降海鲑渗透生理及入海窗口期的研究进展
|
摘要
在过去30年中,降海鲑的发育得到了大量关注,研究降海鲑的发育,特别是如何调控降海鲑发育的时间和品质,以便幼鲑移入海水网箱。洄游型鲑科鱼类幼鲑向降海鲑的转变包含了一系列的行为、形态、生理的改变,这都为降海洄游进入河口做准备。降海鲑发育受几种激素的促进,包括生长激素、胰岛素样生长因子-I、皮质醇、甲状腺素,而催乳素通常会抑制发育。鳃氯细胞中特殊的离子转运蛋白(NKA、Na+/K+/Cl-协同转运蛋白和Cl-通道)丰度的增加引起盐分泌能力的提高,从而有利于降海鲑在海水中生长、游泳和存活。具银化期的鲑科鱼类(如大西洋鲑(Salmo salar)、硬头鳟(Oncorhyunchus mykiss)、银大麻哈鱼(Oncorhynchus kisutch)至少在淡水中生活一年,达到临界最小规格之后才可改变其对光周期的响应。如果在银化前未达到这个最小规格,则需等到下一年(或更长)。鲑科鱼类如何在一年中合适的时间确定其规格或生长率,并增加对光周期的响应性,这一重要的领域还未得到大量的关注。因此,了解降海鲑渗透生理及入海窗口期,查明不同鱼类适宜生长盐度,入海时间和方式非常重要,可为鲑科鱼类(如虹鳟(Oncorhynchus mykiss)、硬头鳟等)在黄海冷水团规模化养殖提供理论支持。
In the past 30 years,the researches on the smolt development have received a lot of attentions.Because of the relationship with the aquaculture development,the original purpose of these researches is to explore the smolt development,especially to control the time and quality of the smolt development into sea cage.The parr-smolt transformation of anadromous salmonids include a suite of behavioral,morphological and physiological changes that are preparatory for downstream migration and seawater entry.Smolt development is stimulated by several hormones including growth hormone,insulin-like growth factor-1,cortisol and thyroid hormones but inhibited generally by prolactin.Increase in the abundances of specific ion transporters(Na+/K+-ATPase,Na+/K+/Cl-cotransporter and apical Cl-channel)in gill ionocytes results in the increase of salt secretory capacity,growth and swimming performance in seawater and marine survival.In smolting salmonids that spend at least a year in FW(Atlantic salmon,steelhead trout,coho salmon),a minimum size must be achieved,after which the animals alter their response to photoperiod.If fish have not achieved this minimum size they will wait another year(or more)before transforming to smolts.How fish determine their size or growth rate at the appropriate time of year and translate this to increased responsiveness to daylength is an important area that has not received substantial attention.As a result,researching smolt osmophysiology and their sea entry time windows and ascertaining more optimal salinity,the way and time into sea of specific size salmonids are very important for providing theoretical supports to breeding rainbow and steelhead trout at Cold Water Mass of Yellow Sea.
In the past 30 years,the researches on the smolt development have received a lot of attentions.Because of the relationship with the aquaculture development,the original purpose of these researches is to explore the smolt development,especially to control the time and quality of the smolt development into sea cage.The parr-smolt transformation of anadromous salmonids include a suite of behavioral,morphological and physiological changes that are preparatory for downstream migration and seawater entry.Smolt development is stimulated by several hormones including growth hormone,insulin-like growth factor-1,cortisol and thyroid hormones but inhibited generally by prolactin.Increase in the abundances of specific ion transporters(Na+/K+-ATPase,Na+/K+/Cl-cotransporter and apical Cl-channel)in gill ionocytes results in the increase of salt secretory capacity,growth and swimming performance in seawater and marine survival.In smolting salmonids that spend at least a year in FW(Atlantic salmon,steelhead trout,coho salmon),a minimum size must be achieved,after which the animals alter their response to photoperiod.If fish have not achieved this minimum size they will wait another year(or more)before transforming to smolts.How fish determine their size or growth rate at the appropriate time of year and translate this to increased responsiveness to daylength is an important area that has not received substantial attention.As a result,researching smolt osmophysiology and their sea entry time windows and ascertaining more optimal salinity,the way and time into sea of specific size salmonids are very important for providing theoretical supports to breeding rainbow and steelhead trout at Cold Water Mass of Yellow Sea.
引文
[1]孙大江,王炳谦.鲑科鱼类及其养殖状况[J].水产学杂志,2010,23(2):56-63.Sun D J,Wang B Q.Salmonidaeand their breeding conditions[J].Chinese Journal of Fisheries,2010,23(2):56-63.
[2]周井娟,陈林兴.世界三文鱼的贸易格局及发展预测[J].农业经济与管理,2012(2):86-92.Zhou J J,Chen L X.The world trade patterns of salmon and the development of prediction[J].Agricltural Economics and Management,2012(2):86-92.
[3]李思忠.中国鲑科鱼类地理分布的探讨[J].动物学杂志,1984(1):36-39.Li S Z.The discussion of salmonidae distributionin China[J].Chinese Journal of Zoology,1984(1):36-39.
[4] Johnston C,Eales J.Purines in the integument of the Atlantic salmon(Salmo salar)during parr-smolt transformation[J].Journal of the Fisheries Board of Canada,1967,24(5):955-964.
[5]Mizuno S,Misaka N,Ando D,et al.Quantitative changes of black pigmentation in the dorsal fin margin during smoltification in masu salmon,Oncorhynchus masou[J].Aquaculture,2004,229(1-4):433-450.
[6] McCormick S D,Saunders R L.Preparatory physiological adaptations for marine life of salmonids:Osmoregulation,growth,and metabolism[J].American Fisheries Society Symposrum,1987,1(21):1-229.
[7] Beeman J,Rondorf D,Tilson M.Assessing smoltification of juvenile spring chinook salmon(Oncorhynchus tshawytscha)using changes in body morphology[J].Canadian Journal of Fisheries and Aquatic Sciences,1994,51(4):836-844.
[8]夏重志,陈金平,牟振波.大西洋鲑的生态学资料[J].水产学杂志,1998(2):61-65.Xia Z Z,Chen J P,Mu Z B.Theecology information of Atlantic salmon[J].Chinese Journal of Fisheries,1998(2):61-65.
