17_20_二羟黄体酮降低虹鳟鱼血浆免疫球蛋白M的水平_英文_

17_20_二羟黄体酮降低虹鳟鱼血浆免疫球蛋白M的水平_英文_ CN 53 - 1040/ Q ISSN 0254 - 5853 动 物 学 研 究 2000 , Apr. 21 ( 2) : 97, 102 Zoological Research Ξαβ17, 202D I HYD ROXY242P REGNENE232O NE S UPP RESSES PL ASMA IgM L EVELS IN RAINBOW TRO UT ( O ncor hynch us mykiss) HOU Ya2yi HAN Xiao2do ng ( )S chool of Medici ne , N a nji n g U ni versi t y , N a nji n g 210093 , Chi na SUZUKI Yuzuru ( )Depa rt ment of A quat ic B iosciences , T he U ni versi t y of Tokyo , Tokyo 113 , J apa n αβαβ() Abstract : The relatio nship s bet ween 17,202di hydro xy242p regnene232o ne 17,202D Pand plasma IgM and to2 tal p rotein levels were investigated in rainbow t rout , O ncorhy nch us m ykiss . IgM and total p rotein levels decreased in bot h αβαβsexes of mat ure rainbow t rout when 17,202D P levels increased during t he spawning seaso n ,while t he elevated 17,202 D P supp ressed IgM levels and reversibly enhanced total p rotein co ncent ratio ns in immat ure t rout . This rep resent s t he first αβαβreport of t he effect of 17,202D P o n fish immunity. These data suggest t hat t he increase of 17, 202D P in spawning seaso n may be related to infectious skin diseases. αβKey words : Rainbow t rout ; Immunoglobulin M ;17,202di hydro xy242p regnene232o ne ; Plasma total p rotein co ncent ratio n ) 1998 ; Ho u et al 1 , 1998 ; besides , steroid ho r mo nes The interactio n bet ween steroid ho r mo nes and ( ( immune f unctio ns was reviewed in mammals Schu2 can supp ress plasma IgM levels Ho u et al 1 , 1999a , ) αβαβ) ( urs et al 1 ,1990. Female mice and hamsters generat2 b. 17, 202dihydro xy242p regnene232o ne 17, 202 ) D Pis co nsidered as a role in t he final gamete mat ura2 ed higher titers of t he immunoglo bulins t han males , ( tio n of bot h male and female salmo nid fish Kime , and t he lessening of antibo dy p ro ductio n in males re2 ) 1993 ; Nagahama , 1997 , and has been identified spo nded to t he increase in sex steroid ho r mo nes at ( ) as t he mat uratio n2inducing ho r mo ne of several fish sexual mat urit y Gro ssman , 1985 . Est rogen might also stimulate t he mammalian humo ral , o r antibo dy species , including salmo nid fishes , inducing meiotic respo nse o r enhance t he mammalian immune respo nse mat uratio n of fish oocytes. Mo reover , o ne of t he ( ) Paavo nen et al 1 ,1981 ; Hu et al 1 ,1988. However , majo r p ro blems in hatchery is t he o ut break of skin w hen t reated wit h t he androgenic male steroid ,it was diseases , w hich may cause high mo rtalit y in rainbow fo und t hat testo stero ne co uld also influence im2 t ro ut in spaw ning seaso n . αmunoglo bulin synt hesis and inhibit t he antibo dy re2 βThe effect s of 17,202D P o n plasma IgM levels ( spo nse of mammalian Fuji et al 1 ,1975 ; St hoeger et and total p rotein co ncent ratio ns were t herefo re inves2 ) al 1 , 1988 . It seems po ssible t hat est rogen can en2 tigated in rainbow t ro ut . hance t he immunit y in mammals and androgen sup2 1 Material s and Methods p ress it . IgM is know n to be t he o nly class of teleo st im2 111 Experimental project ( munoglo bulin . In rainbow t ro ut O ncorhy nch us m y2 αβ 11111 Seaso nal changes in 17,202D P , IgM and to2) kiss,t he decrease in plasma IgM levels was also o b2 tal p rotein levels ( served in spaw ning seaso n Suzuki et al 1 ,1997 ; Ho u , ( Mat ure female average bo dy weight :1 23713 ? Ξ 收稿日期 : 1999 - 06 - 23 ; 修改稿收到日期 : 1999 - 09 - 28 Fo undatio n item : This st udy was suppo rted by J apan Fishery Agency αβ) (,202D P used fo r