36TH FUNCTIO O TH GILL O MAYFLNYMPH FRO DIFFEREN HABITATB C A WINGFIEL - PDF document

36TH FUNCTIO O TH GILL O MAYFLNYMPH FRO DIFFEREN HABITATB C A WINGFIELD1Fro th Zoolog Department Universit o Birmingha(Received 3 April 1939(Wit Thre Text-figuresI INTRODUCTIOTH majorit o mayfl nymph posses evagination o th bod surfac i thabdomina regio whic ar mor o les richl supplie wit trachea an t whicth ter trachea gill ha bee applied Palme (1877 wa o th opinio tha thesstructure wer bloo gill i whic th trachea serve chiefl fo mechanicasuppor an playe n direc par i respiration Vayssier (1882) howeverassigne a definit respirator functio t th tracheae H als pointe ou thai earl instar i whic th trachea gill ar no developed respirator exchangtake plac throug th thi bod wall Dewit (1890 observe tha mayfl nymphfro whic th trachea gill ha bee remove no onl survived bu afte a femonth regenerate thei gills Cue"no (1925) i hi wor o th mayfl nympCloeon dipterum, confirme thi observatio o Dewitz an conclude tha "letrache'o-branchies adjuvant d l respiratio cutand ge"ne"rale peut-etr inutilee hiver e repos dan l'ea extr&memen oxyg6n£e deviennen utile e indispensable lorsqu'un activit plu grand d corp necessit un oxydatio pluactive lorsqu l'ea es moin oxyg^n^e comm e 6t6". Fro experiment similat thos o Cue"not Rem (1925 conclude tha oxyge absorptio too placchiefl throug th trachea gill an tha Cu6no ha underestimate th importanc o thes structure i respiration Baba & Foustk (1907 foun tha ath oxyge concentratio o th wate wa decrease th gill o mayfl nymph mor rapidly an the conclude tha th gill wer specia respiratororgans Workin o mayfl nymph fro differen habitats Dodd & Hisa(1924 demonstrate a definit relatio betwee th gil are an th oxygeconten o th water Pruth (1927) o th basi o hi finding fo a singlspecies Cloeon dipterum, cam t th genera conclusio tha "mayfl larvae anpresumabl othe insect a well ca stan a ver lo concentratio o oxygenbelo roc.c/1." Morga & Grierson 19 2 observe tha i th burr win mayflnymph Hexagenia recurvata, th gill wer essentia t norma life althoug gill-lesnymph wer foun t surviv unde certai conditions Eastha (1937 suggestetha th chie functio o th trachea gill i Ecdyonurus venosus wa t bath witwate th respirator gil tuft attache t thei bases1 Kedde Fletche War Studen o th Universit o LondonJEB-XV ii 2 36 C.'A WlNGFIELApar fro th wor o Morga & Grierson i appear tha n accurat experimenta wor ha bee don o th precis respirator functio o th tracheagill i mayfl nymphs althoug man suggestion a t th probabl natur othi functio hav bee pu forward Furthermore nothin i know o an relatiobetwee functio an habitatI appeare desirabl therefor t investigat th respirator functio o thtrachea gill o a numbe o specie o mayfl nymph fro differen habitatsII MATERIAL AN METHODTh animal use i thi investigatio wer Cloeon dipterwn (L.) Baetis sp.an Ephemera vulgata (L.) Cloeon dipterwn i a typica still-wate form whereaBaetis sp i characteristi o swif streams Ephemera vulgata i a burrowinmayfl nymp foun i stil watersCloeon dipterum wa obtaine fro thre differen localities namely a pona Alvechurch Worcestershire a pon a Sell Park Birmingham an a pon aNewdigate Surrey Baetis sp wa take bot fro a swif strea a BlakedownWorcestershire an fro a simila strea a Alvechurch Worcestershire Ephemeravulgata wa obtaine fro th pon a Newdigate Surrey Th oxyge consumptio o specimen o Cloeon