[9] Edwards S L,Marshall W S.Principles and patterns of osmoregulation and euryhalinity in fishes[J].Fish Physiology,2013,32:1-44.
[10]McCormick S D,Lerner D T,Monette M Y,et al.Taking it with you when you go:How perturbations to the freshwater environment,including temperature,dams,and contaminants,affect marine survival of salmon[J].American Fisheries Society Symposium,2009,69:195-214.
[11] Evans D H,Piermarini P M,Choe K P.The multifunctional fish gill:Dominant site of gas exchange,osmoregulation,acid-base regulation,and excretion of nitrogenous waste[J].Physiological Reviews,2005,85(1):97-177.
[12] Zaugg W S,McLain L R.Adenosinetriphosphatase activity in gills of salmonids:seasonal variations and salt water influence in coho salmon,Oncorhynchus kisutch[J].Comparative Biochemistry and Physiology,1970,35(3):587-596.
[13] Richards J G.Na+/K+-ATPase-isoform switching in gills of rainbow trout(Oncorhynchus mykiss)during salinity transfer[J].Journal of Experimental Biology,2003,206(24):4475-4486.
[14] McCormick S D,Regish A M,Christensen A K.Distinct freshwater and seawater isoforms of Na+/K+-ATPase in gill chloride cells of Atlantic salmon[J].The Journal of Experimental Biology,2009,212(24):3994-4001.
[15] Nilsen T O,Ebbesson L O,Madsen S S,et al.Differential expression of gill Na+,K+-ATPase alpha-and beta-subunits,Na+,K+,2Cl-cotransporter and CFTR anion channel in juvenile anadromous and landlocked Atlantic salmon Salmo salar[J].Journal of Experimental Biology,2007,210(16):2885-2896.
[16] McCormick S D,Sheehan T F,Bj9rnsson B T,et al.Physiological and endocrine changes in Atlantic salmon smolts during hatchery rearing,downstream migration,and ocean entry[J].Canadian Journal of Fisheries and Aquatic Sciences,2013,70(1):105-118.
[17] Tipsmark C K,Mahmmoud Y A,Borski R J,et al.FXYD-11associates with Na+-K+-ATPase in the gill of Atlantic salmon:Regulation and localization in relation to changed ion-regulatory status[J].American Journal of Physiology Regulatory Integrative and Comparative Physiology,2010,299(5):1212-1223.
[18] Pelis R M,Zydlewski J,McCormick S D.Gill Na+-K+-2Cl-cotransporter abundance and location in Atlantic salmon:Effects of seawater and smolting[J].American Journal of Physiology-Regulatory,Integrative and Comparative Physiology,2001,280(6):1844-1852.
[19] Stefansson S O,Nilsen T O,Ebbesson L O E,et al.Molecular mechanisms of continuous light inhibition of Atlantic salmon parrsmolt transformation[J].Aquaculture,2007,273(2-3):235-245.
[20] Singer T D,Clements K M,Semple J W,et al.Seawater tolerance and gene expression in two strains of Atlantic salmon smolts[J].Canadian Journal of Fisheries and Aquatic Sciences,2002,59(1):125-135.
[21] Bern H.Endocrinological studies on normal and abnormal salmon smoltification[J].Comparative Endocrinology,1978,5(6):97-100.
[22] Bjornsson B T,Stefansson S O,McCormick S D.Environmental endocrinology of salmon smoltification[J].General and Comparative Endocrinology,2011,170(2):290-298.
[23]gústsson T,Sundell K,Sakamoto T,et al.Growth hormone endocrinology of Atlantic salmon(Salmo salar):Pituitary gene expression,hormone storage,secretion and plasma levels during parr-smolt transformation[J].Journal of Endocrinology,2001,170(1):227-234.
[24] Parhar I S,Iwata M.Intracerebral expression of gonadotropin-releasing hormone and growth hormone-releasing hormone is delayed until smoltification in the salmon[J].Neuroscience Research,1996,26(3):299-308.
[25] McCormick S D,Moriyama S,Bj9rnsson B T.Low temperature limits photoperiod control of smolting in Atlantic salmon through endocrine mechanisms[J].American Journal of Physiology-Regulatory,Integrative and Comparative Physiology,2000,278(5):1352-1361.
[26] McCormick S D,Shrimpton J M,Moriyama S,et al.Differential hormonal responses of Atlantic salmon parr and smolt to increased daylength:A possible developmental basis for smolting[J].Aquaculture,2007,273(2-3):337-344.
[27] Nilsen T O,Ebbesson L O,Kiilerich P,et al.Endocrine systems in juvenile anadromous and landlocked Atlantic salmon(Salmo salar):Seasonal development and seawater acclimation[J].General and Comparative Endocrinology,2008,155(3):762-72.
[28] Beckman B R,Dickhoff W W.Plasticity of smolting in spring chinook salmon:relation to growth and insulin-like growth factor-I[J].Journal of Fish Biology,1998,53(4):808-826.
[29] Beckman B R,Dickhoff W W,Zaugg W S,et al.Growth,smoltification,and smolt-to-adult return of spring chinook salmon from hatcheries on the Deschutes River,Oregon[J].Transactions of the American Fisheries Society,1999,128(6):1125-1150.
[30] Sakamoto T,Hirano T,Madsen S S,et al.Insulin-like growth factor I gene expression during parr-smolt transformation of coho salmon[J].Zoological Science,1995,12(2):249-252.
[31] Kiilerich P,Kristiansen K,Madsen S S.Hormone receptors in gills of smolting Atlantic salmon,Salmo salar:Expression of growth hormone,prolactin,mineralocorticoid and glucocorticoid receptors and 11beta-hydroxysteroid dehydrogenase type 2[J].General and Comparative Endocrinology,2007,152(2-3):295-303.
[32] Takei Y,McCormick S.Hormonal control of fish euryhalinity[J].Euryhaline Fishes Fish Physiology,2013,32(8):69-123.
[33] Tipsmark C K,Madsen S S.Distinct hormonal regulation of Na+,K+-atpase genes in the gill of Atlantic salmon(Salmo salar L.)[J].Journal of Endocrinology,2009,203(2):301-310.
