comparave ocular morphology of troglomorphic fish€¦ · • ocular anatomy (also, hot off the...
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Compara'veOcularMorphologyof
TroglomorphicFish:Implica'onsforEvolu'on?
ChuckSchobert,DVM
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Plan
• Non‐vertebratechordateeyes,agnatha
• Congoblindfish:Lamprologus lethops • Travelogue:CongoRiverHabitats• OcularAnatomy(also,HotOffthePress!)
• Mexicanblindcavefish:Astyanax mexicanus • Travelogue:Habitat• Modelsystem• OcularAnatomy• Sowhat?:Conclusions,specula'ons,andques'ons
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TeleostFishEyeAnatomy
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TheChordatesinEvolu'on
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Cladogram:Hagfish&Lampreys
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Comparison
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HagfishandLampreys
• Nopairedpectoral(shoulder)orpelvic(hip)fins• Notochordpersistsforlife.• Theyhavenoscales.• Theaxonsoftheirneuronsareunmyelinated.• Lampreyshavebothaninnateimmunesystemandanadap'veimmunesystem,butthelaVerisen'relydifferentfromthatfoundinthejawedvertebrates.
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Hagfish• Eatdeadwhales• Secretecopiousamountsofthickslime(mucous)
• Liveindeepoceanictrenches(>100meters)
• Recommenda'on:YouTube:
“EddieandtheHagfish”
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HagfishEyes
• Nocornea• Nolens• 2‐layeredre'na(neuronal)• Nomelanin• Primi'vephotoreceptors• Wiredtothebrainlikea
pinealgland• Nopinealgland• Photoreceptorssynapse
directlyontoganglioncells
Eye
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TransverseSec'ons
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Hagfish Specimen courtesy of Dr Ivan Schwab
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LampreyEye:Larval
• Larvalformtoadultform:FiveYears!
• Mostlyburiedinmud• Nolensini'ally
• Larvadetails– Similartohagfisheye– Eyecoveredinskin
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LampreyLarvalEye
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LampreyAdult•Cornealargelycon'nuouswiththeskin•Nomusclesofaccommoda'on•Hasmostofthestructuresofthevertebrateeye:
–Lens–3neuronallayersinre'na–Extraocularmuscles–Wiredtobrainlikeavisualeye–MelanininchoroidandRPE
Colorvisionandpinealgland
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LampreyAdultEye:FiveYearsLater
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PhotoreceptorEvolu'on
FromLamb,TDetal.Evolu?on of the Vertebrate eye: opsins, photoreceptors, re?na and eye cup.2007.NatureReviews:Neuroscience,8:960‐975.
Hagfish Lamprey
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TakeHome?
• Awindowtotheearliestdevelopmentofthevertebrateeye– Hagfish:circadianfunc'on– Lamprey:image‐formingeye,vertebrate‐like– Evolu'onarycontroversyaboutrela'onshipbetweenhagfishandlamprey:
– Ifhagfishdivergedearlierthanlampreysfromacommonancestor,opportunitytoseetheearliestexamplesofvertebrateocularevolu'on
– Ifhagfishdivergedfromthelampreyclade,thenoculardifferencesareperhapsdegenerate
• Andofcourse,weirdfood
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Hagfish‐SlimeCheddar‐GruyereScones
4cupsall‐purposeflour2tablespoonsbakingpowder4teaspoonssugar1/2teaspoonsalt1cup(twos'cks)chilledunsaltedbuVer,cutinto1/2‐inchcubes2cups(packed)coarselygratedextra‐sharpyellowcheddarcheese(about9ounces),oramixof6ouncescheddarand3ouncesgruyere.1‐1/2cupschilledheavywhippingcream6tablespoonshagfishslimePreheatovento375F
Inafoodprocessor,blendflour,bakingpowder,sugar,andsalt.CutinthebuVerusingquickpulsesun'lthemixtureresemblescoarsemeal.Addcheeseandcutinusingquickpulses.Inasmallbowl,whisktogetherthecreamandhagfishslime.Withthefoodprocessorrunning,addcreammixturethroughfeedtube.Processun'ldoughjustholdstogether–don’tovermix!
Turndoughoutontoalightlyflouredworksurface.Gatherthedoughtogetheranddivideintoquarters.Pateachquarterintoaroundjustshortof1inchhigh(itshouldbeabout6‐7inchesindiameter).Usingaclean,sharpknife,cuteachroundintosixwedges.Transferhalfthewedgestoungreasedbakingsheetslinedwithparchmentpaper,spacingthemabout2inchesapart.
Bakethefirstbatchofsconesun'ltheedgesjuststarttobrownandatoothpickcomesoutclean,about20minutes.Transferthem,s'llontheirparchmentpaper,toawireracktocoolatleast10minutes,duringwhich'meputinthesecondbatchofscones.