dipterum fro th differen localitie mentione abovwa foun t b th same thi wa als th cas wit Baetis spTh alkalin reserve o th water fro whic th animal wer take wera follows Alvechurc pond 0-004 -W Sell Par pond 0-002 N; Newdigatpond 0-001 N' Blakedow stream 0-003 N; Alvechurc stream 0-004 N.Al experiment wer don i a thermosta a io C Oxyge consumptionwer measure o th da followin tha o whic th animal arrive i thlaboratory Oxyge concentration wer determine b th metho o Fo &Wingfiel (1938) I al experiment o Baetis sp an Ephemera vulgata, th experimenta techniqu use wa essentiall th sam a tha describe i a previoupublicatio (Fo et al. 1937) tha techniqu i referre t i thi pape a th"normal technique Th first tw serie o experiment o Cloeon dipterum werals don i thi way bu i th secon tw serie a differen experimenta techniqu employe whic involve certai structura addition t th thermostat Aframewor consistin o on horizonta an tw vertica arm wa fixed t thinsid bas o th thermostat A axl carryin a larg whee wa mounte o thvertica arm i suc a positio tha whe th thermosta wa filled wit wate whee wa completel immersed Th wheel whic wa perforate b si largholes wa connecte throug a syste o pulley t a electri moto outsid ththermostat Afte th animal ha bee place i th experimenta bottle containin o know oxyge concentration a smal glas ro (0- cm diameter 5 cmlong wa introduce an th stoppe replaced Th bottle wer the insertehorizontall i th hole o th whee o th apparatu describe abov an heli positio b smal piece o cork I thi wa th experimenta bottle coul b Function of the Gills of Mayfly Nymphs from Different Habitats 36rotate i th thermosta throughou th cours o a experiment Th consequencirculatio o th glas ro insid eac bottl serve t se u a artificia watecurrent Thi technique whic i terme th "stirring techniqu i thi paperensure tha a constan curren o wate wa passe ove th bod surfac o thanimal throughou th experimentI eac o th mayfl specie mentione above th oxyge consumptio onorma nymph a variou oxyge concentration wa compare wit tha onymph fro whic th trachea gill ha bee removed Th remova o th gill effecte a follows Animal wer anaesthetize i 0- % urethane Singlanimal wer the transferre t a hollo glas bloc containin dilute anaestheti an th gill remove wit iridectom scissor an fin forcep undeth lo powe o a binocula microscope Th animal wer the place i fullaerate wate an lef t recover recover wa complet withi 3 min Th oxygeconsumptio o gill-les animal wa measure o th sam da a th gill werremove an abou 3- hr afte th operationA fa a i wa possibl t judge th activit o norma an gill-les animalwa th sam i al experiment excep thos wit Ephemera vulgata. I thispecie th gill-les nymph wer noticeabl les activ tha th norma onesIII EXPERIMENTATh dat fo norma an gill-les Baetis sp ar show i Table I an IIan summarize i Fig 1 I wil b see fro th figure tha ove a rang o concentration fro 5- t 8- c.c./l. th oxyge consumptio o normaan gill-les animal i alike N experiment wer don a oxyge concentrationlowe tha 4- c.c./l. a i wa foun tha nymph becam moribun an ultimateldie i subjecte fo an lengt o tim t oxyge concentration lowe tha thi(cf Pruthi 1927) I i clea tha ove th rang o oxyge concentration testeth gill o Baetis sp pla n par i oxyge uptakeTable II an I giv th oxyge consumptio o norma an gill-les Cloeondipterum a variou oxyge concentration usin th norma technique Th dat arsummarize i Fig 