proportion of the potential energy in a feed which is in fact digested.

digestible protein
see digestible protein.
 energy was not possible because of the inability to collect sufficient feces to estimate fecal fecal /fe·cal/ (fe´k'l) pertaining to or of the nature of feces.

Relating to or composed of feces.


pertaining to or of the nature of feces.
 energy. The results suggest that practical diets should include levels of dietary protein that are approximately 35% protein to meet requirements for energy and maximum growth. In addition, carbohydrates Carbohydrates
Compounds, such as cellulose, sugar, and starch, that contain only carbon, hydrogen, and oxygen, and are a major part of the diets of people and other animals.

Mentioned in: Laxatives

 seem to be the preferred energy source and lipid levels should be minimized to the extent that essential fatty acids Essential fatty acids
Sources of fat in the diet, including omega-3 and omega-6 fatty acids.

Mentioned in: Nutritional Supplements
 requirements are met.

KEY WORDS: abalone, starch:lipid ratio; energy utilization, Haliotis corrugata


Abalone aquaculture aquaculture, the raising and harvesting of fresh- and saltwater plants and animals. The most economically important form of aquaculture is fish farming, an industry that accounts for an ever increasing share of world fisheries production.  is a growing commercial enterprise and nutritionally balanced, formulated feed is becoming increasingly important as a substitute for their natural diet of macroalgae. The need for an effective formulated feed is based on not only achieving better growth performances but also avoiding the unpredictable availability of macroalgae. Production of commercial size abalone can take several years, because their nutritional requirements nutritional requirements,
n the food and liquids necessary for normal physiologic function.
 and physiology are yet to be fully understood. Thus, research on abalone nutritional requirements is important to increase efficiency through a combination of higher growth rates Growth Rates

The compounded annualized rate of growth of a company's revenues, earnings, dividends, or other figures.

Remember, historically high growth rates don't always mean a high rate of growth looking into the future.
 and lower costs.

Most animals consume their food to fulfill their energy requirements, and abalone is not an exemption (Gomez-Montes et al. 2003); it has been reported that abalone grow better when fed a formulated diet containing a protein:energy (P:E) ratio of approximately 100.

H. fulgens requires between 59-67 cal/g abalone/day to achieve maximum growth, at a crude protein intake of 6.22 [+ or -] 0.17 mg/g abalone/day with a protein efficiency ratio Protein efficiency ratio (PER) is based on the weight gain of a test subject divided by its intake of a particular food protein during the test period.

From 1919 until very recently, the PER had been a widely used method for evaluating the quality of protein in food.
 (PER) of 3.41 (Gomez-Montes el al. 2003). At higher intakes of protein, PER decreases significantly. For the best dietary P:E ratio, energy derived from nonprotein sources was satisfactory to minimize the use of protein as an energy source. For abalone feeding on a diet with a P:E ratio of 108, more dietary protein was used as a source of energy.

To achieve maximum growth, the deposition of dietary protein in the muscle tissue in abalone must be maximized and a proper balance of appropriate sources of dietary protein and energy will help to achieve this objective. Research conducted with different species of fish has consistently demonstrated an insufficient level of dietary protein yields lower growth rates (Smith 1989), and if the dietary level of energy in the diet is insufficient, then protein is used as a source of energy for maintenance (NRC NRC
1. National Research Council

2. Nuclear Regulatory Commission

Noun 1. NRC - an independent federal agency created in 1974 to license and regulate nuclear power plants
 1983). A diet with the most effective P:E ratio will allow for a reduction in the level of dietary protein without a corresponding reduction in growth.

A modification of the relative proportions of dietary carbohydrate and lipid sources of energy may contribute to an even greater growth response. The dietary lipid requirement of abalone is believed to be low (Mai et al. 1995, Durazo-Beltran et al. 2003); therefore, carbohydrates are probably the preferred source of energy (Knauer et al. 1996, Monje & Viana 1998). A possible reduction in dietary lipid in response to the efficient use of carbohydrates as energy sources must be approached carefully to ensure that satisfaction of the essential fatty acid
    Essential fatty acids, or EFAs, are fatty acids that cannot be constructed within an organism from other components (generally all references are to humans) by any known chemical pathways; and therefore must be obtained from the diet.
     requirements is not compromised. A wide range of dietary starch levels seem to have no adverse effect on the growth of abalone, and cornstarch cornstarch, material made by pulverizing the ground, dried residue of corn grains after preparatory soaking and the removal of the embryo and the outer covering. It is used as laundry starch, in sizing paper, in making adhesives, and in cooking.  is considered to be a good carbohydrate source to balance the energy content of experimental diets (Knauer et al. 1996). Therefore, levels of carbohydrates can be changed to produce different dietary P:E ratios.

    The objective of this work was to understand how dietary energy is used when pink abalone Haliotis corrugata are fed diets containing different dietary starch:lipid ratios and different protein levels.


    Diet Preparation

    Six experimental diets were formulated (Table 1) to contain 3 different protein levels (42%, 36% and 32%) and two different starch:lipid ratio levels, ranging from either 1.5-1.8 and from 3.2-3.6 whereas also trying to maintain the P:E ratios (mg protein: kcal) between 90 and 100. Primary sources of protein were menhaden menhaden: see herring.
     or pogy

    Any of several species of Atlantic coastal fishes (genus Brevoortia of the herring family), used for oil, fish meal (mainly for animal feed), and fertilizer.
     fish meal (EWOS EWOS - European Workshop for Open Systems ) and isolated soybean protein isolated soybean protein

    the pure protein that can be spun into a fiber and then, if it is colored and flavored, has a very close resemblance to meat.
     (92% CP), ingredients were maintained at a 2:1 ratio to ensure a common dietary amino acid amino acid (əmē`nō), any one of a class of simple organic compounds containing carbon, hydrogen, oxygen, nitrogen, and in certain cases sulfur. These compounds are the building blocks of proteins.  profile that generally reflected the proportional composition of soft tissue for abalone (Fleming et al. 1996). Because the dietary protein level was reduced, cornstarch was added to maintain the desired P:E ratio of the diets. Kelp kelp: see seaweed; Phaeophyta.

    Any of about 30 genera of large seaweeds that make up the order Laminariales (brown algae), found in colder seas.
     meal was used as a filler fill·er 1  
    One that fills, as:
    a. Something added to augment weight or size or fill space.

    b. A composition, especially a semisolid that hardens on drying, used to fill pores, cracks, or holes in wood, plaster,
     as needed as needed prn. See prn order. . The structural carbohydrates that constitute a high proportion of this ingredient were not included in the calculations of the starch to lipid ratios, because it is unknown whether these specific carbohydrates can be efficiently digested.

    Each dietary treatment contained cod liver oil cod liver oil

    an oil pressed from the fresh liver of the cod and purified. It is one of the best-known natural sources of vitamin D, and a rich source of vitamin A. Because cod liver oil is more easily absorbed than other oils, it was formerly widely used as a nutrient and tonic,
    , which was added proportionally (from 6.3% to 3.1%) for each treatment. The composition and amounts of vitamin and mineral premixes that were added followed the recommendations of Hahn (1989). All ingredients were mixed thoroughly to produce a homogeneous mixture; water was added to achieve a moisture content of approximately 60%. The dough-like diets were cold extruded through a pasta machine (Rosito Bisani, Los Angeles Los Angeles (lôs ăn`jələs, lŏs, ăn`jəlēz'), city (1990 pop. 3,485,398), seat of Los Angeles co., S Calif.; inc. 1850. ) into 0.5 x 0.7 mm pellets, and then stored at -80[degrees]C until used. Water stability of each of the diets was determined in triplicate before and during measurements of intake by immersing 10 pellets of each diet within seawater held in control buckets (no animals inside). After 12 h of immersion immersion /im·mer·sion/ (i-mer´zhun)
    1. the plunging of a body into a liquid.

