Effects of stocking density on the growth and immunity of Atlantic salmon salmo salar reared in recirculating aquaculture system(RAS)
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摘要
Atlantic salmon reared in recirculating aquaculture system(RAS) may lead to inappropriately high stocking density, because ?sh live in a limited space. Finding the suitable stocking density of Atlantic salmon reared in RAS is very important for RAS industry. In this paper, the in?uence of stocking density on growth and some stress related physiological factors were investigated to evaluate the ef fects of stocking density. The ?sh were reared for 220 days at ?ve densities(A: 24 kg/m~3; B: 21 kg/m~3; C: 15 kg/m~3; D: 9 kg/m~3 and E: 6 kg/m~3). The results show that 30 kg/m~3 might be the maximum density which RAS can af ford in China. The stocking densities under 30 kg/m~3 have no ef fect on mortality of Atlantic salmon reared in RAS. However, the speci?c growth rate(SGR), ?nal weight and weight gain in the high density group were signi?cantly lower than the lower density groups and middle density groups. Moreover, feed conversion rate(FCR) had a negative correlation with density. Plasma hormone T_3 and GH showed signi?cant decrease with the increase of the stocking density of the experiment. Furthermore, thyroid hormone(T 3), GH(growth hormone) activities were decreased with stocking density increase. However, plasma cortisol, GOT(glutamic oxalacetic transaminase) and GPT(glutamic pyruvic transaminase) activities were increase with stocking density increase. And the stocking density has no ef fects on plasma lysozyme of Atlantic salmon reared in RAS. These investigations would also help devise effcient ways to rear adult Atlantic salmon in China and may, in a way, help spread salmon mariculture in China.
Atlantic salmon reared in recirculating aquaculture system(RAS) may lead to inappropriately high stocking density, because ?sh live in a limited space. Finding the suitable stocking density of Atlantic salmon reared in RAS is very important for RAS industry. In this paper, the influence of stocking density on growth and some stress related physiological factors were investigated to evaluate the ef fects of stocking density. The ?sh were reared for 220 days at ?ve densities(A: 24 kg/m~3; B: 21 kg/m~3; C: 15 kg/m~3; D: 9 kg/m~3 and E: 6 kg/m~3). The results show that 30 kg/m~3 might be the maximum density which RAS can af ford in China. The stocking densities under 30 kg/m~3 have no ef fect on mortality of Atlantic salmon reared in RAS. However, the speci?c growth rate(SGR), ?nal weight and weight gain in the high density group were signi?cantly lower than the lower density groups and middle density groups. Moreover, feed conversion rate(FCR) had a negative correlation with density. Plasma hormone T_3 and GH showed signi?cant decrease with the increase of the stocking density of the experiment. Furthermore, thyroid hormone(T 3), GH(growth hormone) activities were decreased with stocking density increase. However, plasma cortisol, GOT(glutamic oxalacetic transaminase) and GPT(glutamic pyruvic transaminase) activities were increase with stocking density increase. And the stocking density has no ef fects on plasma lysozyme of Atlantic salmon reared in RAS. These investigations would also help devise effcient ways to rear adult Atlantic salmon in China and may, in a way, help spread salmon mariculture in China.
Atlantic salmon reared in recirculating aquaculture system(RAS) may lead to inappropriately high stocking density, because ?sh live in a limited space. Finding the suitable stocking density of Atlantic salmon reared in RAS is very important for RAS industry. In this paper, the influence of stocking density on growth and some stress related physiological factors were investigated to evaluate the ef fects of stocking density. The ?sh were reared for 220 days at ?ve densities(A: 24 kg/m~3; B: 21 kg/m~3; C: 15 kg/m~3; D: 9 kg/m~3 and E: 6 kg/m~3). The results show that 30 kg/m~3 might be the maximum density which RAS can af ford in China. The stocking densities under 30 kg/m~3 have no ef fect on mortality of Atlantic salmon reared in RAS. However, the speci?c growth rate(SGR), ?nal weight and weight gain in the high density group were signi?cantly lower than the lower density groups and middle density groups. Moreover, feed conversion rate(FCR) had a negative correlation with density. Plasma hormone T_3 and GH showed signi?cant decrease with the increase of the stocking density of the experiment. Furthermore, thyroid hormone(T 3), GH(growth hormone) activities were decreased with stocking density increase. However, plasma cortisol, GOT(glutamic oxalacetic transaminase) and GPT(glutamic pyruvic transaminase) activities were increase with stocking density increase. And the stocking density has no ef fects on plasma lysozyme of Atlantic salmon reared in RAS. These investigations would also help devise effcient ways to rear adult Atlantic salmon in China and may, in a way, help spread salmon mariculture in China.
