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Departmental Bulletin Paper / 紀要論文
Effect of temperature and salinity on egg
hatch of the rainbow trout <special>Salmo
gairdneri</special>
ニジマス卵の孵化に及ぼす水温と塩分の影響
Kashiwagi, Masaaki; Iwai, Toshio; Saji, Atsushi
三重大学水産学部研究報告 = Bulletin of the Faculty of Fisheries, Mie
University. 1987, 14, p. 41-48.
http://hdl.handle.net/10076/3541
Bull.Fac.Fish‥Mie Univ.
No.14:11】19
September30.1987
First Matur‡ty andl両tia[Growth of Some Common
SpecleS Of Bai’naClesinJapan.
ToshiakiIwAKIand HideshiHATTORI
Faculty of Fisheries,Mie University
11hebまologlCalminimum sizesand growth ratesoぎsまⅩSPeCiesofbarnacles
havebeendeもerminedbyexamirlatまonorspecimenstakenfromraftsorintertまdaユ
piles and by measuring the rostrかCarinaldiame廊Or animals growing on
glasspanelssuspendedfromarart.
ThesizeandageatrirstmaturltyWeredeterminedasfollows;及£mproぴ言古㍑ぶ
4.5mmbasaldiameも饉(17days),鼠αJわgど0ぶとα£㍑S5.2mm(17days),且reとよc比gαとαざ
7.5mm(?),且孔明痛かぬ7.3mm(22days),β.eわ比㌻花e㍑β12月mm(50days)and鼠ゐ0花ぬ−
hoL)ilO.Omm(?).Fromcornparisonoftheseresultswithothcrbarnacles,itfo110WS
that au ofthe barnacles sもudied here show early age and smaller size aもfirst
maもurlty.Iniもは1growもhin rostro−Carinaldまrectionis slow unti16days orso
arter seもtlemenも,barnacles reaching・aPprOXimatelylmmまn basaldiameter・
From6to18daystherat80fgrowth or any species accelerates and barnacles
become3.5to5.Ommlarge at the end ofもhis period・Subsequent decline of the
growth rateisexpectedもO be dueもo the sexualmaもurまもy・UntillO days afも群
metamorphosis barnacles tend to expandもhe basalmargln rather thanもhe
OPerCularportion.
Keyword:Breeding・,Growth,鮎rnacles
In previous papers(IwAiく1,1975;1981),it has already been clarifiedもhat the adult
・!・.∴∴. 一丁‥・・∴∴.、∫.・・∴..り、− こ、:∴・−∴−∴・..・.、−・.・‥−∴・−・二、・‥J・・・、、●‥・・・ご・‥∴ヾ
、; −・ご・、.‥.−‥‥・.‥∴・.、.・ J∴・∴・・・∴ご・1.・.ご・、∴ 二 !.・.‥・‥.、.∴‥∴、.、
TARASOV &ZiミVINA andβα∼α花捉ぶrgと£c昆gαと㍑ぶUl、rNOM】,Showed the different reproducもive
activities,COmParedwiththos80r nOrther−1SpeCies;anyindividualof theabove−mention¢d
sp¢CiescouIdbreedseveraltimesdurirlgthebreedingseasom・
Several papers have been published on the breeding and the growth of other barnacles
(ClヰISr)andPAでlミし1961;BAIINl三SandSTC)NFニ,1973;HINifS,1979)andrecentlytheliteratures
has be8n reViewed by ClそISI)and‡ヨOURGl汀(j986).AYuNG(1976)and Hurtu三Y(1973)
12
γoshiakiIwAlくIand HideshiI−‡A′】、ri、011l
attempted to rationalize th扇r daもa onlirehistories or月αgα花㍑ぶ抒ぼ0花αβandβαぬ花㍑ぶpαCよ−
ficus using MACAitrE−川三Ⅰそand WILSON・s concept and reached the samc conclusion that the
rapidgrowth,earlysexualmaturity,hig・hfecundityandgrcatdispersalabilitieswereadvan−
tageousforadaptivestrategiesoftheso‘opportunistic’or‘fugitivQ’(トSelecもed)species.
