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Astheirnameimplies,membersoftheclassCephalopodahavemodifiedheadfoot
whichbearsanarrayofprehensiletentaclesandarmsatthecranialendofthebody.The
visceralmassislocatedtowardthecaudalend.Onlythenautiluspossessesanexternal
shell;theshelliscompletelylackinginoctopodsandisreducedandinternalinsquids
and cuttlefish. While squids and cuttlefish typically use their muscularized fins for
leisurelylocomotion,theyalsopossesstheabilitytomaneuverquicklybyjetpropulsion
rapidlyexpellingwaterfromtheirmantlecavitythroughtheirtubular siphon. The
evolutionaryrecruitmentofthemantlecavityasthefluidreservoirforjetpropulsionwas
incompatiblewithahardexternalshellandexplainswhythisfeatureisabsentorhighly
reducedinmostcephalopods.Theconsequenceofthisevolutionarytrendwasincreased
vulnerabilitytopredationduetothelackofprotectionaffordedbyashell.Asaresult,
many octopods and squids developed the ability to rapidly change their skin color
throughtheuseofspecializedepidermalcellsallowingthemtoblendinperfectlywith
theirsurroundings.This,inturn,madethemmoreeffectivepredators.Inaddition,most
cephalopods(exceptthenautilus)havean inksac whichcandischargeadark,cloudy
liquidthroughtheanustoconfusepotentialpredators.
1. Obtainapreservedsquidandpositionitinyourdissectingpansothatthesidewith
thesiphonistowardyou.
2. Examinetheexternalanatomyofthesquidandidentifythefollowingstructures:
tentacles,arms,fins,siphon,mantle,eyesandcollar.
3. Using your scissors, make a shallow, longitudinal incision through the mantle
startingatthecollarandextendingtothecaudalendofthebodytube(nearthe
fins).Youmayneedtousepinstokeepthebodytubeopen.
4. Use the illustrations and Table 11.2 to identify the following selected internal
structures of the squid and familiarize yourself with their functions: gills,
esophagus, stomach, pancreas, liver, anus, testis (male only), ovary (female
only), ink sac, siphon retractor muscles, kidney, systemic heart, branchial
hearts,caudal(posterior)venacava,andcranial(anterior)venacava.
Cephalopodshavethemostefficientcirculatorysystemofallmolluscs,supportingthe
relativelyhighmetabolicneedsassociatedwithswiftmoving,activepredators.Theyare
uniqueamongmolluscsinhavinga closedcirculatorysystem inwhichthebloodis
permanentlycontainedwithinanetworkofarteriesandveins.Thisadvancementallows
the blood to contain an unusually high concentration of oxygenbinding respiratory
pigments.Inadditiontoasinglesystemicheart,whichreceivesoxygenatedbloodfrom
the gills and sends it back to the tissues, squids have a pair of branchial hearts
associated with the gills to increase blood pressure and thus blood supply to the
capillariesofthegills.
Perhapsthesinglemostremarkableadaptationthatsquidspossessistheirwelldeveloped
eye.Thestrikingsimilaritybetweencephalopodeyesandtheeyesofvertebrates(birds,
mammals,fishes,etc.)isoneofthemostbeautifulexamplesof convergentevolution
amonganimals;eachgrouphasindependentlyevolvedacute,imageformingeyesthat
areamazinglysimilarinstructure. Theeyeofthesquidcontainsalens,cornea,iris,
ciliarymusclesandaretina,justliketheeyesofvertebrates.Duetotheirindependent
evolution,thecephalopodeyedoesdifferinafewimportantwaysfromthevertebrate
eye.Invertebrateeyesthelensiselasticandvisualimagesarefocusedontheretinaby
alteringtheshapeofthelens.Incephalopodeyesthelensisrigidandimagesarefocused
ontheretinabyalteringtheshapeofthelens.Incephalopodeyesthelensisrigidand
imagesarefocusedontheretinabyalteringtheshapeofthelens.Incephalopodeyesthe
lensisrigidandimagesarefocusedontheretinabyalteringthedistancebetweenthelens
andretina(justlikeinacamera).