引文
Ahmad H I,Verma A K,Rani A M B,Rathore G,Saharan N,Gora A H.2016.Growth,non-speci?c immunity and disease resistance of Labeo rohita against Aeromonas hydrophila in bio?oc systems using dif ferent carbon sources.Aquaculture,457:61-67,https://doi.org/10.1016/j.aquaculture.2016.02.011.
Ashley P J.2007.Fish welfare:current issues in aquaculture.Applied Animal Behaviour Science,104(3-4):199-235,https://doi.org/10.1016/j.applanim.2006.09.001.
Bolasina S,Tagawa M,Yamashita Y,Tanaka M.2006.Ef fect of stocking density on growth,digestive enzyme activity and cortisol level in larvae and juveniles of Japanese?ounder,Paralichthys olivaceus.Aquaculture,259(1-4):432-443,https://doi.org/10.1016/j.aquaculture.2006.05.021.
Bonga S E W.1997.The stress response in?sh.Physiological Reviews,77(3):591-625.
Carter P.1978.A convenient method for the determination of nonspeci?c binding in commercially available solid phase iodine labeled radioimmunoassay kits.Clinical Biochemistry,11(3):97-100,https://doi.org/10.1016/S0009-9120(78)90070-X.
Chi L,Li X,Liu Q H,Liu Y.2017.Photoperiod regulate gonad development via kisspeptin/kissr in hypothalamus and saccus vasculosus of Atlantic salmon(Salmo salar).PLoSOne,12(2):e0169569,https://doi.org/10.1371/journal.pone.0169569.
Costas B,Arag?o C,Dias J,Afonso A,Conceicao L E C.2013.Interactive ef fects of a high-quality protein diet and high stocking density on the stress response and some innate immune parameters of Senegalese sole Solea senegalensis.Fish Physiology and Biochemistry,39:1 141-1 151,https://doi.org/10.1007/s10695-013-9770-1.
Ellis T,North B,Scott A P,Bromage N R,Porter M,Gadd D.2002.The relationships between stocking density and welfare in farmed rainbow trout.Journal of Fish Biology,61(3):493-531,https://doi.org/10.1006/jfbi.2002.2057.
Geng X,Dong X H,Tan B P,Yang Q H,Chi S Y,Liu H Y,Liu X Q.2012.Ef fects of dietary probiotic on the growth performance,non-speci?c immunity and disease resistance of cobia,Rachycentron canadum.Aquacult ure Nutr ition,18(1):46-55,https://doi.org/10.1111/j.1365-2095.2011.00875.x.
Greaves K,Tuene S.2001.The form and context of aggressive behaviour in farmed Atlantic halibut(Hippoglossus hippoglossus L.).Aquaculture,193(1-2):139-147,https://doi.org/10.1016/S0044-8486(00)00476-2.
Hori T S,Gamperl A K,Afonso L O,Johnson S C,Hubert S,Kimball J,Bowman S,Rise M L.2010.Heat-shock responsive genes identi?ed and validated in Atlantic cod(Gadus morhua)liver,head kidney and skeletal muscle using genomic techniques.BMC Genomics,11:72,https://doi.org/7210.1186/1471-2164-11-72.