Fil、Sもmaturity andinitiaigrowもh areimportantitems of basicinformation ror under−
Standingthelifehistoryand reproductivebiology of barnacles.To rnakecarefulconclusions
OnthebiologlCalminimumsizeandthebreedingactivlもyinrelation to sizeor age,Observか
tionsonagreatdealofmaterials coveI・ing・もhewholesizerange and froquent measurements
Ofgrowlngindividuals weremade.
MateriaJs and methods
DuringMay−October1986specimensofβ.αJγ堺たか∼£eand且ゐ071dαたou∠atIzo−uraillthe
recessofMatoyaヱさay and those of且∠〝岬rOUgぶ㍑ぶat Matiya−uIta dueeast of thecampus of
MieUniv即Sityw8r8remOV¢drrom bamboopoles,Which‡ladbeensもuckintotheseabedinth¢
1ittoralzon¢byfarmersforculturlng¢dible red andgreen algae,During A咽tlSt−September,
且eわ比r花eαぶ,β.α批coぶとαと㍑ぶand且 re乙£cαgα£混ぶWere Obtain¢d from styrofoam floats and
Otherartificialstructuresunderthe rafts,Which werealso mooredin themarglmalpユots of
MatoyaBayandAgoBayrorculturingoysterand peal・loysもerrespectivdy.So,thebarnacle
POPulation on the bamboo poles and under the rafts w8reinfluencedbythedifferent hydro−
graphicconditions.
BarnaclesfromthQ¢ntiresizera咽eOfeachspecieswereselectedatrandomforprocesslng
inth¢laboratory.Eachbarnacleswasmeasur¢dalongth¢apicalandbasaldiametQrOfrostro−
Carinalaxis,and exa‡Ⅵined forthepresonce ofegg masses.
Sixrectangularpanelsoftransparentg・1assmeasurlng18×14cmwerepreparedforporiodト
Calinspectionofgrowthrate.Inorderforthelarvatoattachthemselveswell,eaChpanelwas
processedbydrillinganumberofsma11pitsontoもhesurface,Thepaももern of settlemenもCan
bepredeterminedbythismethods,becausethecypridssettlepreferentiallyintohemisphericai
smalipits(ClえIS‡J,1960)Eachtwopanelsholdingablackpaneまbetweenthemselveswerofixed
togetherinto a wooden frame and therlthree sets of panels were suspendod at O.5,2and
4m below the surrace of the sea from a raft mooredin Matoya Bay.Tbe panels w8r¢
examined every three days from Au糾Sもto SQpもQmber andもhereaぎter at somQ mOrelo咽er
intervals.Ateachexamination theposition ofeach settled barnaclewastraced ontoanother
Clearplate,Placedonthebackofasurfaceofthepanelwithgrowlngbarnacles,alladditionai
foulinggrowthwasremovedand therlaPICaland basaldjametcrof rostro−Carinalaxis were
measured underabinocularmicroscopewiもhocularmicrometer.
First maもuriもy andinitまalgrowth of barnacles
13
ResuLts
l)R叩rOductivecondition.
Instudyi咽theareaonthePacificcoastofcentralJapan,ithasbe紺foundthatsucces−
sionalbTOOdsapparentlyoccurduringApril−DecemberinB.improuisus,鼠amphitriteandB・
ebumeusandduringMay−OctoberinB.hondahoui,B.reticulaiusandB.albicostatus・They
show no distincts組SOnalcyclo with synchronous release of young as observedin northern
species(BA】モNES and BARNES,1954;CR‡汎1954;BARN略1959)・Å1lof the samples
examined here which were collected during May−October were thereforein reproductive
condition.
