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Anais da Academia Brasileira de Ciências 2019 914 e20190075 Annals of the Brazilian Academy of Sciences Printed version ISSN 00013765 Online version ISSN 16782690 httpdxdoiorg10159000013765201920190075 wwwscielobraabc wwwfbcomaabcjournal An Acad Bras Cienc 2019 914 AgrArIAN SCIeNCeS Metabolic responses to dietary carbohydratelipids ratios in neotropical hybrid catfish Pseudoplatystoma corruscansPseudoplatystoma fasciatum DANIEL OKAMURA1 RODRIGO FORTESSILVA2 RENAN R PAULINO 1 FELIPE G DE ARAÚJO3 DIEGO V DA COSTA4 RAQUEL T PEREIRA1 and PRISCILA V ROSA1 1Department of Animal Science Federal University of Lavras UFLA Aquenta Sol sn Campus Universitário 37200000 Lavras Mg Brazil 2Department of Animal Science Federal University of Viçosa UFV Av Peter Henry rolfs sn Campus Universitário 36570900 Viçosa Mg Brazil 3Department of Animal Science Federal rural University of Pernambuco UFrPe UAg Av Bom Pastor sn Boa Vista 55292270 garanhuns Pe Brazil 4Institute of Agricultural Sciences ICA University of Minas gerais UFMg Av Universitária 1000 Bairro Universitário 39404547 Montes Claros Mg Brazil Manuscript received on January 24 2019 accepted for publication on May 10 2019 How to cite OKAMUrA D FOrTeSSILVA r PAULINO rr ArAÚJO Fg COSTA DV PereIrA rT AND ROSA PV 2019 Metabolic responses to dietary carbohydratelipids ratios in neotropical hybrid catfish Pseudoplatystoma corruscansPseudoplatystoma fasciatum An Acad Bras Cienc 91 e20190075 DOI 10159000013765201920190075 Abstract The purpose of this study was to assess the effect of different carbohydrate and lipid CHOL ratios on the performance and energy metabolism of hybrid catfish Pseudoplatystoma corruscans Pseudoplatystoma fasciatum One hundred and sixtyeight fish 1049165 g BW were distributed into 24 100L tanks The experiment was performed using a randomised block design with four CHOL ratios 045 100 166 and 285 and six replications growth performance protein content glucose and triglycerides in plasma muscle and liver and the hepatic activity of the malic and glucose6phosphate dehydrogenase G6PD enzymes were measured after 60 feeding days No significant differences were observed in growth performance protein and triglycerides of plasma and muscle hepatic protein or malic enzyme activity CHOL 285 provided the highest plasma and liver glucose while CHOL 045 showed high liver triglycerides A linear effect was found for the hepatic activity of G6PD and dietary starch levels r2093 The hepatic activity of g6PD was greater in CHOL285 than for CHOL 045 In conclusion the limit of dietary starch inclusion and lipids were 15 and 900 respectively CHOL 166 and this level did not affect the energy metabolism of the juvenile hybrid catfish Key words enzymes activity hepatic function intermediary metabolism hybrid catfish Surubim Correspondence to Priscila Vieira rosa Email prisciladzouflabr OrCID httpsorcidorg000000020052079X INTRODUCTION Carbohydratecontaining feedstuffs are less expensive than other nutrients in diet and directly influence economic fish production viability Papoutsoglou and Lyndon 2006 Kamalam et al 2017 Although no nutritional requirements of carbohydrates exist in these animals diets their inclusion at appropriate levels can ensure better efficiency in utilising other nutrients by contributing positively in the diet extrusion process and acting DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 2 9 as an abundant energy source for lipids and proteins Krogdahl et al 2005 NrC 2011 Zhou et al 2015 Kamalam et al 2017 However carbohydrate use in a carnivorous fish diet is complex and not well known for digestion absorption and metabolism processes Krogdahl et al 2005 Such processes depend on the relationships between fish species and carbohydrate type or content Krogdahl et al 2005 Kamalam et al 2017 These species are more efficient in using lipids as an energy source given their metabolisms limited capacity to reduce glycaemia Caseras et al 2007 Hemre et al 2002 The metabolic profile and its pathways are the reflex from diet regardless of performance or muscle and liver function Carbohydrate lipids CHOL ratios of around 131 produce the best growth performance in hybrid catfish Pseudoplatystoma reticulatum x Leiarius marmoratus Bernardes et al 2016 One of the statements about fish growth variation is the relation between carbohydrate levels and their effects on diet viscosity and digestion which can in turn lead to a change in nutrient use TranTu et al 2017 Moreover increased dietary protein and fewer carbohydrates and lipids attenuate glycolytic activity and induce hepatic gluconeogenesis in South American catfish Rhamdia quelen Melo et al 2016 The liver histology was normal in the yellow catfish Pelteobagrus fulvidraco fed the 245 363 and 558 CHOL diets The hepatic steatosis and hepatocyte apoptosis rates increased in the yellow catfish fed with 111 and 167 CHOL than 245 363 and 558 CHOL Wang et al 2014 Thus it is imperative for modern aquaculture to define the optimum dietary ratio between nonprotein energy sources such as carbohydrates and lipids Wang et al 2016 Pseudoplatystoma corruscans and Pseudoplatystoma fasciatum are carnivorous fish and important species with commercial characteristics produced in South America Martino et al 2002a Leonardo et al 2004 along with their hybrids P corruscansP fasciatum Beelen et al 2003 Pseudoplatystoma spp is able to adjust amylase activity in accordance with dietary carbohydrate levels Lundstedt et al 2004 Both plasma glucose and plasma protein concentrations lower when corn gluten meal levels increase in Pfasciatum Bicudo et al 2012 All these South American catfish are species of the same family Pimelodidae However very little is known about this hybrid metabolism function in relation to the nutrition effect In some carnivorous species eg Oncorhynchus mykiss Rachycentron canadum and Mystus nemurus glucose stimulates glucose6phosphate dehydrogenase g6PD and malic enzyme Me activities Alvarez et al 2000 Cui et al 2010 Hamid et al 2011 which are the main enzymes responsible for producing NADPH which is required for lipid synthesis Thus the quantification of metabolites in different tissues and enzyme activity in relation to energy metabolism are important for understanding the use of dietary carbohydrates to optimise growth spare proteins and avoid economical waste while farming species The present work was conducted to assess the influence of different CHOL ratio on the energy metabolism of hybrids Pseudoplatystoma corruscansPseudoplatystoma fasciatum by estimating performance protein concentration glucose and triglycerides in plasma muscle and liver and the hepatic activity of both Me and g6PD MATERIALS AND METHODS ANIMAL HOUSINg The growth trial was performed at the Aquaculture Station of the Federal University of Lavras UFLA Minas gerais Brazil and was approved by the UFLAs ethics Committee on Animal Use and was certified by Protocol 02812 Juvenile catfish Pseudoplatystoma spp were obtained from a farm in Minas gerais Brazil One hundred and DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 3 9 sixtyeight fish initial body weight of 1049165 g meanSD were distributed into 24 100L tanks 7 fish per tank Fish were fed twice until apparent satiation The system comprised a 5μm particle sand filter an ultraviolet filter and a biological filter The water flux in the experimental boxes was sufficient to renew water 3 timeshour During the experiment the water temperature was maintained at 2801oC Dissolved oxygen was measured daily by a portable digital oxymeter F1550A Bernauer Indaial SC Brazil with average values of 5109 mg L1 Ammonia nitrite and pH were measured weekly using a commercial kit Labcon Test Alcon Camboriú SC Brazil with average values of 00090002 ppm 0010001 ppm and 7301 respectively Fish were submitted to a 15day adaptation period before the experiment began eXPerIMeNTAL DIeTS Four isoproteic 460 g kg1 crude protein and isoenergetic 3500 