Anothermajordifferencebetweenthetwoeyesisthewaythelightisreceivedbythe
photoreceptorsintheretina.Inthevertebrateeye,therodsandconespointtowardthe
backoftheeye(awayfromthepupil)solightmustpassthroughthephotoreceptorsand
otherassociatednervecellsandbounceoffthebackoftheretina(backtowardthepupil)
beforeitisdetectedbythephotoreceptors!Anunfortunateconsequenceofthisdesignis
thatalloftheneuronsarenaturallyontheinsideoftheretinaandwheretheyexittheeye
astheopticnervetheycometogetherinalarge,cablelikenervefiberandpushthe
rodsandconesasidetomakeapaththroughthebackoftheeyecreatingablindspotin
ourvisualfield.
Incontrast,thecephalopodeyehasthelightsensitiveendofitsphotoreceptorsoriented
towardthefrontoftheeye,solightenteringthepupilpassesthroughthelensanddirectly
stimulates the photoreceptors. Due to the arrangement of neurons being positioned
behindtheretina,cephalopodshavenoblindspotintheirvisual.Ifyouthinkaboutit,
thisisamuchmorelogicaldesignforaneyethanthearchitectureofthevertebrateeye!
Thisisagoodillustrationofhowdifferentevolutionarymeansmaybeemployedtoreach
the same endin this case a functional, imageforming eye. Remember, natural
selectioncanonlyoperateonexistingvariationwithinpopulations,sothemostlogical
designmaynotalwaysbeachievable. Withoutnaturalvariabilityinatrait(e.g.,the
elasticityofthelens),thattraitwillforeverremainunchangedandothertraitsinwhich
variabilitywithinthepopulationdoesexistaremodifiedtoimprovesurvival(e.g.,ciliary
musclesthatmovethelensbackandforthratherthanchangetheshapeofthelens).
Table11.2AnatomyofaCephalopod(Squid)
Structure Function
Collar Fleshyborderseparatingheadfootfromvisceralmass(mantle)
Eyes Imageformingorgansfordetectingvisualstimuli
Siphon Hollowtubethroughwhichwaterisexpelledfromthemantlecavityathigh
velocitytopropelthesquidthroughthewater
Mantle Bodytubeencirclingvisceralmassformingahollowchamberinwhichwateris
collectedandusedforpropulsion
Arms Shorterappendages(8)usedtomanipulatecapturedpreyandactasarudderfor
navigatingwhileswimming
Tentacles Long,extensible,prehensileappendages(2)forcapturingprey
Fins Triangularshapedextensionsofthecaudalendofthebodytubethatareusedfor
leisurelyswimmingandformaneuveringduringlocomotion
Gills Featheryorgansusedforrespiration
Esophagus Thintubeconnectingthemouthtothestomach
Stomach Smallsaclocatedatcaudalendofthebodytubewherefoodisstoredand
digested;digestionisentirelyextracellularincephalopods
Pancreas Small,granulardigestiveglandthatsecretesenzymesintothestomachtoassistin
thebreakdownoffood
Liver Large,elongatedglandthatreleasessecretionsintothestomachtofacilitate
enzymaticdigestionoffood
Anus Terminalportionofdigestivetractlocatednearsiphon
Testis Producessperm;locatedincaudalendofbodytube
Inksac Largesacthatopensintotheanusandsecretesadarkbrownorblackfluidwhen
theanimalisalarmed
Siphonretractormuscles Longmuscleswhichcontrolthecontractionofthesiphon
Kidneys Adjacentexcretoryorganslocatedbetweenthegills
Systemicheart Large,muscularizedchamberthatreceivesoxygenatedbloodfromthegillsand
pumpsitthroughoutthebody
Branchialhearts Smaller,muscularizedchambersthatreceivedeoxygenatedbloodfromallparts
ofthebodyandpumpbloodtothegills
Caudavenacava Drainsdeoxygenatedbloodfromthebodytubeandmantlebacktothebranchial
hearts
Cranialvenacava Drainsdeoxygenatedbloodfromtheheadfootbacktothebranchialhearts