    2. the use of the microscope with the object and object glass both covered with a liquid.
    , the amount of remaining feed (dry matter stability) was determined.

    Proximate Analysis (Chem.) an analysis which determines the proximate principles of any substance, as contrasted with an ultimate analysis.

    See also: Proximate
     of Diets and Tissue

    Proximate proximate /prox·i·mate/ (prok´si-mit) immediate or nearest.

    Closely related in space, time, or order; very near; proximal.


    immediate; nearest.
     composition of diets and tissue was determined according to according to
    1. As stated or indicated by; on the authority of: according to historians.

    2. In keeping with: according to instructions.

     standard procedures (AOAC AOAC Association of Official Analytical Chemists (now AOAC International)
    AOAC Association of Analytical Communities
    AOAC Association of Analytical Chemists
    AOAC Always On/Always Connected
    AOAC Aero-Optic Evaluation Center
     1990). For diets, moisture content of each diet was calculated from triplicate samples (4-5g) that were dried to constant weight at 60[degrees]C. Mean total nitrogen content was determined by the microKjeldahl method, and percent crude protein was then calculated as % N x 6.25. Mean total dietary crude lipid was determined gravimetrically according to the method of Bligh & Dyer (1959). Fiber was determined by the acid-detergent method described in AOAC (1990). Mean ash content was determined by heating samples to 550[degrees]C for 4 h. The gross energy contents of diets and organisms (tissue and mucus) were determined by direct combustion in an adiabatic ad·i·a·bat·ic  
    Of, relating to, or being a reversible thermodynamic process that occurs without gain or loss of heat and without a change in entropy.
     calorimeter calorimeter: see calorimetry.

    Device for measuring heat produced during a mechanical, electrical, or chemical reaction and for calculating the heat capacity of materials.
     Parr 1281. Nitrogen free extract was calculated by difference (NFE NFE

    nitrogen free extract.
     = 100 - [% crude protein + % crude lipid + % ash + % crude fiber]). Samples of the soft tissue of the abalone from each experimental unit were collected at the end of the experiment, frozen individually at -80[degrees]C and subsequently submitted to the same analysis.

    Experimental Procedure

    Three weeks prior to the initiation of the experiment, juvenile pink abalone H. corrugata were fed a diet consisting of diatoms diatoms

    a series of unicellular algae, microscopic in size, with cell walls containing silica. Members of the family Diatomaceae. Their remains accumulate as geological deposits and are mined. See diatomaceous earth.
     (Navicula incerta and Nizchia closterium). Thereafter, for 1 week, a standard, nutritionally balanced formulated diet, containing fish meal, soybean soybean, soya bean, or soy pea, leguminous plant (Glycine max, G. soja, or Soja max) of the family Leguminosae (pulse family), native to tropical and warm temperate regions of Asia, where it has been  meal, corn meal and kelp meal as the main ingredients (P:E ratio of 88) was fed as recommended by Durazo-Beltran et al. (2004). A total of 432 abalone with an average initial shell length and total weight of 10.70 [+ or -] 0.32 mm and 0.164 [+ or -] 0.01 g, respectively, were selected for use in the growth trial. Each experimental unit was a growing chamber that consisted of an ABS (Automatic Backup System) See backup program.  (acrylonitrile-butadiene-styrene) black pipe tube holding a container fitted with a plastic mesh floor (1 x 1 mm), and located at 3 cm from the pipe bottom. A total of 23 abalone were placed into each chamber that was held within a 4-L black bucket that was part of a flow-through system (250 mL/min). Each chamber was supplied with an air stone to maintain satisfactory levels of oxygen. Water temperature was maintained at 20[degrees]C [+ or -] 1[degrees]C throughout the feeding trial by a heat pump heat pump: see air conditioning.
    heat pump

    Device for transferring heat from a substance or space at one temperature to another at a higher temperature.
     connected to an insulated in·su·late  
    tr.v. in·su·lat·ed, in·su·lat·ing, in·su·lates
    1. To cause to be in a detached or isolated position. See Synonyms at isolate.

     700-L reservoir, from which water was distributed to the growing chambers. Each treatment was run in triplicate for a total of 18 experimental units; treatments were randomly assigned to chambers in the culture system. A photoperiod photoperiod /pho·to·pe·ri·od/ (fo´to-per?e-od) the period of time per day that an organism is exposed to daylight (or to artificial light).photoperiod´ic

     of 12L/12D was maintained throughout the experiment, and diets were fed in excess every night at 20:00 hrs. On the following day at 08:00 h, any unconsumed diet was removed. Total feed intake (F) of each of the triplicate groups of abalone for each treatment was calculated twice during the last 8 days of each month. Daily dry matter loss from pellets in control chambers without abalone (stability) was used to adjust the feed intake that was estimated as:

    F = (G [S/100]) - R (1)

    where G represents the amount of feed offered, S is the amount of feed recovered from the control buckets, and R is the feed remaining in the containers with the experimental abalone.

    The mean daily rate of feed intake for each treatment was then calculated as a percentage of body weight. Every 4 weeks, length and weight of all abalone within each replicate of each treatment were recorded as determined with an electronic digital caliper caliper

    Instrument that consists of two adjustable legs or jaws for measuring the dimensions of material parts. Spring calipers have an adjusting screw and nut; firm-joint calipers use friction at the joint to hold the legs unmoving.
     ([+ or -] 0.01 [micro]m) and an electronic scale (0.01 g error), respectively. The experiment was terminated after 131 days.

    The following nutritional indices were calculated using the feed intake data.

    Feed conversion efficiency,

    FCE FCE First Certificate in English
    FCE Final Cut Express (Apple video editing suite)
    FCE Facultad de Ciencias Económicas (Spanish)
    FCE Functional Capacity Evaluation
    FCE Florida Coastal Everglades
     = 100 (wet weight gain [g])/(feed intake [g]) (2)

    Protein efficiency ratio,

    PER = (wet weight [g] gain)/(protein intake [g]) (3)

    Specific growth rate (SGR SGR Sustainable Growth Rate
    SGR Societa' di Gestione del Risparmio (Italian: Investment Management Company)
    SGR Specific Growth Rate
    SGR Surgeon General's Report
    SGR Soft Gamma-ray Repeater
    , %/day) was estimated according to Hopkins (1992)

    SGR = (ln [[w.sub.t]] - ln [[w.sub.i]])/t * 100 (4)

    where ln ([w.sub.t]) is the natural logarithm Natural logarithm

    Logarithm to the base e (approximately 2.7183).
     of final body weight, ln ([w.sub.i]) is the natural logarithm of the initial body weight and t is the time in days.

    Oxygen Consumption

    Oxygen consumption was recorded using two computer-controlled polarographic po·lar·og·ra·phy  
    An electrochemical method of quantitative or qualitative analysis based on the relationship between an increasing current passing through the solution being analyzed and the increasing voltage used to produce the current.
     oxygen sensors An oxygen sensor is an electronic device that measures the proportion of oxygen (O2) in the gas or liquid being analyzed. It was developed by Robert Bosch GmbH during the late 1960s under supervision by Dr. Günter Bauman.  (Strathkelvin Instruments, Ireland), each with a capacity of six channels. Prior to use, the sensors were calibrated cal·i·brate  
    tr.v. cal·i·brat·ed, cal·i·brat·ing, cal·i·brates
    1. To check, adjust, or determine by comparison with a standard (the graduations of a quantitative measuring instrument):
     with a 0% oxygen solution (2% sodium sulfite sodium sulfite
    A white crystalline or powdered compound, Na2SO3, used in preserving foods, silvering mirrors, developing photographs, and making dyes.
     in 0.01M sodium borate sodium borate
    A crystalline compound that is the sodium salt of boric acid and is used as an alkalizing agent and as a mild astringent in lotions, gargles, and mouthwashes.
     as buffer) and 100% saturated oxygen water. The sensors were held in the solution/water (21 [+ or -] 1[degrees]C) until variations were no longer recorded.