kcal kg1 diets were formulated to contain the dietary carbohydrate and lipid ratios CHOL 045 100 166 and 285 Table I Canola oil was considered herein because the parietal P fasciatumis well able to use plant oils than fish oil to transform linoleate into arachidonate Arslan et al 2008 Moreover plant oil is used by P coruscans Martino et al 2002b Dietary ingredients were mixed in a paddle mixer Inbramaq São Paulo Brazil with 20 of water pelletized 4 mm and dried for 12 h in an oven at 50C Diets were stored in a freezer 5C until used eXPerIMeNTAL DeSIgN After the adaptation period fish were fed the experimental diets 4 CHOL ratios and 6 replies twice a day at 0800h and 1800h The amount of diet was provided according to apparent animal satiety The delivered amount of diets was quantified daily by weighing before and after the feeding process The trial lasted 60 days At the end of the experimental period fish were starved for 24h weighed and then kept in anaesthetic solution until opercular movement had completely ceased 08 mll1 2phenoxyethanol One millilitre of blood per cardiac punch was also sampled using a heparinised syringe 1 ml needle size of 21g 12 08x40 Blood was centrifuged at 3000 rpm for 10 minutes before isolating plasma by a Pasteur micropipette After this procedure fish were decapitated at the skull insertion point Then the liver and white dorsal muscle samples approximately 3g were collectedfrom each juvenile The liver TABLE I Composition of the experimental diets Ingredients Inclusion levels Fish meal 1 5717 5217 4800 4400 gelatine 500 500 600 700 Albumin 700 700 800 800 Casein 300 600 700 850 Canola oil 1100 1000 900 700 Starch2 500 1000 1500 2000 Dicalcium phosphate 150 150 150 150 Mineral andvitamin 3 030 030 030 030 Antioxidant 003 003 003 003 Cellulose 500 400 317 260 Inert 500 400 200 100 Total 10000 10000 10000 10000 Carbohydrate lipid ratio 045 100 166 285 Approximate composition g100 g dry matter CP4 467 463 463 462 Ce5 kcalkg 1 3525 3548 3595 3550 1Salmon meal 67 crude protein Total Alimentos 2Carbohydrate source maize raw starch Pachá Alimentos 3Composition per kilogram of the mineral and vitamin supplement 1500 IU vitamin A 20 mg vitamin B1 15 mg vitamin B2 1000 IU vitamin B3 10 mcg vitamin B12 25 mg vitamin e 120 mg vitamin PP 2000 mg choline 80 mg calcium pantothenate 2 mg folic acid 80 mg manganese 24 mg iron 50 mg zinc 8 mg copper 3 mg iodine 010 mg selenium and 170 mg BHT 4CP crude protein 5Ce crude energy DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 4 9 was weighed to calculate the hepatossomatic index Samples were stored at 80oC for further analyses The following parameters were calculated Hepatosomatic Index HSI liver weightfish weight100 weight gain WG final weight first weight feed conversion ratio FCR ingested rateweight gain protein efficiency rate PER weight gainingested crude protein and energy efficiency rate EER weight gainingested crude energy Muscle and liver assessments The liver and muscle homogenates were obtained according to Paulino et al 2018 with slight modifications The white muscle and liver samples were homogenized in icecold deionised water and centrifuged 13400 g 10 min 4ºCThe resultant supernatant was collected and stored at 80ºC glucose and triglyceride levels were measured with commercial kits glucose HK Liquiform Cat 85 and Triglycerides Liquiform Cat 87 Soluble protein content was determined according to Bradford 1976 with bovine serum albumin solution Sigma used as a standard The specific activity of both malic enzyme Me eC11140 and glucose6phosphate dehydrogenase g6PD eC11149 was estimated following the methodology outlined in graeve et al 1994 and Spina et al 1970 respectively Hepatic tissue was macerated with a pistil in an icebath After maceration the sample was put in an ultrasonic bath for 10 minutes before being centrifuged at 10000 g and 4oC for 30 minutes before collecting the supernatant The homogenate of supernatants was used to estimate specific G6PD activity All the enzymatic activities are shown as Umg1 protein STATISTICAL ANALYSIS All the obtained data were analyzed using the SAeg statistical package proposed by euclydes 1997 A data analysis was done by a oneway analysis of variance ANOVA after testing for normality and homogeneity of variances with the ShapiroWilk test and the Levene test respectively Significant differences among the means were determined by a StudentNewmanKeuls test A regression equation was obtained to describe the glucose6phosphate dehydrogenase pattern RESULTS No fish mortality was recorded during the experimental period After the 60 days that the trial lasted no growth performance parameter eg weight gain feed conversion rate Hepatosomatic Index protein efficiency rate and energy efficiency rate was significantly affected P005 by the dietary carbohydratelipid CHOL ratio Table II The concentrations of glucose triglycerides and protein in the white muscle of juveniles did not change P005 Table III However CHOL had an effect on glucose in both plasma and the liver as did triglycerides on hepatic tissue P005 The highest plasma and liver glucose contents were found in the fish fed the diet CHOL 285 The catfish juveniles fed the diet CHOL045 showed higher concentrations of hepatic triglycerides compared to other CHOL ratios P005 The plasma and liver protein concentrations were not affected by diet P005 No significant differences were observed for the hepatic activity of Me P005 However g6PD activity was greater in the treatment CHOL 285 compared to CHOL045 P005 Table IV A linear effect was found for G6PD activity which increased with a rising carbohydrate concentration in diet Figure 1 DISCUSSION In this study the fish fed different CHOL ratio in their diet did not show any significant differences in their performance The range within which DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 5 9 dietary carbohydrates can be used to meet energy requirements vastly varies between and even within species depending on many factors which have been categorised into those also related to biology and experimental conditions Kamalam et al 2017 For example the inclusion of dietary soluble polysaccharides does not affect growth performance nor the feed conversion ratio in African catfish Clarias gariepinus Martino et al 2005 found that replacing the energy obtained from soya bean oil in diet with corn flour did not affect the specific growth rate the feed conversion rate total consumption protein and the hepatosomatic index of the parietal Pseudoplatystoma ssp On the other hand the 4 8 and 12 increases in soya bean oil led to enhanced nitrogen fixation in a P coruscans carcass Campos et al 2006 The feed conversion ratio and daily feed consumption showed a marked drop in the inverse proportion to the lipid level in diet in P coruscans Martino et al 2002b In short the present study revealed that the hybrid P corruscansP fasciatum Figure 1 Linear regression of the hepatic glucose6phosphate dehydrogenase activity of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days n18 TABLE II Growth performance feed utilisation and Hepatosomatic Index of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days Lack of superscript letters in the same line implies no differences in data at P005 Parameters Carbohydrate lipid ratios1 045 100 166 285 WG 11742 2749 11943 2467 12543 2100 13297 2281 FCr 111 027 118 022 111 016 115 015 HSI 069 009 060 004 072 010 076 011 Per 202 047 206 042 218 036 231 039 eer 026 006 027 005 028 004 030 005 1Means n18 standard deviation Mean in the same row with different superscript letters are significantly different P 005WG weight gain FCR feed conversion rate HSI hepatosomatic index PER protein efficiency rate EER energy efficiency rate DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 6 9 was efficient in using different dietary amounts of carbohydrate or lipid ratios as energy sources when considering growth parameters In the hybrid catfish the CHOL ratios had an effect