2.1 I wil b see tha th oxyge consumptio o norma angill-les animal remain th sam ove a rang o oxyge concentration fro8- t 3* c.c./l Belo th latte concentratio th oxyge consumptio o gill-lesnymph fall of rapidly bu thi marke decreas doe no occu i th normaanimal unti th oxyge conten o th wate i reduce t approximatel 1- c.c./lI i clea tha th gill i thi specie onl ai oxyge consumptio i wate ooxyge concentratio lowe tha 3- c.c./lI Baetis sp th gill neve beat I Cloeon dipterum th gill bea intermittentla hig oxyge concentrations bu a th oxyge conten o th wate falls thquiescen period becom muc reduced an thu th volum o wate passeove th dorsa surfac o th nymp i a give tim i correspondingl increasedTh norma leve o oxyge consumptio o Cloeon dipterum a lo oxyge1 Thes result hav alread bee communicate i a preliminar repor (Wingfield 1937)34- Tabl I The oxygen consumption of Baeti sp. at various oxygen concentrations.Normal animals: normal techniqueExpnoi234567891Ii232245611291I11Dat4 v 3ititit12 v 3it18 v 3ititit25 v 3»5 v 3it2 vi 3„,,,28 v 3it9 vi 3,,,„Mea drweigh o1 animalM. \0 IIIH Til *mg11111111111111111I11111111Oxyge concentratioc.c/1I exp7-7-777-7-7-7-7-7-7-7-6-6-5'S-5-S-S-5-5-4-44-4-Mea7-7-6-S-4Oxyge consumptioommI exp268234256273199204197231236205212284213224287239154172229156158195263159152202154232/g-/hrMea258221263190180Tabl II The oxygen consumption of Baeti sp. at various oxygen concentrations.Gill-less animals: normal techniqueExpno1234I0911111122211I23Dat3 " 3�11 ii 3.14 iv 3,„16 iv 3•� 21 iv 3,�5 v 3it�t28 iv 3»15 vi 3itMea drweigh o1 animalmg111912211I1I1I111I919Oxyge concentratioc.c./lI exp777rlTS7TS7'666-Mea7-7-6-6-6-6-5- 5-5-S'5-4-5-Oxyge consumptioc.mm./g./hrI exp204224190255226204184234232252260274273221256250237215iS7139178Mea218222255226158 Function of the Gills of Mayfly Nymphs from Different Habitats 363 4 5 6 7 8 9Oxyge concentration c.c./lFig i Th oxyge consumptio o norma an gill-les Baetis spa variou oxyge concentrationsI1 3 4 5 6 7 8 9Oxyge concentration Fig a Th oxyge consumptio o norma an gill-let Cloeon dipteruma variou oxyge concentrations Norma technique• Norma animals O Gill-les animals. 36 C A WlNGFIELTabl III The oxygen consumption of Cloeo dipteru at various oxygenconcentrations. Normal animals: normal techniqueExpno1234567891111I1111222222222333Dat16 xi 3i�»,»2 xii 3�3 xii 3I„»t»20 xi 3»,�21 xi 3l25 xi 3)II28 xi 3»,I26 xi 328 xi 3»Mea drweigh o1 animalmg22322232333333433334333333333733Oxyge concentratioc.c./lI exp777-7-7-7-766-6'66-66-44'44'112-I2-2-2-i-1-••1 "i-i'0i-Mea7'6-4-2-i*I'Oxyge consumptioc.mmI exp98147148III129141139137155157I53I351271261251289311599109102123121m13310014310077716075/g-/hrMea129141115m11971concentration migh b maintaine b eithe o th followin processes Th gillmigh functio a tru respirator organs actua gaseou exchang takin plac ath surfac o th gills or alternatively th gill migh ac a paddle passin agreate volum o wate ove th norma respirator surface o th animal I havteste th possibilit o th secon alternativ b measurin th oxyge consumptio a variou oxyge concentration o norma an gill-les animal i aartificia wate curren produce b rotatin th experimenta bottle i th stirrinapparatu mentione above Th result ar give i Table V an VI an summarize i Fig 3 I shoul b note tha th oxyge consumptio o norma angill-les animal a relativel hig concentration o oxyge wa approximatelth sam usin eithe techniqu (cf