︵辞︶ト0⊂¢⊃bむ︼軋
l00
80
dO
AO
20
0
O 1 2 3 4 5 d 7 8 9 】0 1l
O 1 2 3 d 5 る 7 8 9 】O1112】314 15
Api¢a暮 diameter a10ng th◎rO$trO−Car【nalaxIs(mm)
Fig.1.Percentage occurence of barnacles bearing egg masses・
In Fig.1the percentages of animals with egg massesまn the samples are shown as a
flユnCtion oftheapユCaldiameもer.The size ra咽eS and frequQnCies pertainlng tO the smallest
individualscarrylnぎeggmaSSeSVaried withspecies−Precisedimensjons ofthesmallest ones
w8re2.15mmforβ.£汀岬rOUZぶ㍑ざ(Nニ474),2.54mmforβ.α翫coぶまα£㍑ざ(N=509),3.56mmfor且re己Z−
c㍑∼α£混ざ(N=510),3.65mmfor鼠α?叩ゐかよ£g(N=842),4.85mmforβ.ゐ0花血如u∠(Nニ999)and
5.08mmforB.ebunLeuS(N=ニ660).Inan adult,eSPeCially southernspeciesofbarnaclewhichis
14
ToshiakiI\VAKiand HideshiHArユ「′ユ、ORr
PrOducing broods rapidly and repeatedly durir唱the breedingseason,thefertilization occurs
immediately afterma七uration ofgonads(CRIS盲〕and DAVほS,柑55).Ifitdid soinanimals
Which had jus七㌻組Ched maturity for thefirst timein theirlife,thevaluesglVen above for
each species can be regarded as the biologユCalminirnum size.Thereafter the proportions
Ofanimalsbearingfertilizedeggsincreasedwithincreaseinsize,andonreachingcertainsize
becamealmostasteadystateindistinguishablefromthoseoflargerindividuals.Thesizeat
Which theぎrequencies ceasQd toincrea3e COuld be rouが11y decided as rbllows;4.Omm for
β.わ研rOぴ∼ぶ混ざ,5.Ommfor且α路お0β£α己揖,5.5mmfor及re£去c昆ぬと㍑ぶ,6.Ommforβ.α〝軍兢rよ紘
6.5mmforβ.eあ比r花e㍑ざand8.Ommfor鼠如71血ゐou£.Thes¢Valu8Slikewise suggested to be
Parallelwith maximum sizes of barnacles observedin naturalpopulations,aS Shown by
marginalsizesrangosinFig.1.
2)Growth
BeforodrawlnggrOWthcurv8StberelationsbetweenapICalandbasaldiameterwerefirst
estabiished.Fig.2is based on a number of measurements of animals which were able to
expandthebaserreely underexperimentalconditions.Therelationshipfoundtogivethebest
COrrelation was two allometric8quations that were obtained separateiy,for thc small8r
β●∫
ラ/
0.5
1
2 3 4 5
0.5
1
2 3 4 5
0.5
1
2 3
Apicaldiameter(m†河
Fig.2.Relations between apまcaland basaldiameters measured along the
rostro−Carinalaxes or shell.
First maturity and initial growlh of barnacles
15
Tablel.Regressionformulashowingtheallometricgrowthofapicaldiameter(Ⅹinmm)
relative to basaldiameもer(Yin mm)or barnacles.