in energy metabolism terms The low lipid and high carbohydrateratios increased plasma and hepatic glucose and also the hepatic triglyceride levels in the studied hybrid Lundstedt et al 2004 obtained similar results for P corruscans according to which the incorporation of corn flour into diet significantly increased glycaemia and hepatic glucose The liver is the central organ that controls glycaemia via enzymes that exert the main catabolism and synthesis of energy substrates The metabolic capacity of reducing glycaemia is limited in carnivorous fish Caseras et al 2007 Hemre et al 2002 which can lead to poor carbohydrate absorption and also to an inefficient metabolism and energy profit from glucose No influence of the CHOL ratios was found on the protein content in white muscle liver and plasma However the highest glycaemia and plasma protein rates were low at 033 with a high 124 proteinCHO ratio respectively in the P fasciatum X L marmoratus hybrids Souza et al 2014 In our study although the fish fed the diet containing 15 TABLE III The plasma mgdL1 liver mgg1 and white muscle mgg1 compositions of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days The means in the same row with different superscripts are significantly different at P005 Parameters Carbohydrate lipid ratios1 045 100 166 285 White muscle mg g1 glucose 095 007 112 021 115 018 124 016 Triglycerides 120 025 113 038 110 028 107 025 Protein 3006 507 2480 492 2706 375 2666 408 Plasma mg ml1 glucose 057 005b 071 005b 077 003b 113 008a Triglycerides 177 010 165 017 162 021 130 018 Protein 383 033 410 053 382 035 404 059 Liver mg g1 glucose 1914 198b 1910 416b 2203 271ab 2359 265a Triglycerides 2631 141a 1815 242b 2168 228b 1966 194b Protein 9025 571 7745 687 8015 966 8260 485 1Means n18 standard deviation Mean in the same row with different superscript letters are significantly different P 005 TABLE IV Hepatic enzyme activity of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days The means in the same row with different superscripts are significantly different at P005 Parameters Carbohydrate lipid ratios1 045 100 166 285 Me 345 101 389 065 407 098 371 013 g6PD 3404 301b 4788 396ab 4909 881ab 6168 063a 1Means n18 standard deviation Mean in the same row with different superscript letters are significantly different P 005 DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 7 9 of corn starch CHOL 166 showed high hepatic glucose values this concentration did not influence glycaemia The livers of the fish fed this diet were probably able to metabolize circulating glucose after 24 h of fasting This effect was not found for the treatment that contained 20 corn starch CHOL 285 which probably characterises a liver capacity limit to recover glycaemia homeostasis Our findings revealed no differences in ME when hybrid catfish were fed at different CHOL ratios In agreement with our result lipogenic enzyme g6PD activity was more pronounced than the gluconeogenic enzyme ME in catfish Mystus nemurus Hamid et al 2011 g6PD activity was greater at a high CHOL ratio Jin et al 2014 reported an increase in both the g6PD enzyme expression and hepatic glycaemia levels after an intraperitoneal injection of glucose in rainbow trout of a weak lineage It is the main enzyme in the pentose phosphate pathway to generate NADPH as a reducing power for biosynthesis such as lipid synthesis and has been widely assessed to describe animals adaptation capacity to diets containing a high carbohydrate concentration g6PD and 6PgDH in the catfish liver did not respond to the dietary concentrations of protein carbohydrates and lipids Indeed the only response to be obtained has been found in fasted freshwater South America catfish Randia quelen Melo et al 2016 However this fish is considered an omnivorous species which means that the carbohydrate type appeared to influence the activities of liver enzymes to a greater extent greater g6PD activity has been observed in freshwater blunt snout breams Megalobrama amblycephala fed a glucose diet than those fed a dextrin or wheat starch diet at 2 h postprandial Ren et al 2015 Thus our findings suggest that carnivorous hybrid hepatic cells appear to use carbohydrate properly at lower CHOL ratios The increased metabolism activity for glucose at the cell level may lead to a drop in not only hepatic g6PD the ratelimiting enzyme in the pentosephosphate pathway but also in the production of both NADPH and ribose5phosphates the building blocks of nucleic acids via an initial breakdown of glucose in channel catfish Ictalurus punctatus Schroeter et al 2018 CONCLUSIONS By way of conclusion the present study suggests that a rise in the carbohydratelipid ratio 285 might lead to increasing glycaemia but does not improve either growth or the protein efficiency rate in hybrid catfish However the glycaemia effect was not observed in the diet with a CHOL of 166 which may be the most suitable carbohydrate and lipid inclusion level for this species CHO L285 also affected g6PD activities which somehow suggests that Pseudoplatystoma corruscansPseudoplatystoma fasciatum is unable to efficiently cope with high dietary carbohydrate levels The authors declare that there is no conflict of interest AUTHOR CONTRIBUTIONS rosa was the researcher responsible for the elaboration and leadership of the research Okamura Araújo and Costa conducted the field assay and laboratory analyses Paulino Fortes Silva and Pereira together with the other authors participated mainly in the draft paper statistical analysis and discussion of the results REFERENCES ALVAreZ MJ DÍeZ A LÓPeZBOTe C gALLegO M AND BAUTISTA JM 2000 Shortterm modulation of lipogenesis by macronutrients in rainbow trout Oncorhynchus mykiss hepatocytes Brit J Nutr 84 619 628 ARSLAN M RINCHARD J DABROWSKI K AND POrTeLLA MArIA C 2008 effects of Different Dietary Lipid Sources on the Survival growth and Fatty Acid Composition of South American Catfish Pseudoplatystoma fasciatum Surubim Juveniles J World Aquacult Soc 39 5161 DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 8 9 BeeLeN r BOYD B gArAVeLLO JC PAVANeLLI gC AND AINSWORTH AJ 2003 A cytochemical light and electron microscopic study of the peripheral blood leucocytes of hybrid surubim catfish Pseudoplatystoma corruscans x Pseudoplatystoma fasciatum Comp Clin Path 12 6168 BerNArDeS CL NAVArrO rD gUerrASANTOS B AND FOrTeSSILVA r 2016 effects of dietary carbohydratelipid ratios on growth body composition and nutrient utilization of hybrid catfish Pseudoplatystoma reticulatum x Leiarius marmoratus rev Colom Cienc Pecua 29 5865 BICUDO ÁJDA BOrgHeSI r DAIrIKI JK SADO rY AND CYrINO JeP 2012 Performance of juveniles of Pseudoplatystoma fasciatum fed graded levels of corn gluten meal Pesq Agropec Bras 47 838845 BrADFOrD MM 1976 A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of proteindye binding Anal Biochem 72 248254 CAMPOS P MArTINO rC AND TrUgO LC 2006 Amino acid composition of Brazilian surubim fish Pseudoplatystoma coruscans fed diets with different levels and sources of fat Food Chem 96 126130 CASerAS A MeTÓN I VIVeS C egeA M FerNÁNDeZ F AND BAANANTe IV 2007 Nutritional regulation of glucose6phosphatase gene expression in liver of the gilthead sea bream Sparus aurata Brit J Nutr 88 607 614 CUI XJ ZHOU QC LIANg HO YANg J AND ZHAO LM 2010 effects of dietary carbohydrate sources on the growth performance and hepatic carbohydrate metabolic enzyme activities of juvenile cobia rachycentron canadum Linnaeus Aquaculture research 42 99107 eUCLYDeS rF 1997 Manual de utilização do programa SAeg Sistema para análises estatísticas e genéticas 2ª ed Universidade Federal de Viçosa Viçosa GRAEVE K SCHAEWEN A AND SCHEIBE R 1994 Purification characterization and cDNA sequence of glucose6phosphate dehydrogenase from potato Solatium tuberosum L Plant J 5 353361 HAMID NKA MAHAYAT M AND HASHIM r 2011 Utilization of different carbohydrate sources and starch forms by bagrid catfish Mystus nemurus Cuv Val Aquacult Nutr 17 e10e18 HeMre gI MOMMSeN TP AND KrOgDAHL