Hosfeld C D,Hammer J,Handeland S O,Fivelstad S,Stefansson S O.2009.Ef fects of?sh density on growth and smolti?cation in intensive production of Atlantic salmon(Salmo salar L.).Aquaculture,294(3-4):236-241,https://doi.org/10.1016/j.aquaculture.2009.06.003.
Ingram G A.1980.Substances involved in the natural resistance of?sh to infection-a review.Journal of Fish Biology,16(1):23-60,https://doi.org/10.1111/j.1095-8649.1980.tb03685.x.
Laiz-Carrión R,Viana I R,Cejas J R,Ruiz-Jarabo I,Jerez S,Martos J A,Eduardo A B,Mancera J M.2012.In?uence of food deprivation and high stocking density on energetic metabolism and stress response in red porgy,Pagrus pagrus L.Aquaculture International,20(3):585-599,https://doi.org/10.1007/s10499-011-9488-y.
Laursen D C,Silva P I M,Larsen B K,H?glund E.2013.High oxygen consumption rates and scale loss indicate elevated aggressive behaviour at low rearing density,while elevated brain serotonergic activity suggests chronic stress at high rearing densities in farmed rainbow trout.Physiology&Behavior,122:147-154,https://doi.org/10.1016/j.physbeh.2013.08.026.
Leung L Y,Kwong A K Y,Man A K Y,Woo N Y S.2008.Direct actions of cortisol,thyroxine and growth hormone on IGF-I mRNA expression in sea bream hepatocytes.Comparative Biochemistry and Physiology Part A:Molecular&Integrative Physiology,151(4):705-710,https://doi.org/10.1016/j.cbpa.2008.08.023.
Li M J,Yin Y C,Hua H,Sun X M,Luo T,Wang J A,Jiang YF.2010.The reciprocal regulation ofγ-synuclein and IGF-I receptor expression creates a circuit that modulates IGF-I signaling.Journal of Biological Chemistry,285(40):30 480-30 488,https://doi.org/10.1074/jbc.M110.131698.
Li X,Liu Y,Blancheton J P.2013.Ef fect of stocking density on performances of juvenile turbot(Scophthalmus maximus)in recirculating aquaculture systems.Chinese Journal of Oceanology and Limnology,31(3):514-522,https://doi.org/10.1007/s00343-013-2205-0.
Liu B L,Liu Y,Wang X P.2015.The ef fect of stocking density on growth and seven physiological parameters with assessment of their potential as stress response indicators for the Atlantic salmon(Sa l mo salar).Mar Freshwat Behav Physiol.,48(3):177-192,https://doi.org/10.1080/10236244.2015.1034956.
López-Pati?o M A,Conde-Sieira M,Gesto M,Librán-Pérez M,Soengas J L,Míguez J M.2013.Melatonin partially minimizes the adverse stress ef fects in Senegalese sole(Solea senegalensis).Aquaculture,388-391:165-172,https://doi.org/10.1016/j.aquaculture.2013.01.023.
Martins C I M,Eding E H,Verdegem M C J,Heinsbroek L TN,Schneider O,Blancheton J P,d'Orbcastel E R,Verreth J A J.2010.New developments in recirculating aquaculture systems in Europe:a perspective on environmental sustainability.Aquacultural Engineering,43(3):83-93,https://doi.org/10.1016/j.aquaeng.2010.09.002.
Montero D,Izquierdo M S,Tort L,Robaina L,Vergara J M.1999.High stocking density produces crowding stress altering some physiological and biochemical parameters in gilthead seabream,Sparus aurata,juveniles.Fish Physiology and Biochemistry,20(1):53-60,https://doi.org/10.1023/A:1007719928905.
Nardocci G,Navarro C,Cortés P P,Imarai M,Montoya M,Valenzuela B,Jara P,Acu?a-Castillo C,Fernández R.2014.Neuroendocrine mechanisms for immune system regulation during stress in?sh.Fish&Shell?sh Immunol ogy,40(2):531-538,https://doi.org/10.1016/j.fsi.2014.08.001.