standards were p urchased f ro m 1737718 gand male average bo dy weight : 1 26317 ? ( ) ) Sigma Chemical Co mpany U SA . The rabbit anti2 21414 g rainbow t ro ut were used in t he p resent st udy . These fish were reared in a co ncrete tank wit h αβ17, 202dihydro xy242p regnene232o xime2B SA serum was p urchased f ro m Teiko kozo ki Phar maceutical running sp ring water at abo ut 10 ?,under nat ural day () αβlight ,at t he Oizumi Fisheries Research L abo rato ry of Co mpany J apan. The cro ss reactivit y of 17,202di2 αTo kyo U niversit y of Fisheries , Oizumi , Yamanashi βhydro xy242p regnene232o xime2B SA wit h 17, 202di2 Prefect ure. Samples were o btained at abo ut mo nt hly ββαβhydro xy242p regnene232o ne , 52p regnane23, 17, 202 intervals f ro m J une 1996 to February 1997 . A rando m βαt riol ,2 02hydro xyp rogestero ne and 1 72hydro xyp ro2 selectio n of fish was made f ro m t he fish pop ulatio n in gestero ne were 1 0 0 , 2 15 4 , 1 15 5 , 0 1 8 2 , 0 1 0 2 8 and < 0101 % , respectively. The int ra2assay and inter2as2 t he samples. Three fish of each sex were net ted ,anes2 ( ) t hetized immediately wit h 22p heno xyet hanol o n each say coefficient s of variatio n were 610 % n = 6 and ( ) occasio n. Bloo d samples were taken f ro m t he caudal 711 % n = 7. Recovery rate was 9519 %. The low2 vascult ure using a heparinized syringe wit h 5 min af2 est limit of detectio n was 45 p g/ mL . ( ) ter net ting. Bloo d samples were separated by cent rif u2 11212 Enzyme immuno so rbent assay E IAof IgM The assays were carried o ut in 962well polyst yre2 gatio n at 1 000 ×g at 4 ? fo r 10 min . Plasmas were ( ) ne microtiter plates N unc . The microtiter plates t hen o btained and sto red at - 25 ? until being used. ααβ were coated wit h p urified rabbit anti2rainbow t ro ut 11112 Effect s of 172P o n 17,202gM and to2D P , I IgM Ig G t hat was dissolved in 011 M carbo nate tal p rotein levels (μ) J uvenile rainbow t ro ut used fo r t his st udy were buffer p H 916. A volume of 100L was dispensed o btained f ro m a local f ar m of To kyo . Their bo dy into each well and incubated at roo m temperat ure . Af2 μter 42times wash wit h PB S2Tween , 300 L of Block weight s ranged f ro m 42 g to 76 g. The fish were ( ) Ace diluted wit h PB S 1 ?4was added to each well reared in 802liter ro und plastic tanks supplied wit h fil2 and incubated at roo m temperat ure . Befo re being tered circulating water and p hotoperio d kep t at 14L ? μwashed again ,100L of plasma samples and standard 10D . Twent y fish were equally separated into t wo IgM samples were placed into t he app rop riate wells (tanks o ne as t he co nt rol , anot her as t he experimen2 and incubated over night at 4 ?. Af ter washing , each ) tal. The fish were acclimatized fo r 2 weeks at 10 ? μwell received 100 L of t he mixt ure of a drop of A befo re experiment . During t his perio d , fish were fed (and B f ro m ABC kit avidin and biotinlated ho rsera2 o nce a day wit h co mmercial t ro ut pellet s. Af ter 22 ) dish pero xidase macro molecular co mplex ,Vecto rdis2 αweek acclimatizatio n , a diet co ntaining 1 mg 172P/ g solved in 10 mL PB S2Tween ,and plates were incubat2 diet , o r a co nt rol diet was substit uted fo r t he t wo ed at roo m temperat ure . The plates were washed and gro up s respectively. Diet was p repared as follow s : ) ( μ( 100L of TMB 3 , 3 , 5 , 52tet ramet hylbenzidine1 α 172P was dissolved in 100 % et hanol and t hen t he so2 mg TMB dissolved in 10 mL of 011 M ,p H 515 cit ric lutio n was