Species
Smaller group
Larger groups
鼠∠mprou£ぶ比ぶ
Ln Y=0.692+1.362×Ln X
Ln Y=0.770+0.948×Ln X
鼠 αgむ£coぶ£α£昆ぶ
Ln Y=0.718十1.3ユ8×Ln X
Ln Y=0.827」−0.870×Ln X
且αナナ叩仙「涼
Ln Y=0.744+1.538×Ln X
Ln Y=0−698+1.298×Ln X
Ln Y=0.699」−1.035×Ln X
Ln Y=0.743+0.964XLn Y
風 eわ捉㍗花e比ぶ
Ln Y=0.728+1.384×Ln X
Ln Y=0.833+1.016×Ln X
β.ゐ0花dαゐ0乙l∼
Ln Y=0.636+1.248×Ln X
鼠 re££c紀∠α餌S
groupslessthan2mmandforlargergroupsexceedinglmminapICaldiameter・Theintersections
ofthetwo r曙reSSionlines foreachspecies werenotlar酢1ydまfferent rrom¢aCh other,もhe
co−Ordinates showing the values ofl.14tol.33mmin apicaldiameter and2・40to3・07mm
inbasaldiametQr,QXCQptthat及ref加Jαと昆8hadrathersmallersizedpointand及reとょc㍑gαと昆ぶ
onlygaveasl咽1elin¢aStheresultofinsufficientdat臥Abovetheseresults朋reSummarized
in Tablel.Whereasin any smaller groups the relative growth coefficients,Or eXPOnentS,
werelargQrthanon¢,thosevaluesinlargergroupswereequalto,OrSOmeWhatsmallerthan
one.AnimalscomposlngSmallergroupsthereforehadtheirshellbaseswhichhadachievedat
higherspecificgrowthrate(dy/dt/y)thanthatrate(dx/dt/Ⅹ)oftheorificeofshell,but
inlargergroups both thesecompartments of shells dev8lopedisometrically(dy/dt/y=
dx/dt/Ⅹ).Thisfactcan beunderstood fromtheallometricequationinitsdifferentialform一
In any sp∝ies ofbarnaclesもh¢heavy settユem孤tis observed fromJune to Octoberin
Matoya Bay(ⅠⅥ∫ÅKI,H王別NO and KAWA】【IARノ\,1977).且reと∠c“gαと昆ぶand及£汀軍rOUよぶ㍑ざ
mainlysettledonpanelsatd叩thof2and4mrespectivelyandotherfourspeciespreferred
tosettleonuppermostone.Fig.3givesthegrowthcurvesofbarnaclesbasedonaveragesize
offivetotwelveまndまvidualsforeachsp∝iesat8VQrymeaSurement,butthem組Sur¢men七SOf
鼠ゐoJldαたou£and風花乙£c比gα£昆ぶWerelimitod to a錆w number orindividuals as wellas to
shortperiodsowlngtO alargenumberofmortalities・Itseemedthattherewasashortlag
phaseveryearlyinanygrowthcourse,Showingthegrowthrateofca・0−1mm/dayinthefirst6
days.From6to18daysthegrowthofanyspeciesaccelerated,barnaclesreachingale咽もhof
3.5to5.Ommattheendofthisperiod.でhereaどteritsloweddown,eXCePtthat放め㍑r花eむぶCOn−
tinuedtogrowin thesamemannoTaSbeforeFor30daysormore・Thegrowth rates ofB・
improuisus,B・albicosta己usandB・amPhitriteduringtheintervalfrom15to18dayswereO・28
mm/day蓼0.44mm/dayandO.55mm/dayrespcctivelyandduringthenextintervaltheratesreduced
byhal壬l・CoQfficiQntSOfvariationassociatedwithaveragesizesofbasaldiameterwere13to
18%atthefirstand2】to40%attheendoぎgrowthcourseof飽Chspecies.
Fig.3canbeusedinconjunctionwith bothIPig.1andダig・2givenabovetodetermine
approximatesizeandageatwi−ichthebarnacleぎirstreachesmaturityandthosecorr¢SpOnding
withtheintersectionoftworegrQSSionlinescorrelatingapICaldiameterswithbasaldiameもersL
でoshiakiIwAK王 and HideshiHATTORI
16
8
6
︵∈∈︶㌫芯∈万石
4
嵩め吋皿
0 3 6 9 12 15 18 21 24 27 30
Days after settlement
Fig.3.GrowもhofbarnaclesallowedもOSettleong・1asspanelsinMaもOyaBayfrom
midA咽uStもOmidSepもember.
i:及∼mダrOぴgS比ぶ,e:及eあ㍑r几e㍑ぶ,al:風α路£co8£α払5,am:β.αmp九たr£紬,
k:鼠如花血鬼0ぴ£,r:β.reと∼c捉gαと㍑∂.
Table 2・Sizes and ages corresponding・tOinflection point of allometric growth of
もhe shelland onset of first maturity.
First Maturity
王nflecもion poinも
Species
Apicaldia. Basaldia.
m
m
2
1
2
1
3
1
且 克or乙dαゐ0乙Jよ
2
0
β.eむu/・/!ビ〟S
5883朗4007一
1
β.αmpんか£とe
m
m
l
鼠 re乙∼e比gα£昆ぶ
2128911433⋮
且improL壷uぶ
β.α拍ね08fα£㍑8
Age
Apicaldia. Basaldia.
