cefalopodos cuerpo plano origenes

7/29/2019 Cefalopodos Cuerpo Plano Origenes

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Tanabe, K., Shigeta, Y., Sasaki, T. & Hiano, H. (eds.) 2010. Cephalopods - Present and PastTokai Univesity Pess, Tokyo, p. 23-34.

Introduction: Traditional views on the evolution of

cephalopod body plans

In 1830, two young natualists, Meyanx andLauencet, attempted a compaison of the anatomy ofvetebates and cephalopods, speculating that they havethe same basic stuctual pinciple. While GeoffoySt. Hilaie adopted the idea as poof of his theoy,on the unity of body plan that is composed of shaedcomponents of all animals, Geoges Cuvie ejected itusing questionable esults of his anatomical study ofan octopus (Figue 1; Appel, 1987; Le Guyade, 2004fo eviews). Eve since this pe-Dawinian academic

debate, many oologists have indulged in a long lastingdiscussion of how the cephalopod body plan and theiogan systems can be linked to those of vetebates (e.g.Packad, 1972; ODo and Webbe, 1986).

To addess the l inkage be tween these twophylogenetically distant taxa, data to infe the oiginalcondition was equied. The taditional views on theoigin of cephalopod soft pats lagely elied on theanatomical analysis of extant cephalopods, paticulaly

nautiloids (Owen, 1832; Macdonald, 1855; Giffin,1900; Willey, 1902). The numeous tentacles without

suckes, the leathey hood, the pinhole eyes, the ovoidsacs of statocysts, the cod-like bain, the unfusedfunnel, absence of the ink sac, and also most of theothe chaactes have been consideed to be pimitivestates that enabled us to infe the common ancesto (seeSaundes and Landman, 1987; Budelmann et al., 1997).

Though a synthesis of the data in paleontology,embyology, and adult mophology (Naef, 1921-23,1928), some evolutionay schemes wee constuctedto explain the tansitoy change (Figue 2) with theidea of a pototype of Conchifea and a hypothetical

Na ut il us embyo (Naef, 1928). In this view, the

anteio-posteio (AP) and doso-vental (DV) axesae completely conseved duing the changes foma gastopod to a cephalopod as initially suggestedthough the embyological study of the squid (Books,1880) (the foot o ams define the vental side of the

bodies, and the head defines the anteio side). Alongthe DV axis, the ams, colla, funnel, mantle, and theshell of a cephalopod ae espectively coespondingto the whole foot sole, epipodial egion, mantle, and

The origins of cephalopod body plans: A geometrical and

developmental basis for the evolution of vertebrate-like

organ systems

SHUICHI SHIGENO*1, SASAKI TAKENOrI2AND SIGUrD VON BOLETzKY3

1Department of Neurobiology, University of Chicago, 947 East 58th Street, Chicago, IL 60637, USA(*corresponding author; e-mail: [email protected])2The University Museum, University of Tokyo, Bunkyo-ku, Tokyo 113-0033, Japan (e-mail: [email protected])

3C.N.R.S., Observatoire Ocanologique, Laboratoire Arago, F-66651 Banyuls-sur-Mer, France (e-mail: [email protected])

received May 6, 2008; revised manuscipt accepted Septembe 28, 2008

Abstract. The evolution of cephalopod body plans has been one of the most intriguing topics in zoology. Their

body parts, particularly of coleoids (squids, cuttlefishes, and octopuses), are composed of multiple sets of

components including vertebrate-like analogical systems as a result of convergence. However, in spite of thepotential importance for understanding the evolution and diversity in bilaterians, the origins of cephalopods

have been poorly understood. There is little consensus of opinion for morphological linkage among the plans

of cephalopods, basal molluscs, and other bilaterians. Here, we provide a review and new interpretation

with an emphasis on the topographic transition of the soft parts that is shaped by a shared concentric circle

or ovoid pattern in the embryos and adults of extant or fossil molluscs. The purpose of this article is also to

characterize the cephalopod body plans, set against those of the other bilaterians, in the light of recent data

from paleontology, embryology, and molecular gene expression patterns.

Key words; embryo, evolution, development, molluscs, nervous system, nautilus


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Shuichi Shigeno et al.24

the shell of gastopods. This scheme supposes thatcephalopod ams wee developed fom the whole footegion of a gastopod, and that the colla was deived

fom the epipodial egion. Basically, concening the DVaxis, this view has long been adopted by subsequentauthos (raven, 1958; Seidel, 1960; Fiooni, 1978;Boletky, 2003, 2006; Shigeno et al., 2008). Bandeland Boletky (1988) suggested that cephalopod amswee deivates fom the oveall foot egion, the funnelalso a modification of the posteio foot (Boletky,1989, 1993). Howeve, some diffeent ideas alsohave been poposed about the concept on AP axis,fo example, that the pe-tocheal head egion ofthe spialian tochophoe lava coesponds to thelocation of the shell gland in cephalopods, given theapical position of the animal pole (Fiooni, 1978). Todate, we can use a lage amount of histological datafo undestanding development of each ogan systemin coleoids (see Fiooni, 1978; Budelmann et al.,1997).Howeve, when in the 20th centuy moleculaeseach took the foefont, the developmental study ofcephalopods fell out of favo, due to the lack of a modelspecies amenable to molecula biology. As a esult,unfotunately, the pevious evolutionay hypotheses as

stated above emain lagely untested.In tun, a diffeent appoach has been adopted by