[34] McCormick S D,Regish A M,Christensen A K,et al.Differential regulation of sodium-potassium pump isoforms during smolt development and seawater exposure of Atlantic salmon[J].The Journal of Experimental Biology,2013,216(7):1142-1151.
[35] Wood A W,Duan C,Bern H A.Insulin-like growth factor signaling in fish[J].International Review of Cytology,2005,243(6):215-285.
[36] Sheridan M A.Effects of thyroxin,cortisol,growth hormone,and prolactin on lipid metabolism of coho salmon,Oncorhynchus kisutch,during smoltification[J].General and Comparative Endocrinology,1986,64(2):220-238.
[37] Duan C,Plisetskaya E M,Dickhoff W W.Expression of insulinlike growth factor I in normally and abnormally developing coho salmon(Oncorhynchus kisutch)[J].Endocrinology,1995,136(2):446-452.
[38] Stefansson S,Bj9rnsson B,Sundell K,et al.Physiological characteristics of wild Atlantic salmon post-smolts during estuarine and coastal migration[J].Journal of Fish Biology,2003,63(4):942-955.
[39] Specker J L.Interrenal function and smoltification[J].Aquaculture,1982,28(1):59-66.
[40] Fontaine M,Hatey J.Sur la teneur en 17-hydroxycorticosteroides du plasm desaumon(Salmo salar L.)[J].Comptes Rendus de Academie des Sciences de,1954,239(3):319-321.
[41] Specker J L,Schreck C B.Changes in plasma corticosteroids during smoltification of coho salmon,Oncorhynchus kisutch[J].General and Comparative Endocrinology,1982,46(1):53-58.
[42] Shrimpton J M,Devlin R H,McLean E,et al.Increases in gill cytosolic corticosteroid receptor abundance and saltwater tolerance in juvenile coho salmon(Oncorhynchus kisutch)treated with growth hormone and placental lactogen[J].General and Comparative Endocrinology,1995,98(1):1-15.
[43] Nishioka R S,Bern H A,Lai K V,et al.Changes in the endocrine organs of coho salmon during normal and abnormal smoltification—an electron-microscope study[J].Aquaculture,1982,28(1):21-38.
[44] Prunet P,Boeuf G,Bolton J P,et al.Smoltification and seawater adaptation in Atlantic salmon(Salmo salar):Plasma prolactin,growth hormone,and thyroid hormones[J].General and Comparative Endocrinology,1989,74(3):355-364.
[45] Young G,Bj9rnsson B T,Prunet P,et al.Smoltification and seawater adaptation in coho salmon(Oncorhynchus kisutch):Plasma prolactin,growth hormone,thyroid hormones,and cortisol[J].General and Comparative Endocrinology,1989,74(3):335-345.
[46] Yada T,Takahashi K,Hirano T.Seasonal changes in seawater adaptability and plasma levels of prolactin and growth hormone in landlocked sockeye salmon(Oncorhynchus nerka)and amago salmon(O.rhodurus)[J].General and Comparative Endocrinology,1991,82(1):33-44.
[47] Hgsen H R,Prunet P.Plasma levels of thyroxine,prolactin,and cortisol in migrating and resident wild arctic char,Salvelinus alpinus[J].Canadian Journal of Fisheries and Aquatic Sciences,1997,54(12):2947-2954.
[48] Seidelin M,Madsen S S.Prolactin antagonizes the seawater-adaptive effect of cortisol and growth hormone in anadromous brown trout(Salmo trutta)[J].Zoological Science,1997,14(2):249-256.
[49] Seidelin M,Madsen S.Endocrine control of Na+,K+-ATPase and chloride cell development in brown trout(Salmo trutta):Interaction of insulin-like growth factor-I with prolactin and growth hormone[J].Journal of Endocrinology,1999,162(1):127-135.
[50] Hoar W S.The thyroid gland of the Atlantic salmon[J].Journal of Morphology,1939,65(2):257-295.
[51] Dickhoff W W,Folmar L C,Gorbman A.Changes in plasma thyroxine during smoltification of coho salmon, Oncorhynchus kisutch[J].General and Comparative Endocrinology,1978,36(2):229-232.
[52] Dickhoff W W,Sullivan C V.Involvement of the thyroid gland in smoltification,with special reference to metabolic and developmental processes[J].American Fisheries Society Symposium,1987,1(6):197-210.
[53] Iwata M,Tsuboi H,Yamashita T,et al.Function and trigger of thyroxine surge in migrating chum salmon Oncorhynchus ketafry[J].Aquaculture,2003,222(1-4):315-329.
[54] McCormick S D,O'Dea M F,Moeckel A M,et al.Endocrine and physiological changes in Atlantic salmon smolts following hatchery release[J].Aquaculture,2003,222(1-4):45-57.
[55] Hutchison M J,Iwata M.Effect of thyroxine on the decrease of aggressive behaviour of four salmonids during the parr-smolt transformation[J].Aquaculture,1998,168(1):169-175.
[56] Iwata M.Downstream migratory behavior of salmonids and its relationship with cortisol and thyroid hormones:A review[J].Aquaculture,1995,135(1):131-139.
[57] Ojima D,Iwata M.The relationship between thyroxine surge and onset of downstream migration in chum salmon Oncorhynchus ketafry[J].Aquaculture,2007,273(2-3):185-193.
[58] Ojima D,Iwata M.Central administration of growth hormone-releasing hormone and corticotropin-releasing hormone stimulate downstream movement and thyroxine secretion in fall-smolting coho salmon(Oncorhynchus kisutch)[J].General and Comparative Endocrinology,2010,168(1):82-87.
[59] McCormick S D.Endocrine control of osmoregulation in teleost fish[J].American Zoologist,2001,41(4):781-794.
[60] Shrimpton J M,McCormick S D.Regulation of gill cytosolic corticosteroid receptors in juvenile Atlantic salmon:Interaction effects of growth hormone with prolactin and triiodothyronine[J].General and Comparative Endocrinology,1998,112(2):262-274.
[61] Taylor E B.Environmental correlates of life-history variation in juvenile chinook salmon,Oncorhynchus tshawytscha(Walbaum)[J].Journal of Fish Biology,1990,37(1):1-17.