sp rayed o nto foo d pellet s ; t he solvent was μ) acid buffer co ntaining 115 L of 30 % HOwere 2 2 t hen allowed to evapo rate by air2dry. The co rre2 added to each well fo r enzymatic colo ur reactio n at spo nding co nt rol diet was p repared by sp raying t he roo m temperat ure . The reactio n was stopped af ter 20 t ro ut pellet s wit h et hanol o nly. The experimental di2 μmin by adding 25L of 2 M HSO. The abso rbance 2 4 et was given o nce each fo r a perio d of 5 weeks. Bloo d was measured at 450 nm using an EL ISA Plate reader samples were taken f ro m each fish of each gro up o nce ( ) M PR 4A9 To so u. a week using t he same p rocedure as above . 11213 Q uantitatio n of total p rotein co ncent ratio n 112 Experimental methods Total p rotein co ncent ratio ns were measured us2 ( )11211 Steroid ho r mo ne radioimmunoassay R IA (ing t he Bio2Rad Protein Assay kit Bio2Rad , So ut h ( αβ) L abeled 2 , 4 , 6 , 723 H17, 202D P was p ur2) ( Rich2mo nd , CA wit h bovine serum albumin 2 0 0 , ( ) chased f ro m New England N uclear U SA. U nlabeled μ) 400 , 800 , 1 000 , 1 200 and 1 400g/ mL as t he standard. 11214 Statistical analysis St udent t2test was employed to evaluate t he dif2 ference amo ng t he levels of IgM o r steroid ho r mo nes in each sample . The differences of means amo ng t he gro up s in each sample occasio n were analyzed by Dun2 can’s multiple range test . Co rrelatio n coefficient s were calculated bet ween IgM co ncent ratio n and plas2 αβ ma 17,202D P. 2 Results αβ211 Sea sonal changes in 17, 202D P , IgM and total protein level s αβPlasma 17,202D P levels in mat ure females re2 ( mained low f ro m J une to Sep tember below 115 ng/ ) mL , but started to increase in Octo ber wit h a peak ( ) o bserved in November 1715 ?5109 ng/ mL , w hen t he spaw ning co mmenced. Then t he ho r mo ne level decreased to low levels as o bserved in p re2spaw ning ( ) seaso n Fig11A . Changes in plasma IgM levels in ( ) mat ure female were showed Fig11B. Plasma IgM levels declined f ro m Sep tember , wit h it s minimum () noted in November 0190 ?0137 mg/ mL , t hen re2 sto red f ro m December . The changes in plasma total p rotein levels in mat ure female were showed ) ( Fig11C. Plasma total p rotein level began to rise ( f ro m August , peaked in Sep tember 146148 ?9106 ) mg/ mL , decreased f ro m Octo ber , and reached it s () minimum in J anuary 75169 ?7180 mg/ mL . αβIn mat ure males , plasma 172, 20D P level re2 αβFig11 Changes of 17,202di hydro xy242p regnene232o ne ( ) mained low f ro m J une to August abo ut 1 ng/ mL , () () ( ) A, IgM B,total p rotein Clevels in mat ure ( but t wo peaks were o bserved in Sep tember 15123 ? female rainbow t rout during spawning seaso n Each point rep resent s mean ?S E. 3 P < 0105 co mpared to ot her ) (1184 ng/ mL and o n December 4 17120 ?3167 ng/ sampling point s o n t he same line . ) ( ) mL , t hen decreased Fig12A . Plasma IgM co n2 cent ratio n fell gradually f ro m August to it s lowest le2 ( levels in bot h sexes r > - 01797 , P < 01001 in fe2 () vel o n December 4 0176 ?0119 mg/ mL , and t hen ) males ; r > - 01477 , P < 0105 in males. ro se slowly until t he end of t he experiment ααβ212 Eff ects of 172P on 17, 202D P , IgM and ( ) Fig12B . Plasma total p rotein level began to de2 total protein level s α crease f ro m Octo ber , and showed no recovery till t he The administ ratio n wit h 1 72 P caused evident ( ) end of t he experiment Fig12C. αβ,202D P co ncent ratio ns ,co m2 elevatio n in plasma 17 Significantly