Prusty A K,Kohli M P S,Sahu N P,Pal A K,Saharan N,Mohapatra S,Gupta S K.2011.Ef fect of short term exposure of fenvalerate on biochemical and haematological responses in Labeo rohita(Hamilton)?ngerlings.Pestic ide Biochemistry and Physiology,100(2):124-129,https://doi.org/10.1016/j.pestbp.2011.02.010.
Salas-Leiton E,Anguis V,Martín-Antonio B,Crespo D,Planas J V,Infante C,Ca?avate J P,Manchado M.2010.Ef fects of stocking density and feed ration on growth and gene expression in the Senegalese sole(Solea senegalensis):potential ef fects on the immune response.Fish&Shell?sh Immunol ogy,28(2):296-302,https://doi.org/10.1016/j.fsi.2009.11.006.
Schmid A C,Lutz I,Kloas W,Reinecke M.2003.Thyroid hormone stimulates hepatic IGF-I mRNA expression in a bony?sh,the tilapia Oreochromis mossambicus,in vitro and in vivo.General and Comparative Endocrinology,130(2):129-134,https://doi.org/10.1016/S0016-6480(02)00577-4.
Schmid A C,Reinecke M,Kloas W.2000.Primary cultured hepatocytes of the bony?sh,Oreochromis mossambicus,the tilapia:a valid tool for physiological studies on IGF-Iexpression in liver.The Journal of Endocrinology,166(2):265-273,https://doi.org/10.1677/joe.0.1660265.
Schram E,Van der Heul J W,Kamstra A,Verdegem M C J.2006.Stocking density-dependent growth of Dover sole(Solea solea).Aquaculture,252(2-4):339-347,https://doi.org/10.1016/j.aquaculture.2005.07.011.
Tort L.2011.Stress and immune modulation in?sh.Developmental and Comparative Immunology.,35(12):1 366-1 375,https://doi.org/10.1016/j.dci.2011.07.002.
Yarahmadi P,Miandare H K,Fayaz S,Caipang C M A.2016.Increased stocking density causes changes in expression of selected stress-and immune-related genes,humoral innate immune parameters and stress responses of rainbow trout(Oncorhynchus mykiss).Fish&Shell?sh Immunology,48:43-53,https://doi.org/10.1016/j.fsi.2015.11.007.
Zhang X N,Tian H,Wang W,Ru S G.2014.Monocrotophos pesticide decreases the plasma levels of total 3,3',5-triiodol-thyronine and alters the expression of genes associated with the thyroidal axis in female gold?sh(Carassius auratus).PLoS One.,9(9):e108972,https://doi.org/10.1371/journal.pone.0108972.
Ashley P J.2007.Fish welfare:current issues in aquaculture.Applied Animal Behaviour Science,104(3-4):199-235,https://doi.org/10.1016/j.applanim.2006.09.001.
Bolasina S,Tagawa M,Yamashita Y,Tanaka M.2006.Ef fect of stocking density on growth,digestive enzyme activity and cortisol level in larvae and juveniles of Japanese?ounder,Paralichthys olivaceus.Aquaculture,259(1-4):432-443,https://doi.org/10.1016/j.aquaculture.2006.05.021.
Bonga S E W.1997.The stress response in?sh.Physiological Reviews,77(3):591-625.
Carter P.1978.A convenient method for the determination of nonspeci?c binding in commercially available solid phase iodine labeled radioimmunoassay kits.Clinical Biochemistry,11(3):97-100,https://doi.org/10.1016/S0009-9120(78)90070-X.
Chi L,Li X,Liu Q H,Liu Y.2017.Photoperiod regulate gonad development via kisspeptin/kissr in hypothalamus and saccus vasculosus of Atlantic salmon(Salmo salar).PLoSOne,12(2):e0169569,https://doi.org/10.1371/journal.pone.0169569.
Costas B,Arag?o C,Dias J,Afonso A,Conceicao L E C.2013.Interactive ef fects of a high-quality protein diet and high stocking density on the stress response and some innate immune parameters of Senegalese sole Solea senegalensis.Fish Physiology and Biochemistry,39:1 141-1 151,https://doi.org/10.1007/s10695-013-9770-1.