[62] McCormick S D.Ontogeny and evolution of salinity tolerance in anadromous salmonids:Hormones and heterochrony[J].Estuaries,1994,17(1):26-33.
[63] Gallagher Z S,Bystriansky J S,Farrell A P,et al.A novel pattern of smoltification in the most anadromous salmonid:pink salmon(Oncorhynchus gorbuscha)[J].Canadian Journal of Fisheries and Aquatic Sciences,2013,70(3):349-357.
[64] Zaugg W S,Mclain L R.Influence of water temperature on gill sodium,potassium-stimulated ATPase activity in juvenile coho salmon(Oncorhynchus kisutch)[J].Comparative Biochemistry and Physiology Part A:Physiology,1976,54(4):419-421.
[65] Duston J,Saunders R,Knox D.Effects of increases in freshwater temperature on loss of smolt characteristics in Atlantic salmon(Salmo salar)[J].Canadian Journal of Fisheries and Aquatic Sciences,1991,48(2):164-169.
[66] McCormick S D,Shrimpton J M,Zydlewski J D.Temperature effects on osmoregulatory physiology of juvenile anadromous fish[J].Researche and Exploration in Laboratory,1996,46(3):170-188.
[67] Wagner H H.Photoperiod and temperature regulation of smolting in steelhead trout(Salmo gairdneri)[J].Canadian Journal of Zoology,1974,52(2):219-234.
[68] Zydlewski G B,Haro A,McCormick S D.Evidence for cumulative temperature as an initiating and terminating factor in downstream migratory behavior of Atlantic salmon(Salmo salar)smolts[J].Canadian Journal of Fisheries and Aquatic Sciences,2005,62(1):68-78.
[69] Staurnes M,Lysfjord G,Hansen L,et al.Recapture rates of hatchery-reared Atlantic salmon(Salmo salar)related to smolt development and time of release[J].Aquaculture,1993,118(3-4):327-337.
[70] Lundqvist H,McKinnell S,Fngstam H,et al.The effect of time,size and sex on recapture rates and yield after river releases of Salmo salar smolts[J].Aquaculture,1994,121(1-3):245-257.
[71] Bilton H,Alderdice D,Schnute J.Influence of time and size at release of juvenile coho salmon(Oncorhynchus kisutch)on returns at maturity[J].Canadian Journal of Fisheries and Aquatic Sciences,1982,39(3):426-447.
[72] Virtanen E,S9derholm-Tana L,Soivio A,et al.Effect of physiological condition and smoltification status at smolt release on subsequent catches of adult salmon[J].Aquaculture,1991,97(2):231-257.
[73] McCormick S D,Hansen L P,Quinn T P,et al.Movement,migration,and smolting of Atlantic salmon(Salmo salar)[J].Canadian Journal of Fisheries and Aquatic Sciences,1998,55(S1):77-92.
[74] Orciari R D,Leonard G H.Length characteristics of smolts and Timing of downstream migration among three strains of Atlantic salmon in a Southern new england stream[J].North American Journal of Fisheries Management,1996,16(4):851-860.
[75] McGinnity P,de Eyto E,Cross T F,et al.Population specific smolt development,migration and maturity schedules in Atlantic salmon in a natural river environment[J].Aquaculture,2007,273(2-3):257-268.
[76] Handeland S O,Wilkinson E,SveinsbB,et al.Temperature influence on the development and loss of seawater tolerance in two fast-growing strains of Atlantic salmon[J].Aquaculture,2004,233(1-4):513-529.
[77] Banks J L.Raceway density and water flow as factors affecting spring chinook salmon(Oncorhynchus tshawytscha)during rearing and after release[J].Aquaculture,1994,119(2):201-217.
[78] Miyakoshi Y,Nagata M,Kitada S.Effect of smolt size on postrelease survival of hatchery-reared masu salmon Oncorhynchus masou[J].Fisheries Science,2001,67(1):134-137.
[79] Connor W P,Smith S G,Andersen T,et al.Postrelease performance of hatchery yearling and subyearling fall Chinook salmon released into the Snake River[J].North American Journal of Fisheries Management,2004,24(2):545-560.
[80] Saloniemi I.Survival of reared and wild Atlantic salmon smolts:Size matters more in bad years[J].ICES Journal of Marine Science,2004,61(5):782-787.
[81] Dieperink C.Predation on Atlantic salmon and sea trout during their first days as postsmolts[J].Journal of Fish Biology,2002,61(3):848-852.
[82] Jrvi T.Cumulative acute physiological stress in Atlantic salmon smolts:The effect of osmotic imbalance and the presence of predators[J].Aquaculture,1990,89(3):337-350.
[83] Handeland S,Jrvi T,Fern9A,et al.Osmotic stress,antipredatory behaviour,and mortality of Atlantic salmon(Salmo salar)smolts[J].Canadian Journal of Fisheries and Aquatic Sciences,1996,53(12):2673-2680.
[84] Price C S,Schreck C B.Stress and saltwater-entry behavior of juvenile chinook salmon(Oncorhynchus tshawytscha):Conflicts in physiological motivation[J].Canadian Journal of Fisheries and Aquatic Sciences,2003,60(8):910-918.
[85] Schreck C B,Stahl T P,Davis L E,et al.Mortality estimates of juvenile spring-summer chinook salmon in the lower columbia river and estuary,1992-1998:Evidence for Delayed Mortality?[J].Transactions of the American Fisheries Society,2006,135(2):457-475.
[86] Loge F J,Arkoosh M R,Ginn T R,et al.Impact of environmental stressors on the dynamics of disease transmission[J].Environmental Science&Technology,2005,39(18):7329-7336.
[87] Schreck C B,Solazzi M F,Johnson S L,et al.Transportation stress affects performance of coho salmon,Oncorhynchus kisutch[J].Aquaculture,1989,82(1):15-20.
[88] Staurnes M,Sigholt T,Asgard T,et al.Effects of a temperature shift on seawater challenge test performance in Atlantic salmon(Salmo salar)smolt[J].Aquaculture,2001,201(1-2):153-159.
[89] Duston J.Effect of salinity on survival and growth of Atlantic salmon(Salmo salar)parr and smolts[J].Aquaculture,1994,121(1-3):115-124.