9days 2.15mm
lO−11
2.54
7
3.56
12−13
3.65
5.08
10
4.85
ユ0
Ag・e
4.46mm
5.15
17days
17
7.48
7.32
22
11.99
9.98
ca.50
Ther8SultsaregiveninTable2.It can be seen that the changes c)f positive allometry to
isometryin the relative growth of barnacle shells occur first of a11before first maturity,
andthatasr曙ards及∠mproび血ぶ,及α路gcoぶ£αと揖and且αmダ兢rgとetheageandsize orthe
Smal】eStbarnaclesbearingeggsnearlycoincidewiththoseatwhichthegrowthfelloff(viz.
3ふ5.Omminbasaldiameter;18days aftersettlement),but thereis aslight discrepancy of
Sizcs onlyin B.amphllrite.
First maturlもy andinitialgrowもh or barnacles
17
Discussion
A11individualbarnaclesexaminedhere werefrom the naturalpopulation undercrowded
COnditions,SO they would be capable of cross−fertilizationまn naturalconditions.Barnacles
Separatedfromothers byadistancegreaterthanthatoftheextendedpenisprobablydo not
rertilize(CRiSP,1950).This f飽tureis quite generalin cirripeds,eXC叩tin the case of
Cん£たαmαg昆ぶぶ£eggα£昆ぶ,掩rrαCαぶか℃e′花王αandβαぬ花昆叩eげorα乙昆ぶ(BA‡ヱNESandCRiS】〕,1956),
Inhisstudyorbreedinginβαgα花㍑5わαJα花0£desMool11ミ(1935)suggestedもhatthesterility
WaS foundin thQla柑er Or Olderindividuals,becomlng mOre frequent withincreasl昭Si狗
and was commonin members ofcrowded communities.As regards species studied here,the
barnacles more than two years old wererarQly foundin their habitats wher¢Sampling was
made,becauseintense settlement occurs armually over the greatly extended period andis
largelyresponsiblefordeathofanimalssettledinpreceding・year.ThemajOrityofspecim¢nS
brought ouもin】?ig.1arethereぎoreconsidered to be relativeけyoung and normaleondition
Withouts七倍rility.
Theimportant factors doterrninlng the size attained byindividuals are theintra−and
inter−SPeCificcompetition,the timeofsubmergence and the water movement carrying food
Withinrangeoftheanimals.Crowdingmayslowdown growth and delay the maもuration of
theovaryconsiderably(CRiSP,1961)andtheraftcondiもionsarefarbetterfromthepoinもOf
View ofgrowththan thosewhich areobtained onintertidalsubstrata(CRISP,1964).Using
沌eresultsrecordedherofortherateofgrowthundertheraftcondiもion,thesQWOuldth即efore
glVetheminimumagesoffirstmaturityfor及αJ?Wたか∠絶息如花血ゐou£and及∠J7箪rOUよぶ㍑S,
Whichweresampledfromcrowdedpopulationsonintertidalpiles,
By comparlngmany reSults of this and other studies for breeding of barnacles,it can
beseen that thereareconsiderabledirferencesin age and size of the first maturity of the
Same kind of barnacles from different r喝10nS.New generations of及eゐ昆㌻花e昆ぶin Trinidad
Mangrove Swamp carlreaCh maturitylnless than two weeks,during which the barnacles
growfromabout3to5mminbasaldiamQter(BACON,1971),COmparedwiththetwomonths
required at Woods Hole(GR^Vlラニ,1933)and the50days(12mm)in Matoya Bay.11iRANO
and OKUSHJ(1952)found thatinlate spri咽and summer鼠α汀pゐかよ£grel飽Sedits first
naupliiabout40days after settlementin Aburasubo Bay.In this area the temperature of
theseaduringthesameseasonwasabouも22to27℃,SOthebroodperiodofB.amphitriteis
COnSidered to be 4to 6days whichis estimated from the relationship between the brood
period and temp8ratur¢preViously published for the same varieもies,及 αJれpんかょとe var.