Ellis T,North B,Scott A P,Bromage N R,Porter M,Gadd D.2002.The relationships between stocking density and welfare in farmed rainbow trout.Journal of Fish Biology,61(3):493-531,https://doi.org/10.1006/jfbi.2002.2057.
Geng X,Dong X H,Tan B P,Yang Q H,Chi S Y,Liu H Y,Liu X Q.2012.Ef fects of dietary probiotic on the growth performance,non-speci?c immunity and disease resistance of cobia,Rachycentron canadum.Aquacult ure Nutr ition,18(1):46-55,https://doi.org/10.1111/j.1365-2095.2011.00875.x.
Greaves K,Tuene S.2001.The form and context of aggressive behaviour in farmed Atlantic halibut(Hippoglossus hippoglossus L.).Aquaculture,193(1-2):139-147,https://doi.org/10.1016/S0044-8486(00)00476-2.
Hori T S,Gamperl A K,Afonso L O,Johnson S C,Hubert S,Kimball J,Bowman S,Rise M L.2010.Heat-shock responsive genes identi?ed and validated in Atlantic cod(Gadus morhua)liver,head kidney and skeletal muscle using genomic techniques.BMC Genomics,11:72,https://doi.org/7210.1186/1471-2164-11-72.
Hosfeld C D,Hammer J,Handeland S O,Fivelstad S,Stefansson S O.2009.Ef fects of?sh density on growth and smolti?cation in intensive production of Atlantic salmon(Salmo salar L.).Aquaculture,294(3-4):236-241,https://doi.org/10.1016/j.aquaculture.2009.06.003.
Ingram G A.1980.Substances involved in the natural resistance of?sh to infection-a review.Journal of Fish Biology,16(1):23-60,https://doi.org/10.1111/j.1095-8649.1980.tb03685.x.
Laiz-Carrión R,Viana I R,Cejas J R,Ruiz-Jarabo I,Jerez S,Martos J A,Eduardo A B,Mancera J M.2012.In?uence of food deprivation and high stocking density on energetic metabolism and stress response in red porgy,Pagrus pagrus L.Aquaculture International,20(3):585-599,https://doi.org/10.1007/s10499-011-9488-y.
Laursen D C,Silva P I M,Larsen B K,H?glund E.2013.High oxygen consumption rates and scale loss indicate elevated aggressive behaviour at low rearing density,while elevated brain serotonergic activity suggests chronic stress at high rearing densities in farmed rainbow trout.Physiology&Behavior,122:147-154,https://doi.org/10.1016/j.physbeh.2013.08.026.
Leung L Y,Kwong A K Y,Man A K Y,Woo N Y S.2008.Direct actions of cortisol,thyroxine and growth hormone on IGF-I mRNA expression in sea bream hepatocytes.Comparative Biochemistry and Physiology Part A:Molecular&Integrative Physiology,151(4):705-710,https://doi.org/10.1016/j.cbpa.2008.08.023.
Li M J,Yin Y C,Hua H,Sun X M,Luo T,Wang J A,Jiang YF.2010.The reciprocal regulation ofγ-synuclein and IGF-I receptor expression creates a circuit that modulates IGF-I signaling.Journal of Biological Chemistry,285(40):30 480-30 488,https://doi.org/10.1074/jbc.M110.131698.
Li X,Liu Y,Blancheton J P.2013.Ef fect of stocking density on performances of juvenile turbot(Scophthalmus maximus)in recirculating aquaculture systems.Chinese Journal of Oceanology and Limnology,31(3):514-522,https://doi.org/10.1007/s00343-013-2205-0.
Liu B L,Liu Y,Wang X P.2015.The ef fect of stocking density on growth and seven physiological parameters with assessment of their potential as stress response indicators for the Atlantic salmon(Sa l mo salar).Mar Freshwat Behav Physiol.,48(3):177-192,https://doi.org/10.1080/10236244.2015.1034956.