Å 2002 Carbohydrates in fish nutrition effects on growth glucose metabolism and hepatic enzymes Aquacult Nutr 8 175 194 JIN J MÉDALe F KAMALAM BS AgUIrre P VÉrON V AND PANSerAT S 2014 Comparison of glucose and Lipid Metabolic gene expressions between Fat and Lean Lines of rainbow Trout after a glucose Load PLOS ONe 9 e105548 KAMALAM BS MeDALe F AND PANSerAT S 2017 Utilisation of dietary carbohydrates in farmed fishes New insights on influencing factors biological limitations and future strategies Aquaculture 467 327 KrOgDAHL Å HeMre gI AND MOMMSeN TP 2005 Carbohydrates in fish nutrition digestion and absorption in postlarval stages Aquacult Nutr 11 103122 LeONArDO AFg rOMAgOSA e BOreLLA MI AND BATLOUNI Sr 2004 Induced spawning of hatchery raised Brazilian catfish cachara Pseudoplatystoma fasciatum Linnaeus 1766 Aquaculture 240 451461 LUNDSTeDT LM MeLO JFB AND MOrAeS g 2004 Digestive enzymes and metabolic profile of Pseudoplatystoma corruscans Teleostei Siluriformes in response to diet composition Comp Biochem Phys B 137 331339 MArTINO rC CYrINO JeP POrTZ L AND TrUgO LC 2002a effect of dietary lipid level on nutritional performance of the surubim Pseudoplatystoma coruscans Aquaculture 209 209218 MArTINO rC CYrINO JeP POrTZ L AND TrUgO LC 2002b Performance and fatty acid composition of surubim Pseudoplatystoma coruscans fed diets with animal and plant lipids Aquaculture 209 233246 MArTINO rC CYrINO JeP POrTZ L AND TrUgO LC 2005 Performance carcass composition and nutrient utilization of surubim Pseudoplatystoma coruscans Agassiz fed diets with varying carbohydrate and lipid levels Aquacult Nutr 11 131137 MeLO JFB LUNDSTeDT LM INOUe LAK MeTÓN I BAANANTe IV AND MOrAeS g 2016 glycolysis and gluconeogenesis in the liver of catfish fed with different concentrations of proteins lipids and carbohydrates Arq Bras Med Vet Zoo 68 12511258 NrC 2011 Nutrient requirements of Fish and Shrimp Washington DC The National Academies Press 392 p PAPOUTSOgLOU eS AND LYNDON Ar 2006 Digestive enzymes along the alimentary tract of the parrotfish Sparisoma cretense J Fish Biol 69 130140 PAULINO rr PereIrA rT FONTeS TV OLIVATeLeS A PereS H CArNeIrO DJ AND rOSA PV 2018 Optimal dietary linoleic acid to linolenic acid ratio improved fatty acid profile of the juvenile tambaqui Colossoma macropomum Aquaculture 488 916 reN M HABTeTSION HM XIe J LIU B ZHOU Q ge X PAN L AND CHeN r 2015 effects of dietary carbohydrate source on growth performance diet digestibility and liver glucose enzyme activity in blunt snout bream Megalobrama amblycephala Aquaculture 438 7581 DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 9 9 SCHrOeTer JC PeTerSON BC BLeDSOe J LI M AND SMALL BC 2018 Targeted gene expression panels and microbiota analysis provide insight into the effects of alternative production diet formulations on channel catfish nutritional physiology Aquaculture 489 4655 SOUZA SA SOUZA rC CAMPeCHe DFB CAMPOS rML AND MeLO JFB 2014 relação proteínacarboidrato no desempenho e no metabolismo de híbridos de Pseudoplatystoma fasciatum fêmea X Leiarius marmoratus macho Arq Bras Med Vet Zoo 66 879886 SPINA J BrIgHT HJ AND rOSeNBLOOM J 1970 Purification and properties of Lmalic enzyme from escherichia coli Biochemistry 9 37943801 TRANTU LC HIEN TTT BOSMA RH HEINSBROEK LTN VERRETH JAJ AND SCHRAMA JW 2017 Effect of ingredient particle sizes and dietary viscosity on digestion and faecal waste of striped catfish Pangasianodon hypophthalmus Aquacult Nutr 24 961969 WANG JT JIANG YD HAN T LI XY WANG Y AND LIU YJ 2016 Effects of Dietary CarbohydratetoLipid Ratios on Growth and Body Composition of Orangespotted grouper epinephelus coioides N Am J Aquacult 79 17 WANG LN LIU WB LU KL XU WN CAI DS ZHANG CN AND QIAN Y 2014 effects of dietary carbohydrate lipid ratios on nonspecific immune responses oxidative status and liver histology of juvenile yellow catfish Pelteobagrus fulvidraco Aquaculture 426427 4148 ZHOU C ge X NIU J LIN H HUANg Z AND TAN X 2015 effect of dietary carbohydrate levels on growth performance body composition intestinal and hepatic enzyme activities and growth hormone gene expression of juvenile golden pompano Trachinotus ovatus Aquaculture 437 390397 Resenha Crítica Respostas metabólicas às proporções de carboidratos e lipídios na dieta neotropical Introdução Em relação aos tipos de alimentos os que contém carboidratos em sua concentração são mais baratos o que sugere sua maior utilização na produção e criação de peixes Para saúde e bom desempenho do animal concentrações equilibradas de carboidratos e lipídios deve ser incrementado como fonte de alimento Isso está relacionado à índices equilibrados de proteínas e lipídios hepático musculares Entretanto a ingestão de carboidratos em peixes carnívoros ainda é meio desconhecida no que diz respeito a sua digestão absorção e processos metabólicos Essa relação que se refere ao processo de ingestão à digestão está diretamente ligada a espécie do peixe e a concentração de carboidrato oferecida como alimento Para esses tipos de peixes o mais indicado é a ingestão de lipídios isso devido sua limitação para digestão de carboidratos Sendo assim o perfil metabólico reflete exatamente a dieta proposta para esses animais Existe uma relação entre a concentração de carboidrato proposta e sua quantidade segundo o artigo o aumento de um e diminuição do outro sugere aumento de gliconeogênese e diminuição de atividade glicolítica Sendo assim analisar os efeitos da quantidade e concentração de dietas que envolvam carboidratos eou lipídios é importante Objetivo Avaliar o efeito de diversos tipos de carboidratos e lipídios em relação ao desempenho metabólico de peixes Amostra 168 peixes foram alimentados duas vezes ao dia até sua saciedade ser alcançada Estudo realizado na Aquacultura Estação da Universidade federal de Lavras Minais Gerais Aprovação de Comitê de Ética Método Os animais foram distribuídos em 24 tanques de 100 L cada Os tanques continham água renovada a cada 3 vezes ao dia Esses tanques continham filtro de areia filtro ultravioleta e filtro biológico A temperatura da água foi mantida em 28º O oxigênio disponibilizado para os animais foi medido através de um oxímetro digital portátil O pH da água foi medido semanalmente através de um kit comercial Antes do experimento os animais foram adaptados em um período de 15 dias Utilizouse 4 blocos com quantidades distintas de concentração de carboidrato para assim poder analisar posteriormente as adaptações promovidas pela ingesta 1 bloco 045 2 bloco 1 3 bloco 166 4 bloco 285 Após 60 dias parâmetros metabólicos e estruturais foram analisados Parâmetros estruturais Tamanho estrutural Parâmetros metabólicos Concentração de proteína glicose e triglicerídeos plasmáticos Atividade hepática e muscular ácido málico glicose 6fosfato e enzimas desidrogenase Resultados Nenhuma mortalidade de peixe foi encontrada durante o experimento Não houve diferença significativa relacionada ao tamanho estrutural na concentração de proteínas e triglicerídeos plasmáticos e musculares após o experimento Entretanto no bloco em que a quantidade de carboidrato foi de 285 demonstrou maiores concentrações de glicose plasmática e hepática Já no bloco de 045 a quantidade que demonstrou concentração elevada foi de triglicérides hepáticos ConclusãoCrítica ao artigo Não houve alteração metabólica significativa nos animais a partir do experimento Mudanças metabólicas encontradas em cada animal depende muito da biodisponibilidade de cada um em absorver e processar o que lhe foi ofertado além da condição de cada experimento em si O que sugere então que aumento de quantidade e concentração de carboidratos e lipídeos em uma dieta podem sugerir aumento da glicemia e concentrações de proteína entretanto não sugerem alterações estruturais REFERÊNCIA OKAMURA D FORTESSILVA R PAULINO RR ARAÚJO FG COSTA DV PEREIRA RT ROSA PV Respostas metabólicas às proporções de carboidratos e lipídios na dieta neotropical Anais da Academia Brasileira de Ciências 914 2019