negative co rrelatio ns were o bserved pared wit h t ho se in t he co nt rol af ter 2 weeks. The αα ββbet ween Plasma 1 7, 2 02D P levels and plasma IgM elevated 17,202D P levels remained until t he end of αβFig13 Changes of plasma 17,202di hydro xy242p regnene 32 αβ12 Changes of 17,202di hydro xy242p regnene 32o ne Fig () () ( ) o ne A, IgM B,total p rotein Clevels wit h t reat2 () () ( ) A, IgM B,total p rotein Clevels in mat ure α ment of 172hydro xyp rogestero nemale rainbow t rout during spawning seaso n ? t he co nt rol gro up ; ? t he experimental gro up . Each point rep re2 Each point rep resent s mean ?S E. 3 P < 0105 co mpared to ot her sent s mean ?S E. Point s mar ked 3 are significantly different sampling point s o n t he same line . f ro m t he co nt rol fo r P < 0105 . αβt he experiment , w hile 17,202D P levels in t he co n2 3 Discussion ( ) t rol gro up showed no change Fig13A . Co mpared αβThe changes of 17, 202D P levels in female wit h t he co nt rol af ter 1 week , plasma IgM levels were supp ressed , alt ho ugh IgM levels also declined rainbow t ro ut were similar to t ho se in so me salmo ns ) ( Fig13B. In co nt rast , plasma total p rotein co ncen2 (Dye et al 1 , 1986 ; Kime , 1993 ; Liley et al 1 , 1986b ; αt ratio ns were elevated af ter 3 weeks wit h 172P t reat2 α) Ueda et al 1 ,1983 ; Yo ung et al 1 ,1983. Plasma 17, ment and co ntinued to rise till t he end of t he experi2 β202D P levels were low in t he p ro2spaw ning seaso n , ( ) ment Fig13C. rapidly increased at t he beginning of t he spaw ning seaso n ,and fell p recipito usly af ter ovulatio n in female co rrelated wit h plasma IgM in bot h sexes of mat ure rainbow t ro ut . In male rainbow t ro ut , t he changes of t ro ut in spaw ning seaso n , and severe skin infectio us αβαβ17,202D P levels were different f ro m t ho se in so me t he highest level of 17, 202 disease respo nded to ( IgM level f ro m August to De2 D P. Mo reover , low salmo n w hich were st udied p revio usly Dye et al 1 , αβ ) 1986 ; Kime , 1993 ; Liley et al 1 , 1986a . This differ2 cember co rrespo nded to t wo peaks of 17, 202D P wit hin t he perio d in male . Furt her mo re , t he elevat2 ence may be at t ributed to t he different st rains of fish , αβed 17,202D P significantly reduced plasma IgM le2 in w hich changes in plasma levels of each steroid had vels in immat ure t ro ut , w hich were reared at 15 ?. ( ) respectively annual pat ter ns Suzuki et al 1 , 1997 . αβIt is demo nst rated t hat 17, 202D P may supp ress The p hysiological significance of t he sharp increase in plasma IgM levels and t he decreased plasma IgM may αβplasma 17, 202D P levels at t he time of sper miatio n be o ne of f acto rs causing infectio us diseases in rainbow α is yet unknow n . So me st udies revealed t hat t he 17, t ro ut . β202D P might have a majo r role in t he inductio n of αβThe effect of 17, 202D P o n total p rotein co n2 ( oocyte mat uratio n in several species of teleo st Ueda cent ratio n was not elucidated clearly in t his st udy. ) et al 1 ,1983 ; Nagahama ,1997and act as a p heno me2 The increase in plasma total p rotein levels in mat ure ( et al 1 , nal at t ractant fo r male salmo n Dit t man β female may be caused by 172est radiol , since it s in2) 1993. crease may be related to vitellogenesis. The decreases The decreased plasma IgM may be related to t he in t he total p rotein levels may be at t ributed to t he in2 αβincrease in plasma 17, 202D P levels in spaw ning αβcreases in plasma 17, 202D P in bot h sexes