    A block design with time (3 days) was used. The juvenile pink abalones were first acclimatized in smaller growing chambers for 20 h. The respective experimental diets were fed during the night (20:00 h), and during the following morning, the chambers were cleaned of any organic material that remained. At approximately noon of the day of an evaluation, the water flow of one unit of each treatment was closed and the chamber sealed for an incubation period incubation period
    1. See latent period.

    2. See incubative stage.

    Incubation period 
     of 50 min. The chamber was then opened for 10 min to restore the oxygen levels. This procedure was repeated 4 times. Two chambers were used as controls to determine the amount of oxygen consumed in the absence of abalone. The mean oxygen consumed in the control chambers was subtracted from the oxygen consumed by the experimental abalone. After each measurement, the organisms were then deprived of feed for 3 days and following the previously described procedure, oxygen consumption was measured to calculate the amount of energy used for maintenance (basal metabolism basal metabolism: see metabolism. ).

    After termination of the incubation period, all abalone were removed from the chambers, blot dried with a piece of cloth Noun 1. piece of cloth - a separate part consisting of fabric
    piece of material

    bib - top part of an apron; covering the chest

    chamois cloth - a piece of chamois used for washing windows or cars
    , measured with digital calipers (MAX-CAL, [+ or -] 0.03 mm) at their longest dimension, and weighed using an analytic scale (AND SV-200, [+ or -] 0.01 g). Oxygen consumption rate (metabolic rate Noun 1. metabolic rate - rate of metabolism; the amount of energy expended in a give period
    basal metabolic rate, BMR - the rate at which heat is produced by an individual in a resting state
    , V[O.sub.2]) was estimated from the corrected slope of the oxygen evolution curve (chambers with abalone minus chambers without abalone), after converting the % [O.sub.2] saturation to [micro]mol of dissolved [O.sub.2] in seawater, from known values of oxygen solubility solubility

    Degree to which a substance dissolves in a solvent to make a solution (usually expressed as grams of solute per litre of solvent). Solubility of one fluid (liquid or gas) in another may be complete (totally miscible; e.g.
     (Green & Carritt 1967). The following equation was used for calculating metabolic rate:

    V[O.sub.2e] = (Cs * m * 60)/(100% * n) (5)

    where: V[O.sub.2e] = metabolic rate of the experimental organism (mL [O.sub.2]/g org/h); m = slope of the 02 evolution curve (%[O.sub.2]/min); 60 = factor used to transform from minutes to hours; Cs = Total amount of [O.sub.2] in the incubation incubation /in·cu·ba·tion/ (in?ku-ba´shun)
    1. the provision of proper conditions for growth and development, as for bacterial or tissue cultures.

     chamber at 100% saturation ([micro]L [O.sub.2]); n = Number of organisms in the incubation chamber.

    To determine the rate of oxygen consumption over a 24 h period, 80 organisms were randomly selected and equally divided among 4 chambers (20 per chamber). Oxygen consumption was measured continuously during successive 50 min periods in a closed system (static) followed by 10 min with open flow to recover the oxygen levels, as previously described. Four incubation chambers containing no abalone were used as replicate controls to determine the amount of oxygen that was not consumed by the abalone.

    Ammonia Excretion and Pedal Mucus Production

    Ammonia excretion was determined by introducing the abalone into 800-mL beakers containing Millipore-filtered and air-saturated seawater. To eliminate any nitrogen contamination, all glassware and beakers used were soaked soak  
    v. soaked, soak·ing, soaks
    a. To make thoroughly wet or saturated by or as if by placing in liquid.

    b. To immerse in liquid for a period of time.

     with ammonia free detergent detergent (dētûr`jənt, dĭ–), substance that aids in the removal of dirt. Detergents act mainly on the oily films that trap dirt particles.  (alconox), then 5% hydrochloric acid hydrochloric acid: see hydrogen chloride.
    hydrochloric acid
     or muriatic acid

    Solution in water of hydrogen chloride (HCl), a gaseous inorganic compound.
    , rinsed with distilled water Noun 1. distilled water - water that has been purified by distillation
    H2O, water - binary compound that occurs at room temperature as a clear colorless odorless tasteless liquid; freezes into ice below 0 degrees centigrade and boils above 100 degrees centigrade;
    , and stored in an N-free environment. Beakers were closed with a cover to seal the water and reduce the risk of N contamination during the incubation time. Control beakers, containing seawater but no animals, were incubated at the same time under the same conditions (20[degrees]C). Rates of ammonia excretion were determined as the ammonia content in the beakers with abalone minus the content in the control beakers after incubation at 30 and 60 min. Water samples were removed from the containers and kept at -80[degrees]C until processed in an automatic analyzer (SKALAR) by the Berthelot reaction and read at 640 nm. Rates of excretion were expressed as [micro]g N[H.sub.4.sup.+]/h/g abalone and converted to energy equivalents, using the conversion factor of 0.349 J/[micro]mol N-N[H.sub.4] (Widdows & Salkeld 1993).

    To calculate the pedal mucus, abalone were placed in clean, dry 800-mL beakers that had been previously weighed. After 24 h, abalone were carefully removed and the feces in the beakers were washed away. The beakers were then dried and weighed, and the difference in weight estimated as mucus production (dry weight).

    Statistical Analysis

    To determine whether mean growth, expressed as final body weight (log transformed), fed the different diets were significantly different, a 2-way analysis of variance, 6 diets x 2 periods of 30 day each, was conducted. Diet and interaction were estimated using orthogonal At right angles. The term is used to describe electronic signals that appear at 90 degree angles to each other. It is also widely used to describe conditions that are contradictory, or opposite, rather than in parallel or in sync with each other.  contrasts. The daily feed intake based on measurements during the designated 16 days was calculated and compared among treatments using a 2-way (period and treatment) analysis of variance and multiple comparisons between least square means were performed using Tukey test when treatment showed a P < 0.05. Daily feed intake was used to estimate the amount of crude protein or calories consumed per g abalone. The calculated data was then analyzed as previously stated. FCE and PER of abalone of each treatment were calculated using the daily growth increments and the corresponding feed intake for the 16-day period, dry matter and protein content, respectively. All statistical analyses were performed using SAS-GLM procedures (SAS (1) (SAS Institute Inc., Cary, NC, A software company that specializes in data warehousing and decision support software based on the SAS System. Founded in 1976, SAS is one of the world's largest privately held software companies. See SAS System.  8.2, 2001).


    The analytically determined proximate composition and caloric caloric /ca·lo·ric/ (kah-lor´ik) pertaining to heat or to calories.

    1. Of or relating to calories.

    2. Of or relating to heat.
     content of the diets generally corresponded to the intended compositions and levels based on the formulations (Table 1). Mean stability of the diets representing the different treatments ranged from 88.4% to 77.6% and decreased with increasing levels of the kelp meal ingredient. Diets containing the highest levels of kelp meal had stabilities that were significantly less (P < 0.001) than those of all the other diets (Table 2). Feed intake, expressed as a percentage of body weight, was significantly greater for treatments [T.sub.6] (2.15%) and [T.sub.3] (2.21%) compared with treatments [T.sub.5] and [T.sub.2], (1.96%) and lowest for treatments [T.sub.4] and [T.sub.1] (1.64% and 1.67%, respectively). These values also showed an inverse (mathematics) inverse - Given a function, f : D -> C, a function g : C -> D is called a left inverse for f if for all d in D, g (f d) = d and a right inverse if, for all c in C, f (g c) = c and an inverse if both conditions hold.  trend relative to diet stability. Feed intake, expressed as calories/g abalone ranged between 84-70 and was inversely related to the stability of the diets. Feed intake, expressed as crude protein/g abalone, was not significantly different among treatments (Table 2), ranging from 7.0-7.4 mg CP/g abalone.

    After 131 days (Table 2), within each of the two ranges of starch protein ratios, weight gain decreased as the P:E ratio decreased. For those diets with the highest protein levels, but the differences were not significant. Abalone fed the diets containing the lowest levels of protein and corresponding lowest P:E ratios had the lowest growth rates, significantly lower for those diets containing the high starch:lipid ratio. A similar response was observed for SGRs. The greatest FCEs were observed for abalone from treatments [T.sub.4] and [T.sub.1] (63.8 and 60.6, respectively). FCEs decreased as the water stability of the diets decreased. The somewhat lower levels of dietary fiber dietary fiber
    Coarse, indigestible plant matter, consisting primarily of polysaccharides, that when eaten stimulates intestinal peristalsis.
     in treatments [T.sub.4] and [T.sub.1] may have contributed to the higher FCEs.

    Highest PERs were associated with the diets that contained the highest levels of protein and the highest water stability. The PER for abalone from treatment [T.sub.4] (1.66) was significantly higher than that of abalone from each of treatments [T.sub.3] and [T.sub.2] (1.24 and 1.29, respectively) but not significantly different from those of abalone from the remaining treatments (Table 2). Generally, the proportional composition of shell/tissue of abalone from the different dietary among treatments was very similar except for abalone in treatment [T.sub.2] with a relatively higher and significant tissue proportion (Table 3).

    No differences in rates of oxygen consumption of abalone were observed among dietary treatments with values ranging from 94.3-77.02 [micro]L [O.sub.2]/h/g abalone. In addition, the specific dynamic action (SDA) represented as the oxygen used for abalone to perform digestion for all dietary treatments was not significantly different and represented 52% to 67% (25.7-45.01 [micro]L [O.sub.2]/h/g abalone; calculated from Table 4) of the rate of total oxygen consumed.

    Ammonia production Because of its many uses, ammonia is one of the most highly-produced inorganic chemicals. There are literally dozens of large-scale ammonia production plants worldwide. The worldwide production in 2004 was 109,000,000 metric tons.[1] China produced 28.  of abalone ranged from 4.79-7.87 [micro]g N[H.sub.4]/h/g and were generally similar except for that values for abalone in dietary treatments [T.sub.1] and [T.sub.4] (7.61 and 7.87 N[H.sub.4]/h/g abalone, respectively) were significantly greater than that for abalone from treatment [T.sub.3] (4.79 N[H.sub.4]/h/g abalone). Metabolic rate remained relatively constant, with an average value of 90 [micro]L [O.sub.2]/h/g abalone W wt among treatments. The O:N ratio (molar molar /mo·lar/ (mo´lar)
    1. pertaining to a mole of a substance.

    2. a measure of the concentration of a solute, expressed as the number of moles of solute per liter of solution. Symbol M, , or mol/L.
    ) was similar, ranging from 9.0-10.2, for all dietary treatments with the exception of the significantly higher ratio for treatment [T.sub.3] (14.3). The higher ratio resulted from a comparatively lower rate of ammonia excretion. Mucus production, expressed as mg mucus/ day/g abalone and cal/day/g abalone, was not significantly different among treatments (statistics are not shown, however the percentages are given in as energy balance in Table 4).

    As shown in the distribution of energy, expressed as a percentage of dietary intake (Table 4), energy loss caused by respiration ranged from 5.0% to 7.7% of total intake and SDA was quite similar among all dietary treatments, ranging from 6.7% to 8.5% of the total intake. Energy loss to ammonia production was comparatively low, between 0.52% and 1.03% for all treatments. Digestible energy values calculated as the sum of growth, respiration, ammonia and mucus production corresponded to one fourth of the feed intake with values ranging from 7.8-12.7 (cal/day). However, unexplained unexplained

    strange or unclear because the reason for it is not known

    Adj. 1. unexplained - not explained; "accomplished by some unexplained process"
     energy (including feces) counted for 73.8% to 79.2% from the feed intake.

    Significant (P < 0.001) time-dependent differences in metabolism during night and day hours were observed for the juvenile abalone, reflecting a circadian rhythm circadian rhythm: see rhythm, biological.
    circadian rhythm

    Inherent cycle of approximately 24 hours in length that appears to control or initiate various biological processes, including sleep, wakefulness, and digestive and hormonal activity.
     (Fig. 1). The highest oxygen consumption was observed at night with values up to 120 [micro]L [O.sub.2]/h/g abalone W wt (22:00-04:00 h). The transition period was characterized by rapid changes in metabolic rates, such that abalone exhibited either a night- or light-adapted rate of metabolism. Percent survival remained above 90% throughout the experiment.



    Although formulations of all diets were designed to achieve a P:E ratio that was within a range of 90-100, two of the actual ratios derived from the results of analytic determinations of nutrient nutrient /nu·tri·ent/ (noo´tre-int)
    1. nourishing; providing nutrition.

    2. a food or other substance that provides energy or building material for the survival and growth of a living organism.
     composition for the 32% crude protein diets fell slightly below (87.5 and 88.3) the range. A trend of reduced weight gain as P:E ratios decreased was observed, and affirmed af·firm  
    v. af·firmed, af·firm·ing, af·firms
    1. To declare positively or firmly; maintain to be true.

    2. To support or uphold the validity of; confirm.

     the observations reported by Gomez-Montes et al. (2003). Growth rates of H. corrugata are less than those reported for H. fulgens (Gomez-Montes et al. 2003), and this difference is confirmed from observations on a commercial farm (personal communication, BC Abalone farm Erendira Baja California Baja California, state, Mexico
    Baja California (Span.: bä`hä kälēfōr`nyä), state (1990 pop. 1,660,855), 27,628 sq mi (71,576 sq km), NW Mexico, on the Baja California peninsula. Mexicali is the capital.
    , Mexico).

    Values for feed intake were greater than those reported previously for Haliotis fulgens under the same experimental conditions (Gomez-Montes et al. 2003). Feed intake may have been overestimated for those diets that had the lowest water stability and caloric intake because all organisms were probably similar (approximately 70 cal [g.sup.-] abalone) within the different dietary treatments as suggested by the results previously observed for H. fulgens by Gomez-Montes et al. (2003). Nonetheless, caloric intake was generally higher than that found for H. fulgens at the same P:E ratio (Gomez-Montes et al. 2003). In addition, the calculations for FCE and PER for the diets with reduced water stability are probably overestimates, increasingly magnified as the water stability of an experimental diet decreased. This effect would most probably be enhanced by the disturbance and nutrient loss caused by the feeding activities of the live animals as the pellets become softer.

    The amount of energy consumed was probably overestimated for those diets with reduced water stability, thereby affecting the calculation of the relative proportions of energy channeled to different metabolic activities. Therefore, the proportional losses of energy attributed to feces and unexplained energy (74% to 80%), are probably overestimates. Gomez-Montes et al. (2003) estimated feces and unexplained energy to represent approximately 45% to 55% of the total energy intake.

    When dietary protein levels were approximately 36%, a trend of higher growth of abalone was observed for dietary treatments that contained lower levels of lipid (higher starch to lipid ratios). Previously published reports suggest that carbohydrates are preferentially used as an energy source during starvation starvation, condition in which deprivation of food has forced the body to feed on itself. Causes are famine, fasting, malnutrition, or abnormalities of the mucosal lining of the digestive system.  (Durazo et al. 2004), whereas high levels of lipid (lower carbohydrate to lipid ratios) may actually retard growth (Durazo-Beltran et al. 2003). Despite sufficient carbohydrate being available as an energy source, a relatively stable amount of dietary protein is used to produce energy. Therefore, the dietary level of protein must be sufficient to satisfy the requirement for energy and still support maximum growth. Otherwise, maximum growth cannot be achieved. Maximum growth observed by Gomez-Montes et al. (2003) for H. fulgens fed diets with P:E ratios of 100 and 108 was probably because of provision of sufficient dietary protein (40% to 44%) to satisfy the protein derived energy requirement and to support maximum growth. The actual level of dietary protein needed would also be based on satisfaction of essential amino acid essential amino acid
    An alpha-amino acid that is required for protein synthesis but cannot be synthesized by humans and must be obtained in the diet.

    Respiration rate respiration rate
    Frequency of breathing, expressed as the number of breaths per minute.
     serves as an accurate measurement of the metabolic state of animals. The lack of differences in respiration rate, both in basal metabolism and specific dynamic action (SDA), suggests that the physiologic processing of the nutrients in the diets is not different. The changes in oxygen consumption observed during the night appear to be typical of the entire Haliotis genera. The magnitude of this physiologic increase was greater than that previously reported for H. fulgens (Chacon et al. 2003), suggesting a possible species-specific characteristic.

    Within a projected energy budget, more than 70% of the ingested energy was lost to feces or unexplained energy, and 7.2% to 9.9% was channeled to growth. In a previous investigation (Gomez-Montes et al. 2003), the proportion of consumed energy that was channeled to growth was higher and ranged from 17.8% to 22.8% for comparable P:E ratios. Nonetheless, the percent of digestible (not metabolizable metabolizable

    capable of being converted by metabolism.

    metabolizable energy (ME)
    said of a feed or ration, the net energy available to an animal after the utilization of some energy in the processes of digestion and absorption and
    ) energy, expressed as the sum of growth, ammonia production, mucus production, SDA and basal metabolism, that was channeled to growth for the dietary treatments in our study (33.0% to 40.6%) was comparable to the values (32.6% and 44.6%) determined in the study of Gomez-Montes et al. (2003) with a diet of a similar P:E ratio fed to H. fulgens. However, in contrast to the latter study, digestible energy, estimated as a sum of growth, respiration, ammonia production and mucous mucous /mu·cous/ (mu´kus)
    1. pertaining to or resembling mucus.

    2. covered with mucus.

    3. secreting, producing, or containing mucus.

     production, composed a much lower proportion of the intake energy.

    The inability to determine fecal energy in our investigation prevented a direct estimate of available digestible energy. In the determination of the proportional loss and use of ingested energy, digestibility digestibility

    the proportion of a feed or diet which can be digested by the normal animal of the subject species.

    digestibility coefficient
    see digestibility coefficient.
     is an important evaluation to estimate the amount of digestible energy. Previously the diet digestibility of H. fulgens was estimated (Gomez-Montes et al. 2003). However, collection of sufficient amounts of feces for proper analysis and confidence in results is difficult, and such amounts were not obtained in our study. Therefore, digestible energy was calculated as the sum of all expenditures, including energy loss caused by ammonia and mucus production. The proportional amount of digestible energy loss is much lower than that calculated for H. fulgens and is probably the result of the presumed overestimation o·ver·es·ti·mate  
    tr.v. o·ver·es·ti·mat·ed, o·ver·es·ti·mat·ing, o·ver·es·ti·mates
    1. To estimate too highly.

    2. To esteem too greatly.
     of feed intake and corresponding energy intake.

    The collective results suggest that dietary carbohydrates dietary carbohydrates,
    n the amount of simple and complex sugars consumed; the physical character of the diet. It may tend to produce or modify periodontal disease.
     and proteins are the primary and secondary sources for abalone to suitably satisfy energy requirements. The dietary protein level must be sufficient to adequately meet requirements for energy and growth and the results of this experiment suggest that a protein requirement of approximately 35% will be sufficient, provided requirements of all essential amino acids are appropriately satisfied.

    The range of lipid levels included in our experimental diets for both starch:lipid ratios probably exceeded the level needed to achieve maximum growth when sufficient protein is provided. When the protein level is met, then attention should be directed at reducing the total lipid level in the form of triglycerides Triglycerides
    Fatty compounds synthesized from carbohydrates during the process of digestion and stored in the body's adipose (fat) tissues. High levels of triglycerides in the blood are associated with insulin resistance.
     to no more than 2% to 3%. The lipid requirement may be almost exclusively based on satisfaction of requirements of essential fatty acids. However, the possible growth enhancing effect of phospholipids needs to be investigated. Van Barneveld et al. (1998) reported that high levels of dietary lipids lipids, a broad class of organic products found in living systems. Most are insoluble in water but soluble in nonpolar solvents. The definition excludes the mineral oils and other petroleum products obtained from fossil material.  could negatively impact protein digestibility, but this relationship is not supported by data obtained in our investigation. The lack of a need for high levels of dietary triglycerides is noteworthy and should eliminate a potential problem of high levels of dietary lipids interfering with the successful manufacture of practical diets. An understanding of the digestibility of structural carbohydrates would also assist in the formulation of practical diets. Any future investigations of energy budgets of abalone must incorporate methods to ensure that all diets have strong and equivalent physical integrity so that more accurate determinations of the allocation of energy can be calculated.


    The authors thank BC Abalone farm for their kind donation of the abalone used in our experiment and Roche for donating the vitamin and mineral mixtures. This work was supported by the National Council for Science and Technology (CoNaCyT), Project G28119B.


    AOAC. 1990. 16th ed. Official methods of analysis of AOAC, vol. 1. Association of Official Analytical Chemists, Arlington, VA, USA.

    Bligh, E. G. & W. J. Dyer. 1959. A rapid method of total lipid extraction and purification. Can. J. Biochem. Physiol. 37:911-917.

    Chacon, O., M. T. Viana, A. Farias, C. Vasquez & Z. Garcia-Esquivel. 2003. Circadian metabolic rate and short-term, response of juvenile green abalone (Haliotis fulgens Philippi) to three anesthetics Anesthetics
    Drugs or methodologies used to make a body area free of sensation or pain.

    Mentioned in: Appendectomy
    . J. Shellfish shellfish, popular name for certain edible mollusks (see Mollusca), e.g., oysters, clams, and scallops, and for certain edible crustaceans, e.g., crabs, lobsters, and shrimps. All are aquatic invertebrates with shells; they are not fish.  Res. 22:415-422.

    Durazo-Beltran, E., M. T. Viana, L. R. D'Abramo & J. F. Toro-Vazquez. 2003. Effect of triacylglycerols in formulated diets on growth and fatty acid fatty acid, any of the organic carboxylic acids present in fats and oils as esters of glycerol. Molecular weights of fatty acids vary over a wide range. The carbon skeleton of any fatty acid is unbranched. Some fatty acids are saturated, i.e.  composition in tissue of green abalone (Haliotis fulgens). Aquaculture 224(1-4):257-270.

    Durazo, E., M. T. Viana, L. R. D'Abramo & J. Toro-Vazquez. 2004. Effects of starvation and dietary lipid on the lipid and fatty acid composition of muscle tissue of juvenile green abalone (Haliotis fulgens). Aquaculture 238:328-341.

    Fleming, A. E., R. J. Van Barneveld & P. W. Hone hone,
    v to sharpen.
    . 1996. The development of artificial diets for abalone: A review and future directions. Aquaculture 140:5-63.

    Gomez-Montes, L., Z. Garcia-Esquivel, L. R. D'Abramo, A. Shimada, C. Vasquez-Pelaez & M. T. Viana. 2003. Effect of dietary protein:energy ratio on intake, growth and metabolism of juvenile green abalone Haliotis fulgens. Aquaculture 220:769-780.

    Green, E. & D. Carritt. 1967. New tables for oxygen saturation oxygen saturation sO2 The O2 concentration of blood expressed as a ratio of its total O2-carrying capacity; the OS is a measure of the utilization of O2 transport capacity; sO2  of seawater. J. Mar. Res. 25:140-147.

    Hahn, K. O. 1989. Nutrition and growth of abalone. In: K. O. Hahn, editor. CRC (Cyclical Redundancy Checking) An error checking technique used to ensure the accuracy of transmitting digital data. The transmitted messages are divided into predetermined lengths which, used as dividends, are divided by a fixed divisor.  Handbook of Culture of Abalone and Other Gastropods. Boca Raton Boca Raton (bō`kə rətōn`), city (1990 pop. 61,492), Palm Beach co., SE Fla., on the Atlantic; inc. 1925. Boca Raton is a popular resort and retirement community that experienced significant industrial development in the 1970s and 80s. , FL: CRC Press. pp. 135-156.

    Knauer, J., P. J. Britz & T. Hecht. 1996. Comparative growth performance and digestive enzyme Digestive enzymes are enzymes in the alimentary tract that break down food so that the organism can absorb it. The main sites of action are the oral cavity, the stomach, the duodenum and the jejunum.  activity of juvenile South African abalone, Haliotis midae, fed on diatoms and practical diet. Aquaculture 140:75-85.

    Mai, K., J. P. Mercer & J. P. Donlon. 1995. Comparative studies on the nutrition of two species of abalone, Haliotis tuberculata L. and Haliotis discus discus /dis·cus/ (dis´kus) pl. dis´ci   [L.] disk.

    n. pl. dis·ci
    A flat circular surface; a disk.


    pl. disci [L.]

     hannai Ino. III. Response of abalone to various levels of dietary lipid. Aquaculture 134:65-80.

    Monje, H. & M. T. Viana. 1998. The effect of cellulose cellulose, chief constituent of the cell walls of plants. Chemically, it is a carbohydrate that is a high molecular weight polysaccharide. Raw cotton is composed of 91% pure cellulose; other important natural sources are flax, hemp, jute, straw, and wood.  on the growth and cellulolytic cel·lu·lo·lyt·ic  
    Of, relating to, or causing the hydrolysis of cellulose: cellulolytic organisms.

    [cellulo(se) + -lytic.]
     activity of abalone Haliotisfulgens when used as an ingredient in formulated artificial diets. J. Shellfish Res. 17(3):667-672.

    SAS Institute SAS Institute Inc., headquartered in Cary, North Carolina, USA, has been a major producer of software since it was founded in 1976 by Anthony Barr, James Goodnight, John Sall and Jane Helwig.  Inc. 2001. SAS/STAT. SAS Institute Inc, Cary North Carolina North Carolina, state in the SE United States. It is bordered by the Atlantic Ocean (E), South Carolina and Georgia (S), Tennessee (W), and Virginia (N). Facts and Figures

    Area, 52,586 sq mi (136,198 sq km). Pop.
    , USA.

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    Van Barneveld, R. J., A. E. Fleming, M. A. Vandeeper, J. A. Kruk & P. W. Hone. 1998. Influence of dietary oil type and oil inclusion level in manufactured feeds on the digestibility of nutrients by juvenile greenlip abalone (Haliotis laevigata). J. Shellfish Res. 17(3):649-655.

    Widdows, J. & P. Salkeld. 1993. Practical procedures for the measurement of scope for growth. MAP Tech. Rep. Ser. 71:147-177.


    (1) Programa de Maestria y Doctorado en Oceanografia Costera, Facultad de Ciencias Marinas, Universidad Autonoma de Baja California (UABC UABC Universidad Autónoma de Baja California (Mexico)
    UABC United Aircraft Building Corporation (Russia) 
    ), Ensenada BC, Mexico; (2) Instituto de Investigaciones Oceanologicas, UABC, PO Box 453, Ensenada BC 22860, Mexico; (3) Department of Wildlife and Fisheries fisheries. From earliest times and in practically all countries, fisheries have been of industrial and commercial importance. In the large N Atlantic fishing grounds off Newfoundland and Labrador, for example, European and North American fishing fleets have long , Mississippi State University Mississippi State University, at Mississippi State, near Starkville; land-grant and state supported; coeducational; chartered 1878 as an agricultural and mechanical college, opened 1880. From 1932 to 1958 it was known as Mississippi State College. , PO Box 9690, Mississippi 39762; (4) Laboratorio de Rumiologia y Metabolismo Nutricional, Facultad de Estudios Superiores Cuautitlan, Universidad Nacional Autonoma de Mexico (UNAM), Juriquilla, Qro., Mexico; (5) Coordinacion de la Investigacion Cientifica. Facultad de Medicina Veterinaria y Zootecnia, UNAM, Ciudad Universitaria For Ciudad Universitaria in Venezuela, see Ciudad Universitaria de Caracas.

    Ciudad Universitaria is also an area of Madrid, home to the Complutense University of Madrid

    Ciudad Universitaria
    , DF, Mexico

    * Corresponding author. E-mail:
    TABLE 1.
    Ingredient (% dry weight), proximate composition
    and energy content of six experimental diets
    at two starch:lipid (S:L) ratios and three
    protein levels.
    Ingredients                              Dietary Treatments
                                    [T.sub.1]   [T.sub.2]   [T.sub.3]
    Starch:Lipids (a)                    1.5        21.8        31.6
    Fish meal (b)                       29.50       23.00       16.60
    Soybean protein isolated (c)        14.75       11.50        8.30
    Kelp meal (d)                       17.29       29.94       42.54
    Fish oil                             6.30        5.70        5.10
    Cornstarch                          17.50       15.20       12.80
    Common ingredients (e)              14.66       14.66       14.66
    Proximate composition
    Dry matter (%)                      65.47       68.28       67.51
    Crude protein (%)                   42.06       35.93       32.13
    Total lipids (%)                    11.47        8.35        8.05
    Crude fiber (%)                      2.39        3.14        4.07
    Ash (%)                             15.66       19.59       23.21
    NFEf                                28.57       32.99       32.54
    Energy (Kcal [g.sup.-1])             4.26        4.00        3.67
    P:E                                 98.70       90.00       87.50
    Ingredients                              Dietary Treatments
                                    [T.sub.4]   [T.sub.5]   [T.sub.6]
    Starch:Lipids (a)                   43.3        53.2        63.6
    Fish meal (b)                       31.40       25.00       18.30
    Soybean protein isolated (c)        15.70       12.50        9.15
    Kelp meal (d)                       11.24       24.24       37.79
    Fish oil                             3.70        3.40        3.10
    Cornstarch                          23.30       20.20       17.00
    Common ingredients (e)              14.66       14.66       14.66
    Proximate composition
    Dry matter (%)                      65.74       67.73       67.58
    Crude protein (%)                   42.64       37.57       32.48
    Total lipids (%)                     7.11        6.36        4.67
    Crude fiber (%)                      1.32        3.43        4.05
    Ash (%)                             13.54       17.72       21.92
    NFEf                                35.39       34.92       36.88
    Energy (Kcal [g.sup.-1])             4.26        3.88        3.68
    P:E                                100.10       96.80       88.30
    TABLE 2.
    Biological indices obtained for juvenile pink abalone
    (Haliotis corrugata), feed diets at two starch:lipid
    (S:L) ratios and three protein levels.
                                Dietary Treatments
    Starch:                         [T.sup.1]
    Lipids (a)                         1.5
    Stability (%)            84.45 [+ or -] 0.61 (b)
    Intake (%BW)              1.67 [+ or -] 0.03 (c)
    Intake (cal
      [g.sup.-1] abalone)    71.54 [+ or -] 1.18 (cd)
    Intake (mg PC
      [g.sup.-1] abalone)      7.03 [+ or -] 0.121
    FCE (%)                  60.57 [+ or -] 5.03 (a)
    PER                       1.42 [+ or -] 0.07 (a)
    Weight change
      (mg [d.sup.-1])        2.87 [+ or -] 0.23 (ab)
    Weight gain (%)         226.52 [+ or -] 17.56 (ab)
    SGR (% [d.sup.-1])       0.90 [+ or -] 0.04 (ab)
                                Dietary Treatments
    Starch:                         [T.sup.2]
    Lipids (a)                         1.8
    Stability (%)            82.55 [+ or -] 0.61 (b)
    Intake (%BW)              1.96 [+ or -] 0.03 (b)
    Intake (cal
      [g.sup.-1] abalone)    78.36 [+ or -] 1.18 (b)
    Intake (mg PC
      [g.sup.-1] abalone)     7.04 [+ or -] 0.12 (a)
    FCE (%)                  45.61 [+ or -] 4.59 (ab)
    PER                       1.29 [+ or -] 0.07 (a)
    Weight change
      (mg [d.sup.-1])        3.09 [+ or -] 0.23 (ab)
    Weight gain (%)         225.65 [+ or -] 17.5 (ab)
    SGR (% [d.sup.-1])       0.90 [+ or -] 0.04 (ab)
                                Dietary Treatments
    Starch:                         [T.sup.3]
    Lipids (a)                         1.6
    Stability (%)            77.63 [+ or -] 0.62 (c)
    Intake (%BW)              2.21 [+ or -] 0.03 (a)
    Intake (cal
      [g.sup.-1] abalone)    84.05 [+ or -] 1.18 (a)
    Intake (mg PC
      [g.sup.-1] abalone)     7.26 [+ or -] 0.12 (a)
    FCE (%)                  38.06 [+ or -] 4.59 (b)
    PER                       1.24 [+ or -] 0.07 (b)
    Weight change
      (mg [d.sup.-1])         2.29 [+ or -] 0.23 (b)
    Weight gain (%)         196.05 [+ or -] 17.56 (b)
    SGR (% [d.sup.-1])        0.82 [+ or -] 0.04 (b)
                                Dietary Treatments
    Starch:                         [T.sup.4]
    Lipids (a)                         3.3
    Stability (%)            88.43 [+ or -] 0.62 (a)
    Intake (%BW)              1.64 [+ or -] 0.03 (c)
    Intake (cal
      [g.sup.-1] abalone)    69.90 [+ or -] 1.18 (d)
    Intake (mg PC
      [g.sup.-1] abalone)     7.00 [+ or -] 0.12 (a)
    FCE (%)                  63.83 [+ or -] 4.59 (a)
    PER                       1.66 [+ or -] 0.07 (a)
    Weight change
      (mg [d.sup.-1])         3.72 [+ or -] 0.23 (a)
    Weight gain (%)         276.52 [+ or -] 17.56 (ab)
    SGR (% [d.sup.-1])       1.00 [+ or -] 0.04 (ab)
                                Dietary Treatments
    Starch:                         [T.sup.5]
    Lipids (a)                         3.2
    Stability (%)            83.16 [+ or -] 0.61 (b)
    Intake (%BW)              1.96 [+ or -] 0.03 (b)
    Intake (cal
      [g.sup.-1] abalone)    76.25 [+ or -] 1.18 (bc)
    Intake (mg PC
      [g.sup.-1] abalone)     7.38 [+ or -] 0.12 (a)
    FCE (%)                  53.03 [+ or -] 4.59 (ab)
    PER                      1.41 [+ or -] 0.07 (ab)
    Weight change
      (mg [d.sup.-1])         3.48 [+ or -] 0.23 (a)
    Weight gain (%)         297.06 [+ or -] 17.56 (a)
    SGR (% [d.sup.-1])        1.04 [+ or -] 0.04 (a)
                                Dietary Treatments
    Starch:                         [T.sup.6]
    Lipids (a)                         3.6
    Stability (%)            79.90 [+ or -] 0.65 (c)
    Intake (%BW)              2.15 [+ or -] 0.03 (a)
    Intake (cal
      [g.sup.-1] abalone)    80.09 [+ or -] 1.18 (ab)
    Intake (mg PC
      [g.sup.-1] abalone)     7.08 [+ or -] 0.12 (a)
    FCE (%)                  39.32 [+ or -] 4.59 (b)
    PER                      1.35 [+ or -] 0.07 (ab)
    Weight change
      (mg [d.sup.-1])         2.28 [+ or -] 0.23 (b)
    Weight gain (%)         194.85 [+ or -] 17.56 (b)
    SGR (% [d.sup.-1])        0.82 [+ or -] 0.04 (b)
    (a) Starch:lipid corn starch added:lipid content
    ratio. Standard errors are given.
    Values in the same row with different superscripts
    are statistically different P < 0.05.
    TABLE 3.
    Characteristics of shell and soft tissue for juvenile pink
    abalone (Haliotis corrugata), fed diets at two starch:lipid (S:L)
    ratios and three protein levels.
                                 Dietary Treatments
    Dietary Treatments               [T.sub.1]
    Starch:Lipids (a)                   1.5
    Shell (% dry
       basis weight)          74.25 [+ or -] 1.47 (ab)
    Soft tissue
      (% dry bais weight)     25.75 [+ or -] 1.47 (ab)
    Dry soft tissue
      (% of live weight)      12.57 [+ or -] 0.53 (b)
    Energy (Kcal [g.sub.-1]
      soft tissue)             4.83 [+ or -] 0.05 (a)
                                 Dietary Treatments
    Dietary Treatments               [T.sub.2]
    Starch:Lipids (a)                   1.8
    Shell (% dry
       basis weight)          69.46 [+ or -] 0.85 (b)
    Soft tissue
      (% dry bais weight)     30.53 [+ or -] 0.85 (a)
    Dry soft tissue
      (% of live weight)      15.92 [+ or -] 0.30 (a)
    Energy (Kcal [g.sub.-1]
      soft tissue)             4.82 [+ or -] 0.04 (a)
                                 Dietary Treatments
    Dietary Treatments               [T.sub.3]
    Starch:Lipids (a)                   1.6
    Shell (% dry
       basis weight)          75.54 [+ or -] 0.85 (a)
    Soft tissue
      (% dry bais weight)     24.46 [+ or -] 0.85 (b)
    Dry soft tissue
      (% of live weight)      13.31 [+ or -] 0.30 (b)
    Energy (Kcal [g.sub.-1]
      soft tissue)             4.75 [+ or -] 0.04 (a)
                                 Dietary Treatments
    Dietary Treatments               [T.sub.4]
    Starch:Lipids (a)                   3.3
    Shell (% dry
       basis weight)          74.66 [+ or -] 0.85 (a)
    Soft tissue
      (% dry bais weight)     25.34 [+ or -] 0.85 (b)
    Dry soft tissue
      (% of live weight)      13.83 [+ or -] 0.30 (b)
    Energy (Kcal [g.sub.-1]
      soft tissue)             4.78 [+ or -] 0.04 (a)
                                 Dietary Treatments
    Dietary Treatments               [T.sub.5]
    Starch:Lipids (a)                   3.2
    Shell (% dry
       basis weight)          74.19 [+ or -] 0.85 (a)
    Soft tissue
      (% dry bais weight)     25.80 [+ or -] 0.85 (b)
    Dry soft tissue
      (% of live weight)      14.06 [+ or -] 0.30 (b)
    Energy (Kcal [g.sub.-1]
      soft tissue)             4.85 [+ or -] 0.04 (a)
                                 Dietary Treatments
    Dietary Treatments               [T.sub.6]
    Starch:Lipids (a)                   3.6
    Shell (% dry
       basis weight)          75.74 [+ or -] 0.85 (a)
    Soft tissue
      (% dry bais weight)     25.24 [+ or -] 0.85 (b)
    Dry soft tissue
      (% of live weight)      13.25 [+ or -] 0.30 (b)
    Energy (Kcal [g.sub.-1]
      soft tissue)             4.74 [+ or -] 0.05 (a)
    (a) Starch:lipids corn starch added:lipid content
    ratio. Standard errors are given.
    Values in the same row with different superscripts
    are statistically different P < 0.05.
    TABLE 4.
    Energy budget for juvenile pink abalone (Haliotis corrugata) fed with
    a balanced diet at two starch:lipid (S:L) ratios and three protein
    levels. Energy balance is given per organism in relation to energy
                                                Dietary Treatments
            Starch:Lipids (a)                                   %
    Intake (cal [day.sup.-1])                 38.90          100.00
    Weight change (cal [d.sup.-1])             3.34            8.58
    SDA (cal [day.sup.-1]) (1)                 3.09            7.94
    Maintenance (cal [d.sup.-1]) (2)           2.81            7.22
    Ammonia (cal [d.sup.-1])                   0.38            0.98
    Mucus (cal [d.sup.-1] [g.sup.-1])          0.48            1.23
    Digestible energy (3)                      10.1
    Feces and unexplained energy               28.8           74.03
    O:[N.sub.2] (molar)                       9.4 [+ or -] 1.11 (b)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (4)               94.30 [+ or -] 6.97 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (5)               49.29 [+ or -] 1.89 (a)
    Ammonia ([micro]g N[H.sub.4.sup.+]
      [h.sup.-1] [g.sup.-1] abalone)         7.61 [+ or -] 0.44 (a)
                                                  Dietary Treatments
            Starch:Lipids (a)                                         %
    Intake (cal [day.sup.-1])                 45.79                100.00
    Weight change (cal [d.sup.-1])             4.55                  9.94
    SDA (cal [day.sup.-1]) (1)                 3.07                  6.70
    Maintenance (cal [d.sup.-1]) (2)           2.83                  6.18
    Ammonia (cal [d.sup.-1])                   0.34                  0.74
    Mucus (cal [d.sup.-1] [g.sup.-1])          0.39                  0.85
    Digestible energy (3)                     11.18
    Feces and unexplained energy              34.61                 75.58
    O:[N.sub.2] (molar)                      10.2 [+ or -] 1.21 (b)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (4)               87.88 [+ or -] 6.97 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (5)               55.34 [+ or -] 1.89 (a)
    Ammonia ([micro]g N[H.sub.4.sup.+]
      [h.sup.-1] [g.sup.-1] abalone)         6.42 [+ or -] 0.44 (ab)
                                               Dietary Treatments
            Starch:Lipids (a)                                   %
    Intake (cal [day.sup.-1])                 38.22          100.00
    Weight change (cal [d.sup.-1])             2.66            6.96
    SDA (cal [day.sup.-1]) (1)                 2.87            7.51
    Maintenance (cal [d.sup.-1]) (2)           1.89            4.95
    Ammonia (cal [d.sup.-1])                   0.20            0.52
    Mucus (cal [d.sup.-1] [g.sup.-1])          0.32            0.84
    Digestible energy (3)                      7.95
    Feces and unexplained energy              30.27           79.20
    O:[N.sub.2] (molar)                      14.3 [+ or -] 0.74 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (4)               90.98 [+ or -] 6.97 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (5)               54.94 [+ or -] 1.89 (a)
    Ammonia ([micro]g N[H.sub.4.sup.+]
      [h.sup.-1] [g.sup.-1] abalone)         4.79 [+ or -] 0.44 (b)
                                               Dietary Treatments
            Starch:Lipids (a)                                   %
    Intake (cal [day.sup.-1])                 46.69          100.00
    Weight change (cal [d.sup.-1])             4.51            9.70
    SDA (cal [day.sup.-1]) (1)                 3.94            8.47
    Maintenance (cal [d.sup.-1]) (2)           3.19            6.86
    Ammonia (cal [d.sup.-1])                   0.48            1.03
    Mucus (cal [d.sup.-1] [g.sup.-1])          0.54            1.16
    Digestible energy (3)                     12.66
    Feces and unexplained energy              33.83           72.77
    O:[N.sub.2] (molar)                       9.0 [+ or -] 0.72 (b)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (4)               93.18 [+ or -] 6.97 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (5)               51.52 [+ or -] 1.89 (a)
    Ammonia ([micro]g N[H.sub.4.sup.+]
      [h.sup.-1] [g.sup.-1] abalone)         7.87 [+ or -] 0.44 (a)
                                               Dietary Treatments
            Starch:Lipids (a)                                   %
    Intake (cal [day.sup.-1])                 46.61          100.00
    Weight change (cal [d.sup.-1])             4.35            9.33
    SDA (cal [day.sup.-1]) (1)                 3.56            7.64
    Maintenance (cal [d.sup.-1]) (2)           3.58            7.68
    Ammonia (cal [d.sup.-1])                   0.38            0.82
    Mucus (cal [d.sup.-1] [g.sup.-1])          0.34            0.73
    Digestible energy (3)                     12.21
    Feces and unexplained energy              34.40           73.80
    O:[N.sub.2] (molar)                      10.1 [+ or -] 1.4 (b)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (4)               90.48 [+ or -] 6.97 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (5)               50.60 [+ or -] 1.89 (a)
    Ammonia ([micro]g N[H.sub.4.sup.+]
      [h.sup.-1] [g.sup.-1] abalone)         6.75 [+ or -] 0.44 (ab)
                                               Dietary Treatments
            Starch:Lipids (a)                                   %
    Intake (cal [day.sup.-1])                 36.23          100.00
    Weight change (cal [d.sup.-1])             2.62            7.23
    SDA (cal [day.sup.-1]) (1)                 2.67            7.37
    Maintenance (cal [d.sup.-1]) (2)           1.87            5.16
    Ammonia (cal [d.sup.-1])                   0.26            0.72
    Mucus (cal [d.sup.-1] [g.sup.-1])          0.38            1.05
    Digestible energy (3)                      7.80
    Feces and unexplained energy              28.43           78.47
    O:[N.sub.2] (molar)                      9.11 [+ or -] 0.41 (b)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (4)               77.02 [+ or -] 6.97 (a)
    MR ([micro]L [O.sub.2] [h.sup.-1]
      [g.sup.-1] abalone) (5)               51.35 [+ or -] 1.89 (a)
    Ammonia ([micro]g N[H.sub.4.sup.+]
      [h.sup.-1] [g.sup.-1] abalone)         6.35 [+ or -] 0.44  (ab)
    (a) Starch:lipids; corn starch added:lipid content ratio.
    (1) Calculated by difference between fed and unfed abalone.
    (2) Respiration from abalone left under inanition for three days.
    (3) Calculated as the sum of growth, respiration, ammonia and
    mucus production.
    (4) Oxygen consumption from fed abalone.
    (5) Oxygen consumption from unfed abalone.
    (a,b) Values in the same row with different
    superscripts are statistically different P < 0.05.
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    Author:Vasquez-Pelaez, Carlos
    Publication:Journal of Shellfish Research
    Date:Dec 1, 2005
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