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Anais da Academia Brasileira de Ciências 2019 914 e20190075 Annals of the Brazilian Academy of Sciences Printed version ISSN 00013765 Online version ISSN 16782690 httpdxdoiorg10159000013765201920190075 wwwscielobraabc wwwfbcomaabcjournal An Acad Bras Cienc 2019 914 AgrArIAN SCIeNCeS Metabolic responses to dietary carbohydratelipids ratios in neotropical hybrid catfish Pseudoplatystoma corruscansPseudoplatystoma fasciatum DANIEL OKAMURA1 RODRIGO FORTESSILVA2 RENAN R PAULINO 1 FELIPE G DE ARAÚJO3 DIEGO V DA COSTA4 RAQUEL T PEREIRA1 and PRISCILA V ROSA1 1Department of Animal Science Federal University of Lavras UFLA Aquenta Sol sn Campus Universitário 37200000 Lavras Mg Brazil 2Department of Animal Science Federal University of Viçosa UFV Av Peter Henry rolfs sn Campus Universitário 36570900 Viçosa Mg Brazil 3Department of Animal Science Federal rural University of Pernambuco UFrPe UAg Av Bom Pastor sn Boa Vista 55292270 garanhuns Pe Brazil 4Institute of Agricultural Sciences ICA University of Minas gerais UFMg Av Universitária 1000 Bairro Universitário 39404547 Montes Claros Mg Brazil Manuscript received on January 24 2019 accepted for publication on May 10 2019 How to cite OKAMUrA D FOrTeSSILVA r PAULINO rr ArAÚJO Fg COSTA DV PereIrA rT AND ROSA PV 2019 Metabolic responses to dietary carbohydratelipids ratios in neotropical hybrid catfish Pseudoplatystoma corruscansPseudoplatystoma fasciatum An Acad Bras Cienc 91 e20190075 DOI 10159000013765201920190075 Abstract The purpose of this study was to assess the effect of different carbohydrate and lipid CHOL ratios on the performance and energy metabolism of hybrid catfish Pseudoplatystoma corruscans Pseudoplatystoma fasciatum One hundred and sixtyeight fish 1049165 g BW were distributed into 24 100L tanks The experiment was performed using a randomised block design with four CHOL ratios 045 100 166 and 285 and six replications growth performance protein content glucose and triglycerides in plasma muscle and liver and the hepatic activity of the malic and glucose6phosphate dehydrogenase G6PD enzymes were measured after 60 feeding days No significant differences were observed in growth performance protein and triglycerides of plasma and muscle hepatic protein or malic enzyme activity CHOL 285 provided the highest plasma and liver glucose while CHOL 045 showed high liver triglycerides A linear effect was found for the hepatic activity of G6PD and dietary starch levels r2093 The hepatic activity of g6PD was greater in CHOL285 than for CHOL 045 In conclusion the limit of dietary starch inclusion and lipids were 15 and 900 respectively CHOL 166 and this level did not affect the energy metabolism of the juvenile hybrid catfish Key words enzymes activity hepatic function intermediary metabolism hybrid catfish Surubim Correspondence to Priscila Vieira rosa Email prisciladzouflabr OrCID httpsorcidorg000000020052079X INTRODUCTION Carbohydratecontaining feedstuffs are less expensive than other nutrients in diet and directly influence economic fish production viability Papoutsoglou and Lyndon 2006 Kamalam et al 2017 Although no nutritional requirements of carbohydrates exist in these animals diets their inclusion at appropriate levels can ensure better efficiency in utilising other nutrients by contributing positively in the diet extrusion process and acting DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 2 9 as an abundant energy source for lipids and proteins Krogdahl et al 2005 NrC 2011 Zhou et al 2015 Kamalam et al 2017 However carbohydrate use in a carnivorous fish diet is complex and not well known for digestion absorption and metabolism processes Krogdahl et al 2005 Such processes depend on the relationships between fish species and carbohydrate type or content Krogdahl et al 2005 Kamalam et al 2017 These species are more efficient in using lipids as an energy source given their metabolisms limited capacity to reduce glycaemia Caseras et al 2007 Hemre et al 2002 The metabolic profile and its pathways are the reflex from diet regardless of performance or muscle and liver function Carbohydrate lipids CHOL ratios of around 131 produce the best growth performance in hybrid catfish Pseudoplatystoma reticulatum x Leiarius marmoratus Bernardes et al 2016 One of the statements about fish growth variation is the relation between carbohydrate levels and their effects on diet viscosity and digestion which can in turn lead to a change in nutrient use TranTu et al 2017 Moreover increased dietary protein and fewer carbohydrates and lipids attenuate glycolytic activity and induce hepatic gluconeogenesis in South American catfish Rhamdia quelen Melo et al 2016 The liver histology was normal in the yellow catfish Pelteobagrus fulvidraco fed the 245 363 and 558 CHOL diets The hepatic steatosis and hepatocyte apoptosis rates increased in the yellow catfish fed with 111 and 167 CHOL than 245 363 and 558 CHOL Wang et al 2014 Thus it is imperative for modern aquaculture to define the optimum dietary ratio between nonprotein energy sources such as carbohydrates and lipids Wang et al 2016 Pseudoplatystoma corruscans and Pseudoplatystoma fasciatum are carnivorous fish and important species with commercial characteristics produced in South America Martino et al 2002a Leonardo et al 2004 along with their hybrids P corruscansP fasciatum Beelen et al 2003 Pseudoplatystoma spp is able to adjust amylase activity in accordance with dietary carbohydrate levels Lundstedt et al 2004 Both plasma glucose and plasma protein concentrations lower when corn gluten meal levels increase in Pfasciatum Bicudo et al 2012 All these South American catfish are species of the same family Pimelodidae However very little is known about this hybrid metabolism function in relation to the nutrition effect In some carnivorous species eg Oncorhynchus mykiss Rachycentron canadum and Mystus nemurus glucose stimulates glucose6phosphate dehydrogenase g6PD and malic enzyme Me activities Alvarez et al 2000 Cui et al 2010 Hamid et al 2011 which are the main enzymes responsible for producing NADPH which is required for lipid synthesis Thus the quantification of metabolites in different tissues and enzyme activity in relation to energy metabolism are important for understanding the use of dietary carbohydrates to optimise growth spare proteins and avoid economical waste while farming species The present work was conducted to assess the influence of different CHOL ratio on the energy metabolism of hybrids Pseudoplatystoma corruscansPseudoplatystoma fasciatum by estimating performance protein concentration glucose and triglycerides in plasma muscle and liver and the hepatic activity of both Me and g6PD MATERIALS AND METHODS ANIMAL HOUSINg The growth trial was performed at the Aquaculture Station of the Federal University of Lavras UFLA Minas gerais Brazil and was approved by the UFLAs ethics Committee on Animal Use and was certified by Protocol 02812 Juvenile catfish Pseudoplatystoma spp were obtained from a farm in Minas gerais Brazil One hundred and DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 3 9 sixtyeight fish initial body weight of 1049165 g meanSD were distributed into 24 100L tanks 7 fish per tank Fish were fed twice until apparent satiation The system comprised a 5μm particle sand filter an ultraviolet filter and a biological filter The water flux in the experimental boxes was sufficient to renew water 3 timeshour During the experiment the water temperature was maintained at 2801oC Dissolved oxygen was measured daily by a portable digital oxymeter F1550A Bernauer Indaial SC Brazil with average values of 5109 mg L1 Ammonia nitrite and pH were measured weekly using a commercial kit Labcon Test Alcon Camboriú SC Brazil with average values of 00090002 ppm 0010001 ppm and 7301 respectively Fish were submitted to a 15day adaptation period before the experiment began eXPerIMeNTAL DIeTS Four isoproteic 460 g kg1 crude protein and isoenergetic 3500 kcal kg1 diets were formulated to