Servewarmoratroomtemperature.Thesconeswillstandforabout8hours.Donotrefrigerate.Ifyouwanttoreheatthem,warmthemina350Fovenforabout5minutes
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LampreyBordelaiseFor:6servingsTime:2h50Difficulty:medium
IngredientsforLampreyBordelaise:• 1live2kglamprey• 1dloil• 100gbuVer• 1boVleredBordeauxwine• 1smallglassArmagnac• 4shallots• 8onions• 15leeks• 4garliccloves• 600gBayonneham• 2tablespoonsflour• 1glassbroth• 2cloves• 1bouquetgarni• 6slicescountry‐stylebread• salt• Pepper
TechnicalstagesforLampreyBordelaise:
• Bleedthelampreybyhangingitbytheheadandcuqngthetailoveracontainertocollecttheblood.Whenthere'snoblooddrippinganymore,dipthelampreyinboilingwaterfor1minute.
• Takeitoutandpeelit.Cutin4cm‐thick(1,57inches)slices.Puttheslicesinthecontainerofblood.• Cutthewhitepartoftheleeksinto7‐8cm‐long(2,76‐3,15inches)whistlesandsteamtheminbuVer.• Addthedicedham,theshallotsandmincedonions.Sprinklewithflourandroast.• Moistenwiththewineandbroth.Addtheclovesandbouquetgarni.Addpepper,saltaliVle.Addtwocrushedgarliccloves.Bringtoaboilingpoint,thencookfor45minutesoverlow
heat.• 45minuteslater,addthepiecesoflampreyinthesauce,coverandcookagainfor45minutes.• Removetheslicesoffish,andflambtheminanotherpanorskilletwithArmagnac.• Pourtheequivalentofaglassofwarmsauceoverthebloodinordertodiluteit,andpoorintheskillet.S'rwell.Putbackthelamprey,seasonandcookfor10minuteswiththelidon.
BeatthesaucewithbuVer.• Meanwhile,toastthebreadandrubitwiththeremaininggarlic.Putthelampreyinashallowdish,ontheslicesofbread.Puttheleeksallaround.Coatwiththesauce.
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L. lethops andL. ?gripic?lis
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BenthicFish:gasbubbledisease
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“mondellibureau”
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CongoRiver
• Shortfilm
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WhatiscoolabouttheCongoRiver?
• Many,manymicrohabitats• Evolu'onoccurringallthe'me!
• Weirdriver=weirdfish
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SightedvsBlind
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L. Lethops histologyimages
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L. ?gripic?lis
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Immunohistochemistry:opsins
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Hotoffthepress!Mastacembelus brichardi
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Mastacembelus brichardi
• onespecimenisfullydepigmentedandeyeless• secondspecimenisaslightlypigmentedmorphwith'nyeyes:fullydevelopedlens,re'na,incompletecornea,eyesdorsallydisplacedtowardmidline
• intraspecificcomparisonoristheves'gialeyedmorphahybridbetweentheeyelessbrichardiandaneyedcongener
• collectedatthesamelocality,inshallowwaterunderrocksandboulders.
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Astyanax mexicanus
• Blindandsightedinsamespecies• Divergence1mya
• Modelsystemfordevelopment,moleculargene'cs
• Differencesinocularanatomy
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AMatureAdult
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Sighted
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Blind
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Cavehabitat
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Modelsysteminevodevo
• Moleculargene'cswellknown• Embryologicaldevelopmenteasilystudied:eyebeginstodevelop,lenscellsundergoapoptosis,degenera'on
• 4‐6monthgenera'on'me• Lenstransplantexperimentsshoweyedevelopmentgeness'llpresent:lensfromsightedintoblind,viceversa
• Pleiotropy:singlegenecontrolstwoormorephenotypictraitsthatmayseemunrelated:on‐offswitch….
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Ocularanatomy
• Comparisonmontage
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A. Mexicanus images:blind
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A. mexicanus:sighted
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Implica'onsforevolu'on?
• Evolu'onnotalwaysconstruc've• Regressiveevolu'onoccurs• Differenthabitatsproducevariedselec'onpressures
• Fishadaptstodarknessthroughselec'onof“numerousgrada'ons”leadingtoincreasedolfac'on,enhancedmechanorecep'on,alteredcraniofacialstructure
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PhenotypicPlas'city
• Phenotype=genotype+environment• Light‐darkexperimentswithepigeanAstyanax fasciatuslarvae
• Eyesoflarvaeraisedindarknessfor30daremuchlessdevelopedthanthoseraisedinlight
• Alternatedevelopmentalpathwaysbasedonenvironmentalcondi'ons……..
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Spin‐offs
• Why develop a morphological catalog of blind/par?ally‐sighted fish?
• Relatethesemorphologicaldifferencestogenomedifferencestoaidunderstandingofeyeevolu'on,developmentanddisease
• Usewell‐studiedsystemssuchasZebrafishandAstyanax spp tounderstandtherela'onshipbetweengene'csandmorphologywhenlookingatwildfish
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Manythanks!
• DrDickDubielzig• DrMelanieS'assnyandcolleagues,AMNH• DrWilliamR.Jeffery,UMD• DrIvanSchwab• DrLeandroTeixeira• DrJenDreyfus• MrScoVEarnest• MsKateLieber• Ms.JaniceLokken
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Ques'ons,specula'ons?