[90] Duncan N J,Auchinachie N,Robertson D,et al.Growth,maturation and survival of out-of-season 0+and 1+Atlantic salmon(Salmo salar)smolts[J].Aquaculture,1998,168(1):325-339.
[91] Johnsson J,Clarke W.Development of seawater adaptation in juvenile steelhead trout(Salmo gairdneri)and domesticated rainbow trout(Salmo gairdneri)-effects of size,temperature and photoperiod[J].Aquaculture,1988,71(3):247-263.
[92] Sedgwick S D.Rainbow trout farming in Scotland,Farming trout in salt water[J].Scott Agric,1970,49:180-185.
[93] Landless P J.Acclimation of rainbow trout to sea water[J].Aquaculture,1976,7(2):173-179.
[94]董双林.论我国水产养殖业生态集约化发展[J].中国渔业经济,2015,33(5):4-9.Dong S L.On ecological intensification of aquaculture systems in China[J].Chinese Fisheries Economics,2015,33(5):4-9.
[95]韩立民,郭永超,董双林.开发黄海冷水团建立国家离岸养殖试验区的研究[J].太平洋学报,2016,24(5):79-85.Han L M,Guo Y C,Dong S L.Research on establishing a national offshore aquaculture experimental zone based on the development of the Yellow Sea Cold Water Mass[J].Pacific Journal,2016.24(5):79-85.
[96] Iwata M,Ogura H,Komatsu S,et al.Changes in salinity preference of chum and coho salmon during development[J].Aquaculture,1985,45(1-4):380-381.
[97] Clarke W C,Nagahama Y.Effect of premature transfer to sea water on growth and morphology of[J].Canadian Journal of Zoology,1977,55(10):1620-1630.
[98] Folmar L C,Dickhoff W W,Mahnken C V W,et al.Stunting and parr-reversion during smoltification of coho salmon(Oncorhynchus kisutch)[J].Aquaculture,1982,28(1-2):91-104.
[99] Boeuf G,Harache Y.Criteria for adaptation of salmonids to high salinity seawater in France[J].Aquaculture,1982,28(1):163-176.
[100] Koch H J A.Hemoglobin changes with size in the Atlantic salmon(Salmo salar L.)[J].Aquaculture,1982,28(1-2):231-240.
[2]周井娟,陈林兴.世界三文鱼的贸易格局及发展预测[J].农业经济与管理,2012(2):86-92.Zhou J J,Chen L X.The world trade patterns of salmon and the development of prediction[J].Agricltural Economics and Management,2012(2):86-92.
[3]李思忠.中国鲑科鱼类地理分布的探讨[J].动物学杂志,1984(1):36-39.Li S Z.The discussion of salmonidae distributionin China[J].Chinese Journal of Zoology,1984(1):36-39.
[4] Johnston C,Eales J.Purines in the integument of the Atlantic salmon(Salmo salar)during parr-smolt transformation[J].Journal of the Fisheries Board of Canada,1967,24(5):955-964.
[5]Mizuno S,Misaka N,Ando D,et al.Quantitative changes of black pigmentation in the dorsal fin margin during smoltification in masu salmon,Oncorhynchus masou[J].Aquaculture,2004,229(1-4):433-450.
[6] McCormick S D,Saunders R L.Preparatory physiological adaptations for marine life of salmonids:Osmoregulation,growth,and metabolism[J].American Fisheries Society Symposrum,1987,1(21):1-229.
[7] Beeman J,Rondorf D,Tilson M.Assessing smoltification of juvenile spring chinook salmon(Oncorhynchus tshawytscha)using changes in body morphology[J].Canadian Journal of Fisheries and Aquatic Sciences,1994,51(4):836-844.
[8]夏重志,陈金平,牟振波.大西洋鲑的生态学资料[J].水产学杂志,1998(2):61-65.Xia Z Z,Chen J P,Mu Z B.Theecology information of Atlantic salmon[J].Chinese Journal of Fisheries,1998(2):61-65.
[9] Edwards S L,Marshall W S.Principles and patterns of osmoregulation and euryhalinity in fishes[J].Fish Physiology,2013,32:1-44.
[10]McCormick S D,Lerner D T,Monette M Y,et al.Taking it with you when you go:How perturbations to the freshwater environment,including temperature,dams,and contaminants,affect marine survival of salmon[J].American Fisheries Society Symposium,2009,69:195-214.
[11] Evans D H,Piermarini P M,Choe K P.The multifunctional fish gill:Dominant site of gas exchange,osmoregulation,acid-base regulation,and excretion of nitrogenous waste[J].Physiological Reviews,2005,85(1):97-177.
[12] Zaugg W S,McLain L R.Adenosinetriphosphatase activity in gills of salmonids:seasonal variations and salt water influence in coho salmon,Oncorhynchus kisutch[J].Comparative Biochemistry and Physiology,1970,35(3):587-596.
[13] Richards J G.Na+/K+-ATPase-isoform switching in gills of rainbow trout(Oncorhynchus mykiss)during salinity transfer[J].Journal of Experimental Biology,2003,206(24):4475-4486.
[14] McCormick S D,Regish A M,Christensen A K.Distinct freshwater and seawater isoforms of Na+/K+-ATPase in gill chloride cells of Atlantic salmon[J].The Journal of Experimental Biology,2009,212(24):3994-4001.
[15] Nilsen T O,Ebbesson L O,Madsen S S,et al.Differential expression of gill Na+,K+-ATPase alpha-and beta-subunits,Na+,K+,2Cl-cotransporter and CFTR anion channel in juvenile anadromous and landlocked Atlantic salmon Salmo salar[J].Journal of Experimental Biology,2007,210(16):2885-2896.
[16] McCormick S D,Sheehan T F,Bj9rnsson B T,et al.Physiological and endocrine changes in Atlantic salmon smolts during hatchery rearing,downstream migration,and ocean entry[J].Canadian Journal of Fisheries and Aquatic Sciences,2013,70(1):105-118.
[17] Tipsmark C K,Mahmmoud Y A,Borski R J,et al.FXYD-11associates with Na+-K+-ATPase in the gill of Atlantic salmon:Regulation and localization in relation to changed ion-regulatory status[J].American Journal of Physiology Regulatory Integrative and Comparative Physiology,2010,299(5):1212-1223.