Figs 2 an 3)I wil b see fro Fig 3 that usin th stirrin technique th oxygeconsumptio o norma an gill-les nymph remain th sam ove th whol rango oxyge concentration studied I i clea the tha th rapi decreas i oxygeconsumptio o th gill-les nymph a oxyge concentration lowe tha 3. c.c./l Function of the Gills of Mayfly Nymphs from Different Habitats 36Tabl IV The oxygen consumption of Cloeo dipteru at various oxygenconcentrations. Gill-less animals: normal techniqueExpnoi23456I91I111I111122Dat10 xii 311 xii 312 xii 39 i 3II7 i 3,8 i 3I16 i 3I29 i 3IIIS i 3,28 i 3IIMea drweigh o1 animalmg422333231333233l22133223Oxyge concentratioc.c./lI exp8888-7-4443-3-2-2-•[•[•'•[••1-Mea8-4-3-2-1-I'Oxyge consumptioc.mmI exp14315615514112815812015010714769599062622925202719/g./hrMea1471351326834£D0 I 2 3 5 6 7 6 9Oxyge concentration c.c./lFig 3 Th oxyge consumptio o norma an gill-les Cloeon dipteruma variou oxyge concentrations Stirrin techniqueNorma animals O Gill-les animals 37C A WlNGFIELTabl V The oxygen consumption of Cloeo dipteru at various oxygenconcentrations. Normal animals: stirring techniqueExpnoi23456I91I111111112Dat29 vi 3i„1 vii 3,5 vii 3,•12 vii 3„,9 vii 3,114 vii 3,,7 vii 3»Mea drweigh o1 animalmg11911111111191111111Oxyge concentratioc.c./lI exp7-7-7-7-7-7-7-7-7-S-5-S-33322'I-I-I-Mea7'5-3-2I-Oxyge consumptioc.mm./g./hrI exp209146119167148133125158123122131115121105707962Mea14612310810770Tabl VT The oxygen consumption of Cloeo dipteru at various oxygenconcentrations. Gill-less animals: stirring techniqueExpno34567891111111Dat16 vii 31»»19 vii 3,,M21 vii 3�,123 vii 3„Mea drweigh o1 animalmg1111911119199Oxyge concentratioc.c./lI exp7-7-7-s-5-5-2-2-2-20-i-i-Mea7s-2-I-Oxyge consumptioc.mm./g./hrI exp15214713214113112512712190115117776874Mea143128III73 Function of the Gills of Mayfly Nymphs from Different Habitats 37whic wa foun whe th "normal techniqu wa used i du t th absenco a wate curren ove th bod surfac o th animals Thu i Cloeon dipterum,th gill d no functio a tru respirator organ eve a lo oxyge concentrations and unde thes conditions th norma leve o oxyge uptak i maintaineb th increase movemen o th gill whic cause a large volum o wate tb passe ove th respirator surface o th animal A n indicatio wa founo th presenc o ope spiracle i thi species i appear probabl gaseouexchang take plac throug th integumentRem (1925 explaine th surviva o gill-les nymph b supposin tha thai store i th trachea wa sufficien fo norma respiratio t b continue fosom tim afte th remova o th gills Th norma rat o oxyge consumptioo gill-les Cloeon an Baetis a hig oxyge concentration migh b explaine ia simila way Suc a suppositio woul no howeve explai th differenc ioxyge consumptio betwee norma an gill-les Cloeon a lo oxyge concentration (norma technique and moreover th disappearanc o tha differencwhe th stirrin techniqu i usedTh oxyge consumptio o norma an gill-les Ephemera vulgata a abou thsam oxyge concentratio (7- an 6- c.c./l. i show i Table VI an VIIIFurthe experiment a lowe oxyge concentration wer no possibl a threquisit numbe o animal coul no b obtained I wil b see fro th table th oxyge consumptio o gill-les animal i abou one-quarte tha o normaanimals I thi specie i appear tha th gill ai respiratio eve a hig oxygeconcentrations Morga & Grierso (1932 obtaine a simila resul wit Hexagemarecurvata, a nea relativ o Ephemera vulgata.Cloeon dipterum, whic ha plate-lik trachea