contain the dietary carbohydrate and lipid ratios CHOL 045 100 166 and 285 Table I Canola oil was considered herein because the parietal P fasciatumis well able to use plant oils than fish oil to transform linoleate into arachidonate Arslan et al 2008 Moreover plant oil is used by P coruscans Martino et al 2002b Dietary ingredients were mixed in a paddle mixer Inbramaq São Paulo Brazil with 20 of water pelletized 4 mm and dried for 12 h in an oven at 50C Diets were stored in a freezer 5C until used eXPerIMeNTAL DeSIgN After the adaptation period fish were fed the experimental diets 4 CHOL ratios and 6 replies twice a day at 0800h and 1800h The amount of diet was provided according to apparent animal satiety The delivered amount of diets was quantified daily by weighing before and after the feeding process The trial lasted 60 days At the end of the experimental period fish were starved for 24h weighed and then kept in anaesthetic solution until opercular movement had completely ceased 08 mll1 2phenoxyethanol One millilitre of blood per cardiac punch was also sampled using a heparinised syringe 1 ml needle size of 21g 12 08x40 Blood was centrifuged at 3000 rpm for 10 minutes before isolating plasma by a Pasteur micropipette After this procedure fish were decapitated at the skull insertion point Then the liver and white dorsal muscle samples approximately 3g were collectedfrom each juvenile The liver TABLE I Composition of the experimental diets Ingredients Inclusion levels Fish meal 1 5717 5217 4800 4400 gelatine 500 500 600 700 Albumin 700 700 800 800 Casein 300 600 700 850 Canola oil 1100 1000 900 700 Starch2 500 1000 1500 2000 Dicalcium phosphate 150 150 150 150 Mineral andvitamin 3 030 030 030 030 Antioxidant 003 003 003 003 Cellulose 500 400 317 260 Inert 500 400 200 100 Total 10000 10000 10000 10000 Carbohydrate lipid ratio 045 100 166 285 Approximate composition g100 g dry matter CP4 467 463 463 462 Ce5 kcalkg 1 3525 3548 3595 3550 1Salmon meal 67 crude protein Total Alimentos 2Carbohydrate source maize raw starch Pachá Alimentos 3Composition per kilogram of the mineral and vitamin supplement 1500 IU vitamin A 20 mg vitamin B1 15 mg vitamin B2 1000 IU vitamin B3 10 mcg vitamin B12 25 mg vitamin e 120 mg vitamin PP 2000 mg choline 80 mg calcium pantothenate 2 mg folic acid 80 mg manganese 24 mg iron 50 mg zinc 8 mg copper 3 mg iodine 010 mg selenium and 170 mg BHT 4CP crude protein 5Ce crude energy DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 4 9 was weighed to calculate the hepatossomatic index Samples were stored at 80oC for further analyses The following parameters were calculated Hepatosomatic Index HSI liver weightfish weight100 weight gain WG final weight first weight feed conversion ratio FCR ingested rateweight gain protein efficiency rate PER weight gainingested crude protein and energy efficiency rate EER weight gainingested crude energy Muscle and liver assessments The liver and muscle homogenates were obtained according to Paulino et al 2018 with slight modifications The white muscle and liver samples were homogenized in icecold deionised water and centrifuged 13400 g 10 min 4ºCThe resultant supernatant was collected and stored at 80ºC glucose and triglyceride levels were measured with commercial kits glucose HK Liquiform Cat 85 and Triglycerides Liquiform Cat 87 Soluble protein content was determined according to Bradford 1976 with bovine serum albumin solution Sigma used as a standard The specific activity of both malic enzyme Me eC11140 and glucose6phosphate dehydrogenase g6PD eC11149 was estimated following the methodology outlined in graeve et al 1994 and Spina et al 1970 respectively Hepatic tissue was macerated with a pistil in an icebath After maceration the sample was put in an ultrasonic bath for 10 minutes before being centrifuged at 10000 g and 4oC for 30 minutes before collecting the supernatant The homogenate of supernatants was used to estimate specific G6PD activity All the enzymatic activities are shown as Umg1 protein STATISTICAL ANALYSIS All the obtained data were analyzed using the SAeg statistical package proposed by euclydes 1997 A data analysis was done by a oneway analysis of variance ANOVA after testing for normality and homogeneity of variances with the ShapiroWilk test and the Levene test respectively Significant differences among the means were determined by a StudentNewmanKeuls test A regression equation was obtained to describe the glucose6phosphate dehydrogenase pattern RESULTS No fish mortality was recorded during the experimental period After the 60 days that the trial lasted no growth performance parameter eg weight gain feed conversion rate Hepatosomatic Index protein efficiency rate and energy efficiency rate was significantly affected P005 by the dietary carbohydratelipid CHOL ratio Table II The concentrations of glucose triglycerides and protein in the white muscle of juveniles did not change P005 Table III However CHOL had an effect on glucose in both plasma and the liver as did triglycerides on hepatic tissue P005 The highest plasma and liver glucose contents were found in the fish fed the diet CHOL 285 The catfish juveniles fed the diet CHOL045 showed higher concentrations of hepatic triglycerides compared to other CHOL ratios P005 The plasma and liver protein concentrations were not affected by diet P005 No significant differences were observed for the hepatic activity of Me P005 However g6PD activity was greater in the treatment CHOL 285 compared to CHOL045 P005 Table IV A linear effect was found for G6PD activity which increased with a rising carbohydrate concentration in diet Figure 1 DISCUSSION In this study the fish fed different CHOL ratio in their diet did not show any significant differences in their performance The range within which DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 5 9 dietary carbohydrates can be used to meet energy requirements vastly varies between and even within species depending on many factors which have been categorised into those also related to biology and experimental conditions Kamalam et al 2017 For example the inclusion of dietary soluble polysaccharides does not affect growth performance nor the feed conversion ratio in African catfish Clarias gariepinus Martino et al 2005 found that replacing the energy obtained from soya bean oil in diet with corn flour did not affect the specific growth rate the feed conversion rate total consumption protein and the hepatosomatic index of the parietal Pseudoplatystoma ssp On the other hand the 4 8 and 12 increases in soya bean oil led to enhanced nitrogen fixation in a P coruscans carcass Campos et al 2006 The feed conversion ratio and daily feed consumption showed a marked drop in the inverse proportion to the lipid level in diet in P coruscans Martino et al 2002b In short the present study revealed that the hybrid P corruscansP fasciatum Figure 1 Linear regression of the hepatic glucose6phosphate dehydrogenase activity of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days n18 TABLE II Growth performance feed utilisation and Hepatosomatic Index of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days Lack of superscript letters in the same line implies no differences in data at P005 Parameters Carbohydrate lipid ratios1 045 100 166 285 WG 11742 2749 11943 2467 12543 2100 13297 2281 FCr 111 027 118 022 111 016 115 015 HSI 069 009 060 004 072 010 076 011 Per 202 047 206 042 218 036 231 039 eer 026 006 027 005 028 004 030 005 1Means n18 standard deviation Mean in the same row with different superscript letters are significantly different P 005WG weight gain FCR feed conversion rate HSI hepatosomatic index PER protein efficiency rate EER energy efficiency rate DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 6 9 was efficient in using different dietary amounts of carbohydrate or lipid ratios as energy sources when considering growth parameters In the hybrid catfish the CHOL ratios had an effect in energy metabolism terms The low