López-Pati?o M A,Conde-Sieira M,Gesto M,Librán-Pérez M,Soengas J L,Míguez J M.2013.Melatonin partially minimizes the adverse stress ef fects in Senegalese sole(Solea senegalensis).Aquaculture,388-391:165-172,https://doi.org/10.1016/j.aquaculture.2013.01.023.
Martins C I M,Eding E H,Verdegem M C J,Heinsbroek L TN,Schneider O,Blancheton J P,d'Orbcastel E R,Verreth J A J.2010.New developments in recirculating aquaculture systems in Europe:a perspective on environmental sustainability.Aquacultural Engineering,43(3):83-93,https://doi.org/10.1016/j.aquaeng.2010.09.002.
Montero D,Izquierdo M S,Tort L,Robaina L,Vergara J M.1999.High stocking density produces crowding stress altering some physiological and biochemical parameters in gilthead seabream,Sparus aurata,juveniles.Fish Physiology and Biochemistry,20(1):53-60,https://doi.org/10.1023/A:1007719928905.
Nardocci G,Navarro C,Cortés P P,Imarai M,Montoya M,Valenzuela B,Jara P,Acu?a-Castillo C,Fernández R.2014.Neuroendocrine mechanisms for immune system regulation during stress in?sh.Fish&Shell?sh Immunol ogy,40(2):531-538,https://doi.org/10.1016/j.fsi.2014.08.001.
Prusty A K,Kohli M P S,Sahu N P,Pal A K,Saharan N,Mohapatra S,Gupta S K.2011.Ef fect of short term exposure of fenvalerate on biochemical and haematological responses in Labeo rohita(Hamilton)?ngerlings.Pestic ide Biochemistry and Physiology,100(2):124-129,https://doi.org/10.1016/j.pestbp.2011.02.010.
Salas-Leiton E,Anguis V,Martín-Antonio B,Crespo D,Planas J V,Infante C,Ca?avate J P,Manchado M.2010.Ef fects of stocking density and feed ration on growth and gene expression in the Senegalese sole(Solea senegalensis):potential ef fects on the immune response.Fish&Shell?sh Immunol ogy,28(2):296-302,https://doi.org/10.1016/j.fsi.2009.11.006.
Schmid A C,Lutz I,Kloas W,Reinecke M.2003.Thyroid hormone stimulates hepatic IGF-I mRNA expression in a bony?sh,the tilapia Oreochromis mossambicus,in vitro and in vivo.General and Comparative Endocrinology,130(2):129-134,https://doi.org/10.1016/S0016-6480(02)00577-4.
Schmid A C,Reinecke M,Kloas W.2000.Primary cultured hepatocytes of the bony?sh,Oreochromis mossambicus,the tilapia:a valid tool for physiological studies on IGF-Iexpression in liver.The Journal of Endocrinology,166(2):265-273,https://doi.org/10.1677/joe.0.1660265.
Schram E,Van der Heul J W,Kamstra A,Verdegem M C J.2006.Stocking density-dependent growth of Dover sole(Solea solea).Aquaculture,252(2-4):339-347,https://doi.org/10.1016/j.aquaculture.2005.07.011.
Tort L.2011.Stress and immune modulation in?sh.Developmental and Comparative Immunology.,35(12):1 366-1 375,https://doi.org/10.1016/j.dci.2011.07.002.
Yarahmadi P,Miandare H K,Fayaz S,Caipang C M A.2016.Increased stocking density causes changes in expression of selected stress-and immune-related genes,humoral innate immune parameters and stress responses of rainbow trout(Oncorhynchus mykiss).Fish&Shell?sh Immunology,48:43-53,https://doi.org/10.1016/j.fsi.2015.11.007.
Zhang X N,Tian H,Wang W,Ru S G.2014.Monocrotophos pesticide decreases the plasma levels of total 3,3',5-triiodol-thyronine and alters the expression of genes associated with the thyroidal axis in female gold?sh(Carassius auratus).PLoS One.,9(9):e108972,https://doi.org/10.1371/journal.pone.0108972.