some paleontologists and compaative malacologists(Figue 2; Yochelson et al., 1973; Holland, 1987;Salvini-Plawen and Steine, 1996; Lee et al., 2003).Thei scenaio was lagely based on the impotantndings of the fossils of monoplacophoran-like shells,and the hypothetical econstuction of the soft pats(Yochelson et al., 1973). In this view, the cephalopodtentacles/ams ae deivates of both the diffeentiatedcephalic egion and anteio pat of the foot. The

posteio foot tansfoms into the funnel fold. The funnelis situated at the posteio end of the body; theefoe,the posteio foot pat of benthic ancesto is equied toshift to moe posteio and dosal egions of the bodyof cephalopods. Unfotunately, no fossil ecod of soft

pats has been found so fa to infe the soft pats in thebasal cephalopods. Theefoe, no evidence exists to

suppot this hypothesis (Boletky, 2006). Howeve, it isnotable that the pesence of multiple paied muscle scason some fossil shells (Yochelson et al., 1973) suggeststhat the ancesto might have distinct pedal etactomuscles (o so called doso-vental musculatue) likeothe molluscan goups (Haspuna and Wanninge,2000).

Nautilus embryos:

their unique concentric patterns

How can we cove a lage gap between the complexcephalopod body plans and the much simple ones of

othe molluscs such as gastopods? The embyonicmophology is often a bette guide to highe-gadeevolutionary afnity than adult morphologies that varygeatly in thei shapes. The plans of ealy stages ofcoleoid embyos (e.g., squid, Figue 3D) still exhibitsubstantial diffeences, because of thei lage eyeegions, posteioly situated colla with the funnel,and the am buds ae aanged along the tansveselyelongated ovoid pattens of the embyonic bodies,impeding diect compaison with those of a gastopodtochophoe lava and adults.

An attempt to esolve this poblem came fom theobsevation of living Nautilus embyos (Anold andCason, 1986; Anold, 1987). The six vaiously stagedembyos ofNautilus belauensis wee successfullycollected and the extenal mophology was obseved.The embyos wee diect developes chaacteied bya vey lage oute yolk sac. The egg sie is the biggestof all invetebates, and nealy one yea is needed fothei development to hatching. The embyonic bodiesae moe elongate along the AP axis compaed to those

Figure 1. A classic example fo the compaison of inte-nal organs between a bird and an octopus. Cuvier used this gure

to suppot his functionalism opposed to Geoffoys mophology.The dosal side of octopus body coesponds to the vental sideof the vetebate, showing simila U-shaped digestive ogans.Howeve, the lage pat of the bain is located dosally in the ve-tebrate in contrast to the opposite side. The gure is from Cuvier

(1830).


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Oigins of cephalopod body plans 25

of coleoids. The cephalic compatment is situated at themost anteio, and the tentacle buds ae aanged alongeach lateal side of the embyonic bodies. Followingthese ndings, a more detailed analysis was conductedon the embyonic shell (Anold et al., 1987; Tanabe andUchiyama, 1997). recently, the unknown ealie stagesofNautilus pompilius wee analyed, and the outline ofdevelopmental sequence was finally descibed (Figue3A-C; Shigeno et al., 2008).

The ealy aangement ofNautilus embyonic bodiesobviously exhibits a shaed oveall patten with thatof adult gastopods and of coleoid embyos (Figue3D). The pimodia of body pats ae aanged in aconcentic cicle geomety; the cental (the shell fieldand mantle), medio-lateal (the epipodial egion ocolla with funnel), and the lateal most (the foot otentacle/am buds). Compaed to the pattens of anadult gastopod, Nautilus and coleoid embyo show;(1) a posteio enlagement of the head o cephaliccompatment, (2) a mophological distinction of the

colla and funnel compatment, and (3) the appeaanceof the segmental buds in the whole foot compatment.This conseved configuation of ogan systems seemsto constuct a topogaphical basis to explain howmolluscan body plans evolved, since this patten istypical and ubiquitous in molluscan goups such asmonoplacophoans and polyplacophoans.

The origins and early diversity of body plans:

linkage to extant and fossil molluscs

recent paleontological findings have demonstatedunexpected divesity of body plans in the Ediacaanand Cambrian radiation of molluscs (Buttereld, 2006;Caon, et al. 2006; Sigwat and Sutton, 2007). Theevidence fom Kimberella (Fedonkin and Waggone,1997), Wiwaxia (Eibye-Jacobsen, 2004; Caon et al.,2006), Halkieria (Conway Mois and Peel, 1990;Vinthe and Nielsen, 2005), Orthrozanclus (ConwayMois and Caon, 2007), and Odontogriphus (Caon

Figure 2. Taditional views to explain the evolutionay tansition of cephalopod body plans. Above: the ams as foot hypothesis byNaef (1928). The cephalopod ams ae deived fom the oveall egion of the foot. Below: the ams as head hypothesis by Holland (1987)based on Yochelson (1973). The ve arm pairs are derived from the head and an anterior region of the foot surrounding the mouth. The

posteio foot tansfoms into the funnel folds of the cephalopods. In this scenaio, the am-head needs to otate ventally since the funnelof cephalopods is located more dorso-posterior part of the body. (Figures were modied from the original of Naef (1928) and Holland (1987)

with pemission by Geological Society of London).


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et al., 2006) is suppoting the hypothesis that theealy stem-goup of molluscs had always a bilateallysymmetical concentic body and the doso-ventallycompessed pattens as seen in a pototype of Mollusca(Haspuna, 1992). Inteestingly, Kimberