Effects of water temperature and dissolved oxygen …...Water temperature, dissolved oxygen...

Ž .Aquaculture 182 2000 339–352www.elsevier.nlrlocateraqua-online

Effects of water temperature and dissolved oxygenon daily feed consumption, feed utilization and

ž /growth of channel catfish Ictalurus punctatus

J. Alejandro Buentello, Delbert M. Gatlin III ), William H. NeillDepartment of Wildlife and Fisheries Sciences, Texas A&M UniÕersity System, College Station, TX

77843-2258 USA

Accepted 26 July 1999

Abstract

Ž . Ž . Ž . Ž .Feed intake FI , feed efficiency FE , protein efficiency ratio PER and weight gain WG ofŽ .juvenile channel catfish initially weighing 15.0"0.23 g 10–12 cm initial total length were

Ževaluated under three regimes of time-varying water temperature mean daily water temperature. Žfor Stoneville, MS, USA; mean q3; and, mean y38C and three of dissolved oxygen DO; 100,

.70 and 30% air saturation, at each temperature . Two independent experiments lasting 6 and 8weeks, respectively, simulated spring and fall temperature and photoperiod patterns. Tempera-ture–oxygen regimes were applied to randomly assigned duplicate 110-l aquaria with individualbiofilters. Fish were fed once daily, to slight excess, a commercial catfish floating feed. After 45min, uneaten feed was collected, dried and weighed; then FI was calculated by difference. Fish ineach aquarium were weighed collectively, biweekly. In both experiments higher temperatures andhigher DO levels produced increased feed consumption. A multivariate regression model wasfitted to the data to describe the combined effects of temperature and DO on FI. There was aprogressive reduction in FI as DO declined from 100 to 30% air saturation. In both trials weightgain was higher for fish held 38C above the mean water temperature at 100% air saturation valuesof DO; whereas, the lowest values of WG were obtained for fish held at 30% air saturation and38C below the mean water temperature. Weight gain increased with temperature, with maximumrates reached at 27.18C with DO equivalent to 100% air saturation. In contrast, when DO was 30%air saturation, growth rates plateaued as ambient temperature exceeded 22.88C. This model willassist in optimization of feeding practices in channel catfish aquaculture. q 2000 Elsevier ScienceB.V. All rights reserved.

Keywords: Water temperature; Dissolved oxygen; Feed consumption; Channel catfish

) Corresponding author. Tel.: q1-409-847-9333; fax: q1-409-845-4096; E-mail: [email protected]

0044-8486r00r$ - see front matter q 2000 Elsevier Science B.V. All rights reserved.Ž .PII: S0044-8486 99 00274-4

( )J.A. Buentello et al.rAquaculture 182 2000 339–352340

1. Introduction

Ž .Water temperature, dissolved oxygen concentration DO , and photoperiod are potentinfluences on feed consumption, metabolic rate and energy expenditure, and thus, on

Žgrowth of poikilothermic vertebrates, including fish Brett, 1979; Elliott, 1982; Dutta,.1994; Bhikajee and Gobin, 1998 . Therefore, the effects of these environmental factors

on fish growth and metabolism warrant thorough investigation.Feeding is the single largest cost in raising channel catfish, often exceeding 50% of

Ž .the operating expense Giachelli et al., 1982 . For aquaculture to be profitable it isimportant that the cultured organism’s rates of feed intake and growth be both high and

Ž .uniform Brett, 1979; Elliott, 1982; Koskela et al., 1997 . In practice, however,environmental conditions such as temperature, salinity, light and DO are often difficultto control; so, they may be only monitored or maintained at tolerable, as opposed to

Ž .optimum, levels Burel et al., 1996 .The present study was designed to investigate the effects of seasonally changing DO,

temperature and their interaction, on feed consumption and weight gain of juvenilechannel catfish. Attempts were made to estimate optimal temperature-DO combinationsfor feed consumption, feed conversion efficiency and growth.

2. Materials and methods

Ž . Ž . Ž .Feed intake FI , feed efficiency FE , protein efficiency ratio PER and weight gainŽ . Ž .WG of juvenile channel catfish Ictalurus punctatus were evaluated under three watertemperature regimes typical of the southeastern United States where a majority of the

Žchannel catfish production occurs mean daily water temperature for Stoneville, Missis-. Žsippi, USA; mean q3; and, mean y38C and three of DO 100, 70 and 30% air

.saturation, at each temperature . In two independent feeding trials, photoperiod andmean temperature combinations were designed to mimic spring and fall patterns

Ž .occurring at 34.28 N and 928 longitude Fig. 1 . Spring and fall trials were carried outover 6 and 8 weeks, respectively.

Ž .Fig. 1. A Patterns of water temperature during spring and fall experiments; q3 and y3 denote 38C aboveŽ .and below the mean water temperature, respectively. B Photoperiod profiles for Stoneville, Mississippi,

USA, that were simulated in the experiment.

( )J.A. Buentello et al.rAquaculture 182 2000 339–352 341

ŽJuvenile channel catfish with an initial weight of 15.0"0.23 g mean"S.D.; 10–12.cm initial total length were obtained from ponds at the Aquacultural Research and

Ž .Teaching Facility of the Texas A&M University System Burleson, TX . One hundredand eighty fish in each trial were randomly allocated to the nine temperature-DOtreatments, with two replicates per treatment. Each group of 10 fish was placed in a110-l glass aquarium containing aerated and recirculated well water. Temperature–oxygen regimes were applied to randomly assigned aquaria equipped with individualbiofilters. Water temperature was maintained at nominal values "18C by opposing roomair refrigeration with thermostatically controlled heaters in the individual aquaria.

ŽCompressed air, pure oxygen andror pure nitrogen were used to maintain DO within.5% accuracy at the three treatment levels. Temperature and DO were measured once

daily in each aquarium for maintenance of selected treatment levels. Individual aquariaŽ .were sampled constantly one aquarium each day for temperature and DO, to evaluate

differences in oxygen consumption of fish in the post-prandial state. All measurementsof water temperature and DO were performed via a YSI 610 multi-parameter environ-mental monitoring system. Weekly measurements of water quality were carried out forpH, ammonia, nitrite and nitrate levels and were kept within suitable levels for channel

Ž .catfish Lee, 1991 . Water in all aquaria was completely replaced with fresh wellwaterevery 2 weeks. Fish were acclimatized to experimental conditions for 2 weeks prior toinitiation of each feeding trial.

Ž .Fish were fed once daily 0830 an excess ration of a commercial catfish floating dietŽ .containing 40% crude protein Rangen, Angelton, TX . After 45 min, uneaten feed was

collected, dried and weighed; then FI was estimated by difference. Fish in each tankwere collectively weighed biweekly.

Ž .Established formulas Halver, 1989 were used to compute and express FI, FE, PERand WG of juvenile channel catfish. All statistical analyses were conducted using the

Ž .Statistical Analysis System SAS Institute, 1996 . Possible differences among treatmentswere tested using a nested ANOVA model for FI, WG, FE and PER. A P value of lessthan 0.05 was taken to indicate statistical significance.

3. Results and discussion

Survival of fish was 100% in both trials. The FI of fish was influenced both by waterŽ .temperature and DO in both trials Fig. 2 . As the temperature increased from 15.7 to

31.78C, FI also increased. Due to limitations of the experimental system, we were notŽ .able to produce a descending limb supra-optimal temperature on the rate-temperature

curve for feed consumption. Feed intake was significantly lower at 15.78C than at allother temperatures, and FI during the spring experiment was higher at 308C and 100%air saturation than at any other combination in either experiment. There was a progres-sive reduction in FI as DO declined from 100 to 30% saturation.

Growth patterns for juvenile channel catfish in terms of WG increment for both thespring and fall feeding trials are shown in Fig. 3. In both trials WG was greatest for fishheld 38C above the mean water temperature at 100% air saturation values of DO;whereas, lowest values were obtained for fish held at 30% air saturation and 38C belowthe central water temperature. Weight gain increased with temperature, with maximum


Ž Ž . . Ž .Fig. 2. Feed intake of channel catfish gr100 g body weight BW rday obtained for both spring A and fallŽ .B experiments. Each data point represents the mean of two replicates for each temperatureroxygencombination averaged over the duration of each trial.

Ž .rates averaged for both trials reached at 27.18C and 100% air saturation. In contrast,WG almost plateaued as ambient temperature exceeded 22.88C when DO was 30% air

Ž .saturation. Marked differences in WG were observed at each level of DO Fig. 4 andŽ .temperature Fig. 5 . In both experiments of the present study, apparent FE ranged from

Ž .0.1 to 1.0 g gainrg fed , and showed significant differences related to temperature andŽ .oxygen levels Fig. 6 . The ranges of PER were temperatureroxygen-dependent as well.

The association between WG, water temperature and DO for channel catfish found inŽ . Ž .this study and disregarding transient-state effects is described by Eq. 1 :

WGs3.061y0.099 T y0.0663 O q0.004258317 T )O 1Ž . Ž . Ž . Ž .Ž . Ž .where: WGsWeight gain g , TsWater temperature 8C , OsDissolved oxygen

Ž .% air saturation .

Ž .Fig. 3. Total weight gain increment expressed as percent of initial weight for juvenile channel catfishŽ . Ž . Ž .10 fishraquarium; two replicates per treatment during spring A and fall B experiments. Symbols with

Ž .different superscript letters indicate intergroup statistical differences P -0.05 among treatments at the end ofweek 6. Mean indicates mean daily water temperature for Stoneville, Mississippi, USA, with q3 and y3indicating 38C above and below the mean temperature, respectively. Dissolved oxygen concentration is notedas 100, 70 or 30% air saturation.


Ž .Fig. 4. Dissolved oxygen effects on weight gain increment percent of initial weight for channel catfishŽ . Ž10 fishraquarium; two replicates per treatment averaged for both spring and fall experiments each at

.6-weeks duration . Superscript letters express intergroup statistical differences between percent air saturationlevels of dissolved oxygen. Means not sharing a common letter are significantly different from each otherŽ .P -0.05 .

Ž .Fig. 5. Temperature effects on weight gain increment percent of initial weight for channel catfishŽ . Ž10 fishraquarium; two replicates per treatment averaged for both fall and spring experiments each at

.6-weeks duration . Superscript letters express intergroup statistical differences among temperature regimes.Ž .Means not sharing a common letter are significantly different from each other P -0.05 . Mean indicates

mean daily water temperature for Stoneville, Mississippi, USA, with q3 and y3 indicating 38C above andbelow the mean temperature, respectively.


Ž .Fig. 6. Feed efficiency values for channel catfish 10 fishraquarium; two replicates per treatment obtained forŽ . Ž .both spring A and fall B experiments. Each data point represents the mean of two replicates for each

temperatureroxygen combination averaged over the duration of each trial.

To better examine data from the present study, an XYZ coordinate system was usedŽ . Žto locate the response Z in a tridimensional 3D scatter plot. Interaction tests SAS

.Institute, 1996 were used to evaluate statistical significance of individual factors andŽinteractions among them. Then, selected factors temperature, oxygen and

. Žtemperature)oxygen were submitted to a general linear model procedure SAS Insti-.tute, 1996 to obtain numerical coefficients for the predictive equation. This model

generated new data points through the use of the inverse distance algorithm systemŽ .SigmaPlot, 1994 , which evaluated every original data point, interpolated it, and thus

Ž .converted the original 3D scatter data into a Cartesian mesh plot Fig. 7 . This allows forincreased predictive power within the tested ranges. Because interpolated data points aregraphed as the intersection of the mesh grid, pinpointing predicted values within thismesh is greatly simplified. As expected, increases in temperature and oxygen levelsgenerally resulted in increases in the rates of physiological processes.

3.1. Temperature effect

The regulatory mechanisms underlying the relationship between growth rate andtemperature are likely related to the enzymatic modulation of metabolic processesŽ .Sumpter, 1992; Al-Asgah and Ali, 1997 . The general relationship between temperatureand growth found in our experiment is illustrated in Fig. 7. There was very limited WG

Ž .in fish held at the lowest temperature 15.78C ; above this, the total increment in weightincreased with temperature. The results of the present study are in line with typical

Žrate–temperature curves for warmwater species West, 1965; Andrews and Stickney,1972; Cox and Coutant, 1981; Cuenco et al., 1985; Carrieri et al., 1989; Tandler et al.,

.1989; Jobling, 1993; Iwata et al., 1994; Jobling, 1995; Requena et al., 1997 . Thegrowth rate of most warm water fish species increases with increasing temperature, to avalue a few degrees below the upper lethal limit, provided the feed supply is not limited


Fig. 7. Predictive Cartesian mesh plot for feed consumption based on results of fall and spring experiments.Ž . Ž . Ž .Grid was fitted to the predictive equation FCs0.163y0.001) T y0.002) O q0.00009) T )O , where

T s temperature and Osdissolved oxygen. New data points were generated within tested ranges onlyŽ .15–328C and 30–100% oxygen saturation through the use of the inverse distance algorithm systemŽ .SigmaPlot, 1994 .

Ž .Corey et al., 1983; Heap and Thorpe, 1987; Talbot, 1993 . Such temperature-dependentincrease in growth rate reflects increases in appetite, foraging efficiency, and biochemi-

Žcal reaction rates within the thermal tolerance range of the fish Brett and Groves, 1979;.Cossins and Bowler, 1987; Jobling, 1993 . Maximum growth rate for the present

Ž .experiment was achieved at approximately 288C. According to Jobling 1993 , appetitepeaks at temperatures approaching the upper thermal tolerance limit of fish speciesfalling dramatically above the optimum and decreasing more gradually below theoptimum. At higher water temperatures, fish feed more vigorously and their digestive

Ž .processes are accelerated Cossins and Bowler, 1987 as was observed in the presentŽ .study. Heap and Thorpe 1987 observed that turbot and turbot-brill hybrids grew faster

at higher environmental temperatures, mainly because of much improved appetite, butŽ .also perhaps due to an increase in feed efficiency. Cai and Curtis 1990 , however,

reported that growth rate and feed consumption, but not assimilation efficiency oftriploid grass carp increased with environmental temperature.

3.2. Light-temperature cycling

In the present study, WG was greater during spring than during fall for equivalentŽ .temperaturerDO regimes Fig. 3 . Increasing photoperiod has been shown to increase


growth rates of several fish species such as juvenile Atlantic salmon and green sunfishŽ .Jobling, 1993 . Photoperiod could act as an inducer of daily rhythms observed in fish

Ž .hormonal secretions Requena et al., 1997 . Corroborative evidence for the growth-pro-moting effect of light-temperature cycling has been demonstrated in studies with

Žjuvenile turbot, reared under different temperature and photoperiod regimes Imsland et.al., 1995 . Also, juvenile sockeye salmon usually grow more rapidly under conditions of

photothermal cycling than when held in continuously warm or cool water withoutŽ .photoperiod manipulation Biette and Green, 1980; Jobling, 1995 .

3.3. Oxygen effect

ŽIncreasing DO, up to some limiting value, results in enhanced growth of fish Brett.and Groves, 1979; Cuenco et al., 1985; Neill and Bryan, 1991 . The association betweenŽ Ž ..weight gain and DO inferred for channel catfish in this study Eq. 1 is in agreement

Ž . Žwith that measured for largemouth bass Stewart et al., 1967 , common carp Chiba,. Ž . Ž1966 , coho salmon Hermann et al., 1962; Fisher, 1963 , northern pike Adelman and

. Ž .Smith, 1970 and brook trout Whitworth, 1968 .Fish need oxygen to aerobically generate energy for body maintenance, locomotion

Ž .and biosynthetic processes Van Dam and Pauly, 1995 . The sharp limitation of WGŽ .brought about by lowest values of DO in the present study Fig. 4 can be explained by

the effects of oxygen availability on appetite, FE and metabolism. Attempts have beenŽmade to determine levels of DO at which FI and growth become affected Hogendoorn

. Ž .et al., 1983; Cui and Wootton, 1988 . According to Cuenco et al. 1985 , increasing DOup to some limiting level, at constant levels of all other factors, increases feedconsumption. In the present study, limiting DO levels appeared to be around 70% airsaturation for both spring and fall regimes. Below this level, FI values were sharplyreduced and temperature seemed to have lesser influence on the fish’s appetite. Similar

Ž .results have been obtained with rainbow trout Jobling, 1993 reared under variousconditions of DO. In that case FI was reduced when the DO fell below 60% airsaturation. Also, results from the present experiment corroborate those of Andrews and

Ž .Matsuda 1975 , who showed reduced rates of FI and WG of channel catfish held at268C in water with 60% air saturation. Taken together, these data appear to indicate thatDO below 70% air saturation limits feed consumption of channel catfish. Data relatinglimiting DO levels for FI in other species of fish are sparse, but from the evidenceavailable it appears that limiting levels of DO are usually between 50 and 70% air

Žsaturation, with values for salmonids being towards the top end of the range Jobling,.1995 . It is important to state that the above suggested limiting level of oxygen

saturation was determined for channel catfish under conditions of unrestricted feedingand, consequently, the level of DO that limits growth would be gradually reduced asfeed becomes more restricted.

Ž Ž ..The relationship between WG and DO for channel catfish Eq. 1 is consistent withŽ . Ž .that measured for coho salmon by Fisher 1963 . Garces-Botacio 1991 hypothesized

that the effect of DO on feed consumption of the marine red drum was in part an effecton appetite. This relationship indeed may follow a similar pattern for various fishspecies. Reduced DO may lead to restriction of feeding activity, andror changes in


digestion and assimilation efficiency. The reduction in FI rates found in the presentstudy also can be explained by the effects of insufficient DO in slowing stomach

Ž .evacuation Smith, 1989; Becerra, 1998 .The rate of oxygen uptake by channel catfish has been evaluated for fed and unfed

Ž .individuals Andrews and Matsuda, 1975 . The oxygen consumption value for satiated,Ž100-g fish was 1.45 times higher than that of unfed fish 0.81 and 0.56 g O rkg fishrh,2

.respectively . During unfed conditions, lipid and protein oxidation account for 90–95%Ž .of total oxygen use Van Dam and Pauly, 1995 ; whereas, in fed fish, amino acid

Ž . Ž .oxidation replaces lipid oxidation Dutta, 1994 . According to Jobling 1981b; 1983increased postprandial oxygen demand should be attributed mostly to digestive andpost-absorptive effects, with only a minor fraction being ascribed to peristalsis andenzymatic secretions. It has been determined that high-protein diets notably increase

Ž . Ž .oxygen consumption and specific dynamic action SDA Matty and Lone, 1985 . In thepresent study, differences in oxygen consumption were suggested by the depression inDO after feeding. Magnitudes of the depression indicated that channel catfish subjected

Ž .to the q38C, 100% air-saturation treatment that had a greater FI had oxygenconsumption rates about 1.5 times greater than those of fish in the y38C, 100% regime.

Ž .These results are entirely consistent with those of Andrews and Matsuda 1975 .

3.4. Weight gain and feed intake

Channel catfish in the present study grew fastest when water temperature wasbetween 27 and 288C. Generally, channel catfish do not feed consistently in ponds whenwater temperature drops below 218C. Although they will feed at temperatures as low as108C, the amount and frequency of feeding are greatly reduced. Therefore, in coolerwater, growth rate is markedly depressed. The most favorable temperature indicated

Ž .above is slightly lower than that proposed by Andrews and Stickney 1972 , whosuggested ‘‘approximately’’ 308C as the optimal temperature for catfish growth. This is

Ž .probably influenced by differences in fish size 4 vs. 15 g in the present experiment . Itis generally accepted that as a fish increases in size, its temperature optimum for

Žmaximum growth shifts to lower temperatures Elliott, 1976; Brett, 1979; Cuenco et al.,.1985; Jobling, 1995; Koskela et al., 1997 .

Growth of fish is largely dependent on consumption of feed, its assimilation andŽconversion into body tissues Nikolski, 1963; Brett and Groves, 1979; Dutta, 1994;

.Burel et al., 1996 . It is clearly evident from the present data that growth rate wasŽstrictly dependent on FI. The slower growth rates in treatment y38, 30% in comparison

.with q38, 100% were due mainly to lower FI. A similar relation of growth to rate of FIŽ . Žhas been demonstrated in largemouth bass Thompson, 1941 , brown trout Pentelow,

. Ž1939; Elliott, 1975, 1976 , sockeye salmon Brett et al., 1969; Brett, 1971; Brett and. Ž .Shelbourne, 1975; Biette and Green, 1980 , common carp Huisman, 1974 , margined

Ž . Ž . Žsculpin Davis and Warren, 1965 , coho salmon Stauffer, 1973 , channel catfish West,. Ž .1965; Andrews and Stickney, 1972 and striped bass Cox and Coutant, 1981 .

To prevent feed limitation and excessive competition, the present study provided atemporal ‘clumping’ of feed supply. Consequently, dominant individuals, if any, were


unable to defend the feed source and other fish within the group were able to obtaintheir fair share of the supplied feed. Under this experimental protocol, there was nosocial hierarchy established and fish weight dispersion was limited. It has been demon-strated that high feeding rates usually result in reduced individual growth variationŽ .Fauconneau et al., 1998 .

Because feed supply was not limiting in the present study, feed consumption waspositively correlated with ambient temperature and DO. Feed consumption was morethan two-fold higher at 28.88C than at all temperatures below 228C. Also, thermaltreatments combined with 30% air saturation of DO resulted in intake levels at the low

Ž .end of the spectrum, for both the spring and fall experiments Fig. 2 . The decrease inappetite of fish was most evident for the low temperaturerlow oxygen treatments andfrom a practical stand point, there could be a risk of overfeeding when water tempera-ture drops below 208C and DO levels decline below 60% air saturation. Feed intake isregulated partially by the speed with which digesta pass through the gut, which, aspreviously indicated, is affected both by temperature and DO. In a general way, passage

Ž .increases as temperature Flowerdew and Grove, 1979; Kaushik, 1981 and DO increase.Elevated water temperature generally increases metabolic energy demands; thus, highfeeding rates are typically required to meet these demands. At the same time, increased

Žmetabolic rates require increased oxygen availability and transport rates Van Dam and.Pauly, 1995 .

Ž .The lowest FI less than 1.4 gr100 g BWrday was obtained in the present studyŽ .with the low temperaturerlow DO combination Fig. 2 . Feed intake progressively

increased with increases in temperature and DO, up to 5 gr100 g BWrday at 308C and100% air saturation. In this study, rates of FI are comparable to those measured by

Ž .Andrews and Stickney 1972 for channel catfish held at 26–278C. These results also areŽ .in agreement with those of Platt and Hausser 1978 , who found increases in feeding rate

of Tilapia zillii when the temperature was changed from 28.88C to 31.48C and that athigher or lower temperatures, feeding rate and swimming stamina were reduced.

Based on the preceding information and the general profiles of the Cartesian plot forŽ . Žfeed consumption Fig. 7 , it can be inferred that maximum FI rates 3.4–4.6 gr100 g

.BWrday are likely to occur at 31–328C and 90–100% air saturation. A practicalapplication of these data indicate that a shift in environmental conditions from 318C,50% air saturation to 318C, 70% could increase consumption rates from 1.7 to 3.1gr100 g BWrday, almost a two-fold increase.

Results of the present study indicate that FI kept increasing up to a temperatureŽ .2–38C higher than the optimum for channel catfish growth e.g., 30 vs. 288C . Because

Žof temperature’s differential effects on standard and active metabolic rates Neill and.Bryan, 1991 , the optimum temperature for fish growth tends to be a few degrees lower

Žthan the temperature at which FI is maximal Brett, 1971; Flowerdew and Grove, 1979;Cox and Coutant, 1981; Kaushik, 1981; Elliott, 1982; Wurtsbaugh and Cech, 1983;

.Keast, 1985; Xiao-Jun and Ruyung, 1992; Jobling, 1993; Koskela et al., 1997 and datafrom the present study agree with this conclusion. These results are similar to those of

Ž .Woiwode and Adelman 1991 who reported that feed consumption of hybrid stripedbass increased with temperature up to 29.28C; whereas, the optimum temperature forgrowth was 26.88C, and FE peaked at 21.28C.


3.5. Feed efficiency

Feed efficiency depends on several factors including feeding level and water tempera-Ž .ture Jobling, 1993 . In fish fed maximum rations, FE increases with increasing

temperature up to a maximum and then decreases as temperature continues to rise. TheŽbest FE is generally found at a lower temperature than that for optimum growth Elliott,

.1976; Brett, 1979; Jobling, 1993, 1995 . Results from the present study are in accor-Ž .dance with this. Feed efficiency was best 1.0 for spring and fall for channel catfish

Ž .held at 24–268C Fig. 6 ; whereas, the optimum temperature for growth was estimatedto be around 288C. The values of FE found for channel catfish followed a trend similar

Ž .to the one reported for rainbow trout Staples and Nomura, 1976; Makinen, 1994 . It isgenerally accepted that adaptive mechanisms exist which regulate the capacity of fish toutilize feed for growth over a range of temperature, as described in other species, such

Ž . Ž .as chinook salmon Brett, 1971 , walking catfish Hogendoorn et al., 1983 andŽ .European minnow Cui and Wootton, 1988 .

3.6. Temperature–oxygen interaction

Step-wise regression analyses, indicated that the variables temperature, DO, andtemperature)DO accounted for most of the variation in FI, WG, FE and PER of channelcatfish in the present study. In general, the inclusion of such variables in the model

Ž .resulted in highly significant P-0.0001 effects when evaluated for interaction. When2 Ž .all were included they raised R to 0.92. Garces-Botacio 1991 , when modeling the

growth performance of red drum, found that DO and temperature were statistically morerelevant than ammonia, nitrite, phosphorus, and pH. Thus, it seems evident from thepresent study that temperature, DO and their interaction directly affect the physiologicalpatterns of juvenile channel catfish by altering the metabolic rate of this species. As eachfactor deviates from the optimal level it becomes more important in limiting responsessuch as FI, WG, FE and PER. For example, under conditions of low temperature, the

Ž .ingestion rate is low despite high oxygen availability Fig. 7 . High DO by itself doesnot promote high FI because there is no need for high energy supply when the

Ž .metabolism temperature-controlled is depressed. Similarly, when high temperaturesinduce increased metabolic rates, DO is needed to support high energy demands.Because the present experimental design included three fixed levels of DO and two of

Ž Ž ..those were at or above adequate, it is reasonable that the observed relationship Eq. 1Ž .involved a coefficient for temperature of a higher magnitude 1.50 fold than the

coefficient for DO.On the other hand, when DO went below 70%, FI rates were drastically reduced and

temperature seemed to have less influence on the amount of feed consumed. It is logicalto assume that, given the opportunity, fish will choose the least compromised combina-tion of these two factors. This assumption agrees with results of behavioral regulation

Ž . Ž . Ž .studies reviewed by Brett 1979 , Brett and Blackburn 1981 , Jobling 1981a , KelloggŽ . Ž .and Gift 1983 and Weber and Kramer 1983 , and is consistent with the model

Ž .proposed by Neill and Bryan 1991 . Thus, the impression is gained that there is a closecorrespondence between environmental preferenda and the joint optima of temperatureand DO needed to achieve maximal physiological performance.


Acknowledgements

This study was supported in part by the Southern Regional Aquaculture Centerthrough grant number 95-38500-1411 from the United States Department of Agriculture.Funding for Mr. Buentello was provided in part by the Consejo Nacional para la Ciencia

Ž .y la Tecnologıa CONACYT-Mexico .´ ´

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