[18] Pelis R M,Zydlewski J,McCormick S D.Gill Na+-K+-2Cl-cotransporter abundance and location in Atlantic salmon:Effects of seawater and smolting[J].American Journal of Physiology-Regulatory,Integrative and Comparative Physiology,2001,280(6):1844-1852.
[19] Stefansson S O,Nilsen T O,Ebbesson L O E,et al.Molecular mechanisms of continuous light inhibition of Atlantic salmon parrsmolt transformation[J].Aquaculture,2007,273(2-3):235-245.
[20] Singer T D,Clements K M,Semple J W,et al.Seawater tolerance and gene expression in two strains of Atlantic salmon smolts[J].Canadian Journal of Fisheries and Aquatic Sciences,2002,59(1):125-135.
[21] Bern H.Endocrinological studies on normal and abnormal salmon smoltification[J].Comparative Endocrinology,1978,5(6):97-100.
[22] Bjornsson B T,Stefansson S O,McCormick S D.Environmental endocrinology of salmon smoltification[J].General and Comparative Endocrinology,2011,170(2):290-298.
[23]gústsson T,Sundell K,Sakamoto T,et al.Growth hormone endocrinology of Atlantic salmon(Salmo salar):Pituitary gene expression,hormone storage,secretion and plasma levels during parr-smolt transformation[J].Journal of Endocrinology,2001,170(1):227-234.
[24] Parhar I S,Iwata M.Intracerebral expression of gonadotropin-releasing hormone and growth hormone-releasing hormone is delayed until smoltification in the salmon[J].Neuroscience Research,1996,26(3):299-308.
[25] McCormick S D,Moriyama S,Bj9rnsson B T.Low temperature limits photoperiod control of smolting in Atlantic salmon through endocrine mechanisms[J].American Journal of Physiology-Regulatory,Integrative and Comparative Physiology,2000,278(5):1352-1361.
[26] McCormick S D,Shrimpton J M,Moriyama S,et al.Differential hormonal responses of Atlantic salmon parr and smolt to increased daylength:A possible developmental basis for smolting[J].Aquaculture,2007,273(2-3):337-344.
[27] Nilsen T O,Ebbesson L O,Kiilerich P,et al.Endocrine systems in juvenile anadromous and landlocked Atlantic salmon(Salmo salar):Seasonal development and seawater acclimation[J].General and Comparative Endocrinology,2008,155(3):762-72.
[28] Beckman B R,Dickhoff W W.Plasticity of smolting in spring chinook salmon:relation to growth and insulin-like growth factor-I[J].Journal of Fish Biology,1998,53(4):808-826.
[29] Beckman B R,Dickhoff W W,Zaugg W S,et al.Growth,smoltification,and smolt-to-adult return of spring chinook salmon from hatcheries on the Deschutes River,Oregon[J].Transactions of the American Fisheries Society,1999,128(6):1125-1150.
[30] Sakamoto T,Hirano T,Madsen S S,et al.Insulin-like growth factor I gene expression during parr-smolt transformation of coho salmon[J].Zoological Science,1995,12(2):249-252.
[31] Kiilerich P,Kristiansen K,Madsen S S.Hormone receptors in gills of smolting Atlantic salmon,Salmo salar:Expression of growth hormone,prolactin,mineralocorticoid and glucocorticoid receptors and 11beta-hydroxysteroid dehydrogenase type 2[J].General and Comparative Endocrinology,2007,152(2-3):295-303.
[32] Takei Y,McCormick S.Hormonal control of fish euryhalinity[J].Euryhaline Fishes Fish Physiology,2013,32(8):69-123.
[33] Tipsmark C K,Madsen S S.Distinct hormonal regulation of Na+,K+-atpase genes in the gill of Atlantic salmon(Salmo salar L.)[J].Journal of Endocrinology,2009,203(2):301-310.
[34] McCormick S D,Regish A M,Christensen A K,et al.Differential regulation of sodium-potassium pump isoforms during smolt development and seawater exposure of Atlantic salmon[J].The Journal of Experimental Biology,2013,216(7):1142-1151.
[35] Wood A W,Duan C,Bern H A.Insulin-like growth factor signaling in fish[J].International Review of Cytology,2005,243(6):215-285.
[36] Sheridan M A.Effects of thyroxin,cortisol,growth hormone,and prolactin on lipid metabolism of coho salmon,Oncorhynchus kisutch,during smoltification[J].General and Comparative Endocrinology,1986,64(2):220-238.
[37] Duan C,Plisetskaya E M,Dickhoff W W.Expression of insulinlike growth factor I in normally and abnormally developing coho salmon(Oncorhynchus kisutch)[J].Endocrinology,1995,136(2):446-452.
[38] Stefansson S,Bj9rnsson B,Sundell K,et al.Physiological characteristics of wild Atlantic salmon post-smolts during estuarine and coastal migration[J].Journal of Fish Biology,2003,63(4):942-955.
[39] Specker J L.Interrenal function and smoltification[J].Aquaculture,1982,28(1):59-66.
[40] Fontaine M,Hatey J.Sur la teneur en 17-hydroxycorticosteroides du plasm desaumon(Salmo salar L.)[J].Comptes Rendus de Academie des Sciences de,1954,239(3):319-321.
[41] Specker J L,Schreck C B.Changes in plasma corticosteroids during smoltification of coho salmon,Oncorhynchus kisutch[J].General and Comparative Endocrinology,1982,46(1):53-58.
[42] Shrimpton J M,Devlin R H,McLean E,et al.Increases in gill cytosolic corticosteroid receptor abundance and saltwater tolerance in juvenile coho salmon(Oncorhynchus kisutch)treated with growth hormone and placental lactogen[J].General and Comparative Endocrinology,1995,98(1):1-15.
[43] Nishioka R S,Bern H A,Lai K V,et al.Changes in the endocrine organs of coho salmon during normal and abnormal smoltification—an electron-microscope study[J].Aquaculture,1982,28(1):21-38.