of ma2 seaso n . The effect of water temperat ure o n IgM levels t ure t ro ut s during t he spaw ning seaso n . However , it may not be co nsidered in t he st udy , since t he fish αβis wo rt h to questio n w hy t he elevated 17, 202D P were reared under a co nstant water temperat ure . The increased t he total p rotein levels in immat ure t ro ut . immuno supp ressio n of steroid ho r mo nes —co rtisol , αβIt may be po ssible t hat 17, 202D P administ ratio n βtesto stero ne , 172est radiol and 112ketotesto stero ne changed t he metabolism of p roteins in immat ure ( have been repo rted p revio usly Slater et al 1 , 1993 ; βt ro ut , since it is know n t hat est radiol2172t reated can Slater et al 1 ,1995 ; Suzuki et al 1 ,1997 ; Ho u et al 1 , induce t he synt hesis of vitellogenin in immat ure fish , ) 1999a , b . The recent st udy revealed t hat fish im2 mune f unctio n may be related to p hotoperoid in im2 and t he increased vitellogenin may not be no r mal to ( ) mat ure rainbow t ro ut Ho u et al 1 , 1999c , but ( 2 immat ure fish Ko bayashi , perso nal co mmunica αβchanges of 17, 202D P levels can not be deter mi ned ) tion. However , mo re info r matio n is needed o n t he αβeffect of 17,202D P in t he metabolism of p rotein in αβin t he immat ure fish . 17, 202D P is co nsidered as a role in t he final gamete mat uratio n of bot h male and fish . αβfemale salmo nid fish ,and is also identified as t he mat2 In co nclusio n , 17, 202D P may supp ress fish uratio n2inducing ho r mo ne of several fish species immunoco mpetence . The decreased plasma IgM may ( ) α be o ne of f acto rs causing infectio us diseases of rainbow Kime ,1993 ; Nagahama ,1997, so t he effect of 17, β 202D P sho uld be also co nsidered in mat ure fish .t ro ut in spaw ning seaso n . αβ We o bserved t hat 17, 202D P was negatively Ref erences Dit t man A H , Quinn T P , 1993 . Avoidance of a p utative p hero mo ne , Dye H M ,Sumpter J P , Fagerlund U H M et al , 1986 . Changes in re2 αβ 17, 202dihydro xy242p regnene232o ne , by p recocio usly mat ure chi2p ro ductive parameters during t he spawning migratio n of pink ) ( ) ( noo k salmo n O ncorhy nch us tshaw ytscha J . Ca n . J . Zool . , 72 : salmo n , O ncorhy nch us gorbuscha WalbaumJ . J . Fish B iol . , 215 - 219 . 29 :167 - 176 . Fuji H ,Nawa Y , Tsuchiya K M et al ,1975 . Effect of a single adminis2 inducing ho r mo ne in fish oocytes : Mechanisms of synt hesis and ac2 ( ) t ratio n of testo stero ne o n t he immune respo nse and lymp hoid tissues tio n J . S teroi ds ,62 1:190 - 196 . in mice J . Cel l . I m m u nol . ,20 :315 - 326 . Paavo nen T ,Andersso n L C ,Adlercreutz H ,1981 . Sex ho r mo ne regula2 Gro ssman C J , 1985 . Interactio ns bet ween t he go nadal steroids and t he tio n of i n v i t ro immune respo nse J . J . Ex p . Med . , 154 : 1935 - immune systemJ . S cience ,227 :257 - 260 . 1945 . Ho u Y , 1998 . Endocrinological aspect s of reduced immunoco mpetence Schuurs A H W M , Ver heul H A M , 1990 . Effect s of gender and sex steroids o n t he immune respo nsesJ . J . S teroi d B ioche m . ,35 :157 wit h go nadal mat uratio n in rainbow t ro ut D . Thesis of t he Doc 2 to rate . The U niversit y of To kyo . - 172 . Slater C H ,Schreck C B ,1993 . Testo stero ne alters t he immune respo nse Ho u Y ,Suzuki Y ,Aida K ,1998 . Effect s of steroid ho r mo nes o n antibo dy p ro ducing activit y in rainbow t ro ut J . Dev . Com p . I m m u nol . ,22 of chinoo k salmo n , O ncorhy nch us tshaw ytscha J . Gen . Com p . En2 ( ) docri nol . ,89 :291 - 298 . 