lipid and high carbohydrateratios increased plasma and hepatic glucose and also the hepatic triglyceride levels in the studied hybrid Lundstedt et al 2004 obtained similar results for P corruscans according to which the incorporation of corn flour into diet significantly increased glycaemia and hepatic glucose The liver is the central organ that controls glycaemia via enzymes that exert the main catabolism and synthesis of energy substrates The metabolic capacity of reducing glycaemia is limited in carnivorous fish Caseras et al 2007 Hemre et al 2002 which can lead to poor carbohydrate absorption and also to an inefficient metabolism and energy profit from glucose No influence of the CHOL ratios was found on the protein content in white muscle liver and plasma However the highest glycaemia and plasma protein rates were low at 033 with a high 124 proteinCHO ratio respectively in the P fasciatum X L marmoratus hybrids Souza et al 2014 In our study although the fish fed the diet containing 15 TABLE III The plasma mgdL1 liver mgg1 and white muscle mgg1 compositions of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days The means in the same row with different superscripts are significantly different at P005 Parameters Carbohydrate lipid ratios1 045 100 166 285 White muscle mg g1 glucose 095 007 112 021 115 018 124 016 Triglycerides 120 025 113 038 110 028 107 025 Protein 3006 507 2480 492 2706 375 2666 408 Plasma mg ml1 glucose 057 005b 071 005b 077 003b 113 008a Triglycerides 177 010 165 017 162 021 130 018 Protein 383 033 410 053 382 035 404 059 Liver mg g1 glucose 1914 198b 1910 416b 2203 271ab 2359 265a Triglycerides 2631 141a 1815 242b 2168 228b 1966 194b Protein 9025 571 7745 687 8015 966 8260 485 1Means n18 standard deviation Mean in the same row with different superscript letters are significantly different P 005 TABLE IV Hepatic enzyme activity of the juvenile Pseudoplatystoma ssp fed the experimental diets for 60 days The means in the same row with different superscripts are significantly different at P005 Parameters Carbohydrate lipid ratios1 045 100 166 285 Me 345 101 389 065 407 098 371 013 g6PD 3404 301b 4788 396ab 4909 881ab 6168 063a 1Means n18 standard deviation Mean in the same row with different superscript letters are significantly different P 005 DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 7 9 of corn starch CHOL 166 showed high hepatic glucose values this concentration did not influence glycaemia The livers of the fish fed this diet were probably able to metabolize circulating glucose after 24 h of fasting This effect was not found for the treatment that contained 20 corn starch CHOL 285 which probably characterises a liver capacity limit to recover glycaemia homeostasis Our findings revealed no differences in ME when hybrid catfish were fed at different CHOL ratios In agreement with our result lipogenic enzyme g6PD activity was more pronounced than the gluconeogenic enzyme ME in catfish Mystus nemurus Hamid et al 2011 g6PD activity was greater at a high CHOL ratio Jin et al 2014 reported an increase in both the g6PD enzyme expression and hepatic glycaemia levels after an intraperitoneal injection of glucose in rainbow trout of a weak lineage It is the main enzyme in the pentose phosphate pathway to generate NADPH as a reducing power for biosynthesis such as lipid synthesis and has been widely assessed to describe animals adaptation capacity to diets containing a high carbohydrate concentration g6PD and 6PgDH in the catfish liver did not respond to the dietary concentrations of protein carbohydrates and lipids Indeed the only response to be obtained has been found in fasted freshwater South America catfish Randia quelen Melo et al 2016 However this fish is considered an omnivorous species which means that the carbohydrate type appeared to influence the activities of liver enzymes to a greater extent greater g6PD activity has been observed in freshwater blunt snout breams Megalobrama amblycephala fed a glucose diet than those fed a dextrin or wheat starch diet at 2 h postprandial Ren et al 2015 Thus our findings suggest that carnivorous hybrid hepatic cells appear to use carbohydrate properly at lower CHOL ratios The increased metabolism activity for glucose at the cell level may lead to a drop in not only hepatic g6PD the ratelimiting enzyme in the pentosephosphate pathway but also in the production of both NADPH and ribose5phosphates the building blocks of nucleic acids via an initial breakdown of glucose in channel catfish Ictalurus punctatus Schroeter et al 2018 CONCLUSIONS By way of conclusion the present study suggests that a rise in the carbohydratelipid ratio 285 might lead to increasing glycaemia but does not improve either growth or the protein efficiency rate in hybrid catfish However the glycaemia effect was not observed in the diet with a CHOL of 166 which may be the most suitable carbohydrate and lipid inclusion level for this species CHO L285 also affected g6PD activities which somehow suggests that Pseudoplatystoma corruscansPseudoplatystoma fasciatum is unable to efficiently cope with high dietary carbohydrate levels The authors declare that there is no conflict of interest AUTHOR CONTRIBUTIONS rosa was the researcher responsible for the elaboration and leadership of the research Okamura Araújo and Costa conducted the field assay and laboratory analyses Paulino Fortes Silva and Pereira together with the other authors participated mainly in the draft paper statistical analysis and discussion of the results REFERENCES ALVAreZ MJ DÍeZ A LÓPeZBOTe C gALLegO M AND BAUTISTA JM 2000 Shortterm modulation of lipogenesis by macronutrients in rainbow trout Oncorhynchus mykiss hepatocytes Brit J Nutr 84 619 628 ARSLAN M RINCHARD J DABROWSKI K AND POrTeLLA MArIA C 2008 effects of Different Dietary Lipid Sources on the Survival growth and Fatty Acid Composition of South American Catfish Pseudoplatystoma fasciatum Surubim Juveniles J World Aquacult Soc 39 5161 DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 8 9 BeeLeN r BOYD B gArAVeLLO JC PAVANeLLI gC AND AINSWORTH AJ 2003 A cytochemical light and electron microscopic study of the peripheral blood leucocytes of hybrid surubim catfish Pseudoplatystoma corruscans x Pseudoplatystoma fasciatum Comp Clin Path 12 6168 BerNArDeS CL NAVArrO rD gUerrASANTOS B AND FOrTeSSILVA r 2016 effects of dietary carbohydratelipid ratios on growth body composition and nutrient utilization of hybrid catfish Pseudoplatystoma reticulatum x Leiarius marmoratus rev Colom Cienc Pecua 29 5865 BICUDO ÁJDA BOrgHeSI r DAIrIKI JK SADO rY AND CYrINO JeP 2012 Performance of juveniles of Pseudoplatystoma fasciatum fed graded levels of corn gluten meal Pesq Agropec Bras 47 838845 BrADFOrD MM 1976 A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of proteindye binding Anal Biochem 72 248254 CAMPOS P MArTINO rC AND TrUgO LC 2006 Amino acid composition of Brazilian surubim fish Pseudoplatystoma coruscans fed diets with different levels and sources of fat Food Chem 96 126130 CASerAS A MeTÓN I VIVeS C egeA M FerNÁNDeZ F AND BAANANTe IV 2007 Nutritional regulation of glucose6phosphatase gene expression in liver of the gilthead sea bream Sparus aurata Brit J Nutr 88 607 614 CUI XJ ZHOU QC LIANg HO YANg J AND ZHAO LM 2010 effects of dietary carbohydrate sources on the growth performance and hepatic carbohydrate metabolic enzyme activities of juvenile cobia rachycentron canadum Linnaeus Aquaculture research 42 99107 eUCLYDeS rF 1997 Manual de utilização do programa SAeg Sistema para análises estatísticas e genéticas 2ª ed Universidade Federal de Viçosa Viçosa GRAEVE K SCHAEWEN A AND SCHEIBE R 1994 Purification characterization and cDNA sequence of glucose6phosphate dehydrogenase from potato Solatium tuberosum L Plant J 5 353361 HAMID NKA MAHAYAT M AND HASHIM r 2011 Utilization of different carbohydrate sources and starch forms by bagrid catfish Mystus nemurus Cuv Val Aquacult Nutr 17 e10e18 HeMre gI MOMMSeN TP AND KrOgDAHL Å 2002 Carbohydrates in fish nutrition effects on growth glucose metabolism and hepatic enzymes Aquacult Nutr 8 175 194 JIN J MÉDALe F KAMALAM BS AgUIrre P VÉrON V AND PANSerAT S 2014 Comparison of glucose and Lipid Metabolic gene expressions between Fat and Lean Lines of rainbow Trout after a glucose Load PLOS ONe 9 e105548 KAMALAM BS MeDALe F AND PANSerAT S 2017 Utilisation of dietary carbohydrates in farmed fishes New insights on influencing factors biological limitations and future strategies Aquaculture 467 327 KrOgDAHL Å HeMre gI AND MOMMSeN TP 2005 Carbohydrates in fish nutrition digestion and absorption in postlarval stages Aquacult Nutr 11 103122 LeONArDO AFg rOMAgOSA e BOreLLA MI AND BATLOUNI Sr 2004 Induced spawning of hatchery raised Brazilian catfish cachara Pseudoplatystoma fasciatum Linnaeus 1766 Aquaculture 240 451461 LUNDSTeDT LM MeLO JFB AND MOrAeS g 2004 Digestive enzymes and metabolic profile of Pseudoplatystoma corruscans Teleostei Siluriformes in response to diet composition Comp Biochem Phys B 137 331339 MArTINO rC CYrINO JeP POrTZ L AND TrUgO LC 2002a effect of dietary lipid level on nutritional performance of the surubim Pseudoplatystoma coruscans Aquaculture 209 209218 MArTINO rC CYrINO JeP POrTZ L AND TrUgO LC 2002b Performance and fatty acid composition of surubim Pseudoplatystoma coruscans fed diets with animal and plant lipids Aquaculture 209 233246 MArTINO rC CYrINO JeP POrTZ L AND TrUgO LC 2005 Performance carcass composition and nutrient utilization of surubim Pseudoplatystoma coruscans Agassiz fed diets with varying carbohydrate and lipid levels Aquacult Nutr 11 131137 MeLO JFB LUNDSTeDT LM INOUe LAK MeTÓN I BAANANTe IV AND MOrAeS g 2016 glycolysis and gluconeogenesis in the liver of catfish fed with different concentrations of proteins lipids and carbohydrates Arq Bras Med Vet Zoo 68 12511258 NrC 2011 Nutrient requirements of Fish and Shrimp Washington DC The National Academies Press 392 p PAPOUTSOgLOU eS AND LYNDON Ar 2006 Digestive enzymes along the alimentary tract of the parrotfish Sparisoma cretense J Fish Biol 69 130140 PAULINO rr PereIrA rT FONTeS TV OLIVATeLeS A PereS H CArNeIrO DJ AND rOSA PV 2018 Optimal dietary linoleic acid to linolenic acid ratio improved fatty acid profile of the juvenile tambaqui Colossoma macropomum Aquaculture 488 916 reN M HABTeTSION HM XIe J LIU B ZHOU Q ge X PAN L AND CHeN r 2015 effects of dietary carbohydrate source on growth performance diet digestibility and liver glucose enzyme activity in blunt snout bream Megalobrama amblycephala Aquaculture 438 7581 DANIeL OKAMUrA et al DIeTArY CArBOHYDrATe LIPIDS rATIOS ON CATFISH An Acad Bras Cienc 2019 914 e20190075 9 9 SCHrOeTer JC PeTerSON BC BLeDSOe J LI M AND SMALL BC 2018 Targeted gene expression panels and microbiota analysis provide insight into the effects of alternative production diet formulations on channel catfish nutritional physiology Aquaculture 489 4655 SOUZA SA SOUZA rC CAMPeCHe DFB CAMPOS rML AND MeLO JFB 2014 relação proteínacarboidrato no desempenho e no metabolismo de híbridos de Pseudoplatystoma fasciatum fêmea X Leiarius marmoratus macho Arq Bras Med Vet Zoo 66 879886 SPINA J BrIgHT HJ AND rOSeNBLOOM J 1970 Purification and properties of Lmalic enzyme from escherichia coli Biochemistry 9 37943801 TRANTU LC HIEN TTT BOSMA RH HEINSBROEK LTN VERRETH JAJ AND SCHRAMA JW 2017 Effect of ingredient particle sizes and dietary viscosity on digestion and faecal waste of striped catfish Pangasianodon hypophthalmus Aquacult Nutr 24 961969 WANG JT JIANG YD HAN T LI XY WANG Y AND LIU YJ 2016 Effects of Dietary CarbohydratetoLipid Ratios on Growth and Body Composition of Orangespotted grouper epinephelus coioides N Am J Aquacult 79 17 WANG LN LIU WB LU KL XU WN CAI DS ZHANG CN AND QIAN Y 2014 effects of dietary carbohydrate lipid ratios on nonspecific immune responses oxidative status and liver histology of juvenile yellow catfish Pelteobagrus fulvidraco Aquaculture 426427 4148 ZHOU C ge X NIU J LIN H HUANg Z AND TAN X 2015 effect of dietary carbohydrate levels on growth performance body composition intestinal and hepatic enzyme activities and growth hormone gene expression of juvenile golden pompano Trachinotus ovatus Aquaculture 437 390397 Resenha Crítica Respostas metabólicas às proporções de carboidratos e lipídios na dieta neotropical Introdução Em relação aos tipos de alimentos os que contém carboidratos em sua concentração são mais baratos o que sugere sua maior utilização na produção e criação de peixes Para saúde e bom desempenho do animal concentrações equilibradas de carboidratos e lipídios deve ser incrementado como fonte de alimento Isso está relacionado à índices equilibrados de proteínas e lipídios hepático musculares Entretanto a ingestão de carboidratos em peixes carnívoros ainda é meio desconhecida no que diz respeito a sua digestão absorção e processos metabólicos Essa relação que se refere ao processo de ingestão à digestão está diretamente ligada a espécie do peixe e a concentração de carboidrato oferecida como alimento Para esses tipos de peixes o mais indicado é a ingestão de lipídios isso devido sua limitação para digestão de carboidratos Sendo assim o perfil metabólico reflete exatamente a dieta proposta para esses animais Existe uma relação entre a concentração de carboidrato proposta e sua quantidade segundo o artigo o aumento de um e diminuição do outro sugere aumento de gliconeogênese e diminuição de atividade glicolítica Sendo assim analisar os efeitos da quantidade e concentração de dietas que envolvam carboidratos eou lipídios é importante Objetivo Avaliar o efeito de diversos tipos de carboidratos e lipídios em relação ao desempenho metabólico de peixes Amostra 168 peixes foram alimentados duas vezes ao dia até sua saciedade ser alcançada Estudo realizado na Aquacultura Estação da Universidade federal de Lavras Minais Gerais Aprovação de Comitê de Ética Método Os animais foram distribuídos em 24 tanques de 100 L cada Os tanques continham água renovada a cada 3 vezes ao dia Esses tanques continham filtro de areia filtro ultravioleta e filtro biológico A temperatura da água foi mantida em 28º O oxigênio disponibilizado para os animais foi medido através de um oxímetro digital portátil O pH da água foi medido semanalmente através de um kit comercial Antes do experimento os animais foram adaptados em um período de 15 dias Utilizouse 4 blocos com quantidades distintas de concentração de carboidrato para assim poder analisar posteriormente as adaptações promovidas pela ingesta 1 bloco 045 2 bloco 1 3 bloco 166 4 bloco 285 Após 60 dias parâmetros metabólicos e estruturais foram analisados Parâmetros estruturais Tamanho estrutural Parâmetros metabólicos Concentração de proteína glicose e triglicerídeos plasmáticos Atividade hepática e muscular ácido málico glicose 6fosfato e enzimas desidrogenase Resultados Nenhuma mortalidade de peixe foi encontrada durante o experimento Não houve diferença significativa relacionada ao tamanho estrutural na concentração de proteínas e triglicerídeos plasmáticos e musculares após o experimento Entretanto no bloco em que a quantidade de carboidrato foi de 285 demonstrou maiores concentrações de glicose plasmática e hepática Já no bloco de 045 a quantidade que demonstrou concentração elevada foi de triglicérides hepáticos ConclusãoCrítica ao artigo Não houve alteração metabólica significativa nos animais a partir do experimento Mudanças metabólicas encontradas em cada animal depende muito da biodisponibilidade de cada um em absorver e processar o que lhe foi ofertado além da condição de cada experimento em si O que sugere então que aumento de quantidade e concentração de carboidratos e lipídeos em uma dieta podem sugerir aumento da glicemia e concentrações de proteína entretanto não sugerem alterações estruturais REFERÊNCIA OKAMURA D FORTESSILVA R PAULINO RR ARAÚJO FG COSTA DV PEREIRA RT ROSA PV Respostas metabólicas às proporções de carboidratos e lipídios na dieta neotropical Anais da Academia Brasileira de Ciências 914 2019