gill wel supplie wit tracheae ifoun i pond wher th oxyge conten fall ver lo (unpublishe data) Fig 2show tha a lo oxyge concentration (1-5—3- c.c./l. th norma leve o oxygeconsumptio canno b maintaine b th bod surfac alone Unde thes condi whic ar ofte experience i nature th motil trachea gill furnis aaccessor respirator mechanis whic enable norma respiratio t b continuedBaetis sp i foun i swif stream wher th oxyge conten rarel fall belo4 c.c./l (unpublishe data) usuall i i considerabl higher Th structur o th i simila t tha o Cloeon dipterum, excep tha wherea i th latte specieeac gil i compose o tw lamellae i Baetis sp i consist o onl a singllamella I wil b see fro Fig 1 tha th respiratio a th bod surfac i Baetissp i sufficien t maintai norma respiratio a al oxyge concentration whicwoul b experience i it natura habitat Unde natura condition thereforther i n nee fo an accessor respirator mechanis i thi species an thnon-motil gill d no functio i respirationTh burrowin mayfl nymp Ephemera vulgata i als foun i pond wherth oxyge conten ma dro ver low it motil gill ar filamentou an sparselprovide wit tracheae Table VI an VII sho tha i thi species althougsom respirator exchang appear t tak plac throug th integument th bodsurfac i insufficien t maintai norma respiratio eve i wate saturate wit 37C A WlNGFIELTabl VII The oxygen consumption of Ephemer vulgataNormal animals: normal techniqueExpno6789II11\l1Dat12 xii 3it14 » 3ftft20 ii 310 iv 3it"Mea drweigh oT animalmg171815IJ1018181719202023Oxyge concentratioc.c./lI exp7-7-77-6-66-6-6-6-6-Mea6Oxyge consumptioc.mm./g./hrI exp147145148149189121139117Mea146Tabl VIII The oxygen consumption of Ephemer vulgataGill-less animals: normal techniqueExpno2346IDat19 • 325 1 3Mea drweigh o1 nnimnlmg1814I713Oxyge concentratioc.c./lI exp7-7-67Mea7Oxyge consumptioc.mm./g./hrI exp4537453535Mea40air I appear tha th accessor respirator mechanis i th for o gill i verwel develope i thi species Thi ca b relate t th ver lo oxyge concentratio o th wate whic ma ofte b experience insid th burrow o thes i nature Th exac natur o th accessor respirator mechanis iEphemera vulgata i unknown bu i vie o th differenc i gil structurbetwee i an bot Cloeon an Baetis, i i possibl tha th gill ma ac a trurespirator organ a wel a paddle producin a respirator currentSUMMAR1 Th oxyge consumptio o norma an gill-les nymph o th mayflieBaetis sp. Cloeon dipterum an Ephemera vulgata ha bee measure a variouoxyge concentrations2 I ha bee foun tha ove th complet rang o oxyge concentrationstudied th trachea gill d no ai oxyge consumptio i Baetis sp I Cloeondipterum, a al oxyge concentration tested n gaseou exchang take placthroug th gills a lo oxyge concentrations however th gill functio a aaccessor respirator mechanis i ventilatin th respirator surfac o th bodan s ai oxyge consumption I Ephemera vulgata th gill ai oxyge consumptio Function of the Gills of Mayfly Nymphs from Different Habitats 37eve a hig oxyge concentrations I thi specie th gill ma functio bota tru respirator organ an a a ventilatin mechanism3 I i show tha th difference i gil functio ca b relate t th oxygeconten o th habita o eac speciesREFERENCEBABAK E & FOUSTKA O (1907) PflOg. Arch. get. Pkytiol. 119 530CUENOT L (192s) L'Adaptation. ParisDEWITZ H (1890) Zool. Anz. 13 535DODDS G & HISAW F (1924) Ecology, 5 262EASTHAM L (1937) J- exp. 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