1:137 . Ho u Y ,Suzuki Y ,Aida K ,1999a . Changes in endocrine and immune ac2 Slater C H , Fitzpat rick M S , Schreck C B , 1995 . Androgens and im2 munoco mpetence in salmo nids : specific binding in a reduced immuno2 tivities during t he spawning seaso n in rainbow t ro ut J . Recent Prog ress i n M olecul a r a n d Com p . En docri n . ,442 - 447 . co mpetence of salmo nid lymp hocytes expo sed to nat ural and synt he2 Ho u Y ,Suzuki Y ,Aida K ,1999b. Effect s of steroid ho r mo nes o n IgM in tic androgensJ . A quacul t u re ,136 :363 - 370 . ( ) rainbow t ro ut J . Fish Physiol . B ioche m . ,20 2:155 - 162 . St hoeger Z M ,Chio razzi N ,L ahita R G ,1988 . Regulatio n of t he immune respo nse by sex ho r mo nes :1 . i n v i t ro effect s of est radiol and testo s2 Ho u Y , Han X D ,Suzuki Y et al ,1999c . Immune f unctio n of immat ure ( ) rainbow t ro ut O ncorhy nch us m ykiss related wit h p hotoperio d tero ne o n po kewwd mitogen2induced human B cell differentatio n ( ) J . Zoological Resea rch ,20 6:401 - 405 . J . J . I m m u nol . ,141 :91 - 98 . Suzuki Y , Otaka T , Sato S et al , 1997 . Rep ro ductio n related im2 Hu S K ,Mitcho Y L , Rat h N C , 1988 . Effect of est radiol o n t he inter2 leukin 1 synt hesis by macrop hages J . I nt . J . I m m u nop ha r m a2 munoglo bulin changes in rainbow t ro ut J . Fish Physiol . col . ,10 :247 - 252 . B ioche m . ,17 :415 - 421 . αβ Kime D E , 1993 .“Classical”and“no n2classical ”rep ro ductive steroids in Ueda H , Yo ung G , Crim L W et al , 1983 . 17, 202dihydro xy242p reg2fish J . Rev . Fish B iol . Fisheries ,3 :160 - 180 . nene232o ne : Plasma levels during sexual mat uratio n and i n v i t ro Liley N R ,Breto n B , Fo stier A et al ,1986a . Endocrine changes associat2 ( p ro ductio n by t he testes of amago salmo n O ncorhy nch us rhod u2 ed wit h spawning behavio r and social stimuli in a wild pop ulatio n of ) ( ) r usand rainbow t ro ut S al m o gai r d neri J . Gen . Com p . En2 ( ) rainbow t ro ut S al m o gai r d neri . ?. Males J . Gen . Com p . En2 docri nol . ,51 :106 - 112 . docri nol . ,62 :145 - 156 . αβYo ung G , Crim L W , Kagawa H et al , 1983 . Plasama 17, 202dihy2 Liley N R ,Breto n B , Fo stier A et al ,1986b. Endocrine changes associat2 dro xy242p regnene232o ne levels during sexual mat uratio n of amago ed wit h spawning behavio r and social stimuli in a wild pop ulatio n of ( ) salmo n O ncorhy nch us rhod u r us : co rrelatio n wit h plasma go2 ( ) rainbow t ro ut S al m o gai r d neri . ?. Females J . Gen . Com p . nadot ropin and i n v i t ro p ro ductio n by ovarian follicles J . Gen . En docri nol . ,62 :157 - 167 . Com p . En docri nol . ,51 :96 - 106 . αβ Nagahama Y ,1997 . 17,202dihydro xy242p regnene232o ne ,a mat uratio n2 αβ17, 20- 二羟黄体酮降低虹鳟鱼血浆免疫球蛋白 M 的水平 侯亚义 韩晓冬铃木让 ( ( ))南京大学医学院 南京 210093东京大学鱼类生理研究室 日本东京都 αβ αβ 摘要 : 本文首次报道 17, 20- 二羟黄体酮对 17, 20- 二羟黄体 可能与产卵或排精时高水平的 α 虹鳟免疫功能的影响 。在对生殖季节虹鳟血浆17,酮对虹鳟生理的调节有关 。而且 , 当人为地提高虹 ααββ 20- 二羟黄体酮 、免疫球蛋白 M 和总蛋白水平的 鳟血浆17, 20- 二 羟 黄 体 酮 的 水 平 时 发 现 , 17, βαβ 调查时发现 , 雌雄虹鳟血浆 17, 20- 二羟黄体酮 20- 二羟黄体酮能降低免疫球蛋白 M 的水平 , 相 αβ 的水平与免疫球蛋白 反地提高总蛋白的水平 。这就进一步证 实 M 的水平负相关 , 即在雌性 17, 20 αβ 相 关 系 数 r > - 0 1 7 9 7 , P < 0 1 0 1 ; 而 在 雄 性 r > - 二羟黄体酮的免疫抑制作用 。但 17, 20- 二羟 αβ- 01477 , P < 0105 。这提示17,20- 二羟黄体酮可 黄体酮与总蛋白之间的作用还不明了 。这些结果提 αβ能抑制免疫球蛋白 M 的合成或分泌 。产卵或排精 示 , 17,20- 二羟黄体酮可能与生殖季节 虹 鳟 的 高皮肤病发生率有关 。 后的血浆总蛋白量低于任何季节 , 这种低蛋白水平 αβ关键词 : 虹鳟 ; 免疫球蛋白 M ; 17,20- 二羟黄体酮 ; 血浆总蛋白 + 文献标识码 : A() 中图分类号 : Q959146 8 文章编号 : 0254 - 5853 200002 - 0097 - 06