deTLticulata(PA{rH7Jand CRISP,1960)and B.amphitrite amphit7・ite(CRTSP and CosTLOW,
1963).Inthiscaseareasonableestimateofthe昭eforthefirstmaturityof及α77軍た£とr∠£eis
thereforeabout34to36days,andthesize at that timequoted from HmANO and OKUSHI’s
growthdatais8tolOm乱AIsoiもcanbefoundinsizeイrequencyhistogramsof及α′叩兢㌻如
inShimizuHarborgivenbyKosAKAandIsmBASm(1980)thatthesmallestanimalshavi咽
eggmassesarefrom3to3,5椚inapicaldiameter.Thesevaluoscorrespondwiththe6to7mm
18
ToshialくまIwAIくIand HideshiH▲へrl、′1、Oiモ】
basaldiameterobtainedbycalculation uslngtheequation shownin Tablel.1n Amsもerdam
afewindividualsofB,improuisusobserved to contain eggmassesare5mnlin basaldiameter
(Bl∧もー三汚Ml;N,1934).Itis approxjmately similar to the values obtained hereゴ予or t五e same
dimension,thoughthegrowth rateis slower than thatin Ma七Oya Bay.Comparisons of the
agesoffirst maturitywithin awide diversity of barnacles show that・thc values quoted for
SOuthern species are generallylower than thoseま−or northern species,SuCh as月.あα∼α柁Ogdeぶ,
鼠βOrCα己昆ぶandβ.九αmer∠that do not release naupliiuntilnearly the end o壬、their first
yoar(ClそISl⊃,1954;Moo兄,1935)The values of size aもrirsもmaturityパVeniflimited to
SOuthern species,Vary markedly with specics from about 2mm for BalaTluS trigonus and
C如九αmαg昆β/よぶぶ㍑ぶtO18mmbasaldiameter ror7甘けαCZgαぶq昆αmOSα(AYjJtNG,1976;H‡NJミS,
1979)WemaycorlCludethata1lofthebarnaclesstudiedherereachtheirmaturitiesatrelative−
1yearierag■eandsmall¢rSize,
Earlygrowthjustaftersettlementwasobservedtobeslowerinthepresentstudy.Similar
resultswereobtainedbyCRIS7〕(1960)forElminiusmodes乙us,Bala71uSCrenatuSandBalanus
balaTとOidesandheascribedthisfacttoreductionoftherateofflowintheimmediatevicinity
Ofthesolidsubstratum.Anotherpossibleexplanationofthcdclay wasthataftersettlement
endogenouscontrolofthegrowthwasforawhilelimitedtolowdegreeinconsequenceofthe
metamorphosis.でhe slowlng downin growth rate aft即maturi七yis obviousまy caused by
meもabolicchanges associated with reproduction.Themostcriticalphasefor survivaloccurs
duri咽Orjustaftermetamorphosis,Whiletheshellplatesarenotfullyformed,harde!led,and
cemented to thesubstratum(ClもIS】〕,1961).Tll¢faster expansion of the basis rather than
apertureofshellisth¢reforeadvantageousforsuchindividualsashaveg喝edlessthanlOdays
Or SO afもer settlement andin crowding condition smallspecimen mtlSt COmPIQte aS raSt aS
POSSibletofixitselffirmlytothesubstratum.
AcknowLedgmenl:
Theauthorswish to expresstheirgratitudeto Mr.T.Ul(A10fMieUniversity for kindly
helplr将1nSamPlir噂atthefields.
References
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β£ogogγ,36:335−342,
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relation to salまnity rluctuatまonsin a Trinまdad Mang−rOVe Swamp.ノ.eズp.mαr.β£oZ.
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0£ゐ0ぶ,5:63−76,
】ざir・St maもurlもy arldinitialgrowth og barnacles
19
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CRほ1〕,D.Jリ1950.Breedir唱anddistributionorCゐ£九αmα∼昆Sぶ紬ggα£比ぶ.Ⅳα£㍑re,柑6:31ト312.
…………‥・
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…………‥・
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