[44] Prunet P,Boeuf G,Bolton J P,et al.Smoltification and seawater adaptation in Atlantic salmon(Salmo salar):Plasma prolactin,growth hormone,and thyroid hormones[J].General and Comparative Endocrinology,1989,74(3):355-364.
[45] Young G,Bj9rnsson B T,Prunet P,et al.Smoltification and seawater adaptation in coho salmon(Oncorhynchus kisutch):Plasma prolactin,growth hormone,thyroid hormones,and cortisol[J].General and Comparative Endocrinology,1989,74(3):335-345.
[46] Yada T,Takahashi K,Hirano T.Seasonal changes in seawater adaptability and plasma levels of prolactin and growth hormone in landlocked sockeye salmon(Oncorhynchus nerka)and amago salmon(O.rhodurus)[J].General and Comparative Endocrinology,1991,82(1):33-44.
[47] Hgsen H R,Prunet P.Plasma levels of thyroxine,prolactin,and cortisol in migrating and resident wild arctic char,Salvelinus alpinus[J].Canadian Journal of Fisheries and Aquatic Sciences,1997,54(12):2947-2954.
[48] Seidelin M,Madsen S S.Prolactin antagonizes the seawater-adaptive effect of cortisol and growth hormone in anadromous brown trout(Salmo trutta)[J].Zoological Science,1997,14(2):249-256.
[49] Seidelin M,Madsen S.Endocrine control of Na+,K+-ATPase and chloride cell development in brown trout(Salmo trutta):Interaction of insulin-like growth factor-I with prolactin and growth hormone[J].Journal of Endocrinology,1999,162(1):127-135.
[50] Hoar W S.The thyroid gland of the Atlantic salmon[J].Journal of Morphology,1939,65(2):257-295.
[51] Dickhoff W W,Folmar L C,Gorbman A.Changes in plasma thyroxine during smoltification of coho salmon, Oncorhynchus kisutch[J].General and Comparative Endocrinology,1978,36(2):229-232.
[52] Dickhoff W W,Sullivan C V.Involvement of the thyroid gland in smoltification,with special reference to metabolic and developmental processes[J].American Fisheries Society Symposium,1987,1(6):197-210.
[53] Iwata M,Tsuboi H,Yamashita T,et al.Function and trigger of thyroxine surge in migrating chum salmon Oncorhynchus ketafry[J].Aquaculture,2003,222(1-4):315-329.
[54] McCormick S D,O'Dea M F,Moeckel A M,et al.Endocrine and physiological changes in Atlantic salmon smolts following hatchery release[J].Aquaculture,2003,222(1-4):45-57.
[55] Hutchison M J,Iwata M.Effect of thyroxine on the decrease of aggressive behaviour of four salmonids during the parr-smolt transformation[J].Aquaculture,1998,168(1):169-175.
[56] Iwata M.Downstream migratory behavior of salmonids and its relationship with cortisol and thyroid hormones:A review[J].Aquaculture,1995,135(1):131-139.
[57] Ojima D,Iwata M.The relationship between thyroxine surge and onset of downstream migration in chum salmon Oncorhynchus ketafry[J].Aquaculture,2007,273(2-3):185-193.
[58] Ojima D,Iwata M.Central administration of growth hormone-releasing hormone and corticotropin-releasing hormone stimulate downstream movement and thyroxine secretion in fall-smolting coho salmon(Oncorhynchus kisutch)[J].General and Comparative Endocrinology,2010,168(1):82-87.
[59] McCormick S D.Endocrine control of osmoregulation in teleost fish[J].American Zoologist,2001,41(4):781-794.
[60] Shrimpton J M,McCormick S D.Regulation of gill cytosolic corticosteroid receptors in juvenile Atlantic salmon:Interaction effects of growth hormone with prolactin and triiodothyronine[J].General and Comparative Endocrinology,1998,112(2):262-274.
[61] Taylor E B.Environmental correlates of life-history variation in juvenile chinook salmon,Oncorhynchus tshawytscha(Walbaum)[J].Journal of Fish Biology,1990,37(1):1-17.
[62] McCormick S D.Ontogeny and evolution of salinity tolerance in anadromous salmonids:Hormones and heterochrony[J].Estuaries,1994,17(1):26-33.
[63] Gallagher Z S,Bystriansky J S,Farrell A P,et al.A novel pattern of smoltification in the most anadromous salmonid:pink salmon(Oncorhynchus gorbuscha)[J].Canadian Journal of Fisheries and Aquatic Sciences,2013,70(3):349-357.
[64] Zaugg W S,Mclain L R.Influence of water temperature on gill sodium,potassium-stimulated ATPase activity in juvenile coho salmon(Oncorhynchus kisutch)[J].Comparative Biochemistry and Physiology Part A:Physiology,1976,54(4):419-421.
[65] Duston J,Saunders R,Knox D.Effects of increases in freshwater temperature on loss of smolt characteristics in Atlantic salmon(Salmo salar)[J].Canadian Journal of Fisheries and Aquatic Sciences,1991,48(2):164-169.
[66] McCormick S D,Shrimpton J M,Zydlewski J D.Temperature effects on osmoregulatory physiology of juvenile anadromous fish[J].Researche and Exploration in Laboratory,1996,46(3):170-188.
[67] Wagner H H.Photoperiod and temperature regulation of smolting in steelhead trout(Salmo gairdneri)[J].Canadian Journal of Zoology,1974,52(2):219-234.
[68] Zydlewski G B,Haro A,McCormick S D.Evidence for cumulative temperature as an initiating and terminating factor in downstream migratory behavior of Atlantic salmon(Salmo salar)smolts[J].Canadian Journal of Fisheries and Aquatic Sciences,2005,62(1):68-78.
[69] Staurnes M,Lysfjord G,Hansen L,et al.Recapture rates of hatchery-reared Atlantic salmon(Salmo salar)related to smolt development and time of release[J].Aquaculture,1993,118(3-4):327-337.
[70] Lundqvist H,McKinnell S,Fngstam H,et al.The effect of time,size and sex on recapture rates and yield after river releases of Salmo salar smolts[J].Aquaculture,1994,121(1-3):245-257.
[71] Bilton H,Alderdice D,Schnute J.Influence of time and size at release of juvenile coho salmon(Oncorhynchus kisutch)on returns at maturity[J].Canadian Journal of Fisheries and Aquatic Sciences,1982,39(3):426-447.
[72] Virtanen E,S9derholm-Tana L,Soivio A,et al.Effect of physiological condition and smoltification status at smolt release on subsequent catches of adult salmon[J].Aquaculture,1991,97(2):231-257.
[73] McCormick S D,Hansen L P,Quinn T P,et al.Movement,migration,and smolting of Atlantic salmon(Salmo salar)[J].Canadian Journal of Fisheries and Aquatic Sciences,1998,55(S1):77-92.
[74] Orciari R D,Leonard G H.Length characteristics of smolts and Timing of downstream migration among three strains of Atlantic salmon in a Southern new england stream[J].North American Journal of Fisheries Management,1996,16(4):851-860.
[75] McGinnity P,de Eyto E,Cross T F,et al.Population specific smolt development,migration and maturity schedules in Atlantic salmon in a natural river environment[J].Aquaculture,2007,273(2-3):257-268.
[76] Handeland S O,Wilkinson E,SveinsbB,et al.Temperature influence on the development and loss of seawater tolerance in two fast-growing strains of Atlantic salmon[J].Aquaculture,2004,233(1-4):513-529.
[77] Banks J L.Raceway density and water flow as factors affecting spring chinook salmon(Oncorhynchus tshawytscha)during rearing and after release[J].Aquaculture,1994,119(2):201-217.
[78] Miyakoshi Y,Nagata M,Kitada S.Effect of smolt size on postrelease survival of hatchery-reared masu salmon Oncorhynchus masou[J].Fisheries Science,2001,67(1):134-137.
[79] Connor W P,Smith S G,Andersen T,et al.Postrelease performance of hatchery yearling and subyearling fall Chinook salmon released into the Snake River[J].North American Journal of Fisheries Management,2004,24(2):545-560.
[80] Saloniemi I.Survival of reared and wild Atlantic salmon smolts:Size matters more in bad years[J].ICES Journal of Marine Science,2004,61(5):782-787.
[81] Dieperink C.Predation on Atlantic salmon and sea trout during their first days as postsmolts[J].Journal of Fish Biology,2002,61(3):848-852.
[82] Jrvi T.Cumulative acute physiological stress in Atlantic salmon smolts:The effect of osmotic imbalance and the presence of predators[J].Aquaculture,1990,89(3):337-350.
[83] Handeland S,Jrvi T,Fern9A,et al.Osmotic stress,antipredatory behaviour,and mortality of Atlantic salmon(Salmo salar)smolts[J].Canadian Journal of Fisheries and Aquatic Sciences,1996,53(12):2673-2680.
[84] Price C S,Schreck C B.Stress and saltwater-entry behavior of juvenile chinook salmon(Oncorhynchus tshawytscha):Conflicts in physiological motivation[J].Canadian Journal of Fisheries and Aquatic Sciences,2003,60(8):910-918.
[85] Schreck C B,Stahl T P,Davis L E,et al.Mortality estimates of juvenile spring-summer chinook salmon in the lower columbia river and estuary,1992-1998:Evidence for Delayed Mortality?[J].Transactions of the American Fisheries Society,2006,135(2):457-475.
[86] Loge F J,Arkoosh M R,Ginn T R,et al.Impact of environmental stressors on the dynamics of disease transmission[J].Environmental Science&Technology,2005,39(18):7329-7336.
[87] Schreck C B,Solazzi M F,Johnson S L,et al.Transportation stress affects performance of coho salmon,Oncorhynchus kisutch[J].Aquaculture,1989,82(1):15-20.
[88] Staurnes M,Sigholt T,Asgard T,et al.Effects of a temperature shift on seawater challenge test performance in Atlantic salmon(Salmo salar)smolt[J].Aquaculture,2001,201(1-2):153-159.
[89] Duston J.Effect of salinity on survival and growth of Atlantic salmon(Salmo salar)parr and smolts[J].Aquaculture,1994,121(1-3):115-124.
[90] Duncan N J,Auchinachie N,Robertson D,et al.Growth,maturation and survival of out-of-season 0+and 1+Atlantic salmon(Salmo salar)smolts[J].Aquaculture,1998,168(1):325-339.
[91] Johnsson J,Clarke W.Development of seawater adaptation in juvenile steelhead trout(Salmo gairdneri)and domesticated rainbow trout(Salmo gairdneri)-effects of size,temperature and photoperiod[J].Aquaculture,1988,71(3):247-263.
[92] Sedgwick S D.Rainbow trout farming in Scotland,Farming trout in salt water[J].Scott Agric,1970,49:180-185.
[93] Landless P J.Acclimation of rainbow trout to sea water[J].Aquaculture,1976,7(2):173-179.
[94]董双林.论我国水产养殖业生态集约化发展[J].中国渔业经济,2015,33(5):4-9.Dong S L.On ecological intensification of aquaculture systems in China[J].Chinese Fisheries Economics,2015,33(5):4-9.
[95]韩立民,郭永超,董双林.开发黄海冷水团建立国家离岸养殖试验区的研究[J].太平洋学报,2016,24(5):79-85.Han L M,Guo Y C,Dong S L.Research on establishing a national offshore aquaculture experimental zone based on the development of the Yellow Sea Cold Water Mass[J].Pacific Journal,2016.24(5):79-85.
[96] Iwata M,Ogura H,Komatsu S,et al.Changes in salinity preference of chum and coho salmon during development[J].Aquaculture,1985,45(1-4):380-381.
[97] Clarke W C,Nagahama Y.Effect of premature transfer to sea water on growth and morphology of[J].Canadian Journal of Zoology,1977,55(10):1620-1630.
[98] Folmar L C,Dickhoff W W,Mahnken C V W,et al.Stunting and parr-reversion during smoltification of coho salmon(Oncorhynchus kisutch)[J].Aquaculture,1982,28(1-2):91-104.
[99] Boeuf G,Harache Y.Criteria for adaptation of salmonids to high salinity seawater in France[J].Aquaculture,1982,28(1):163-176.
[100] Koch H J A.Hemoglobin changes with size in the Atlantic salmon(Salmo salar L.)[J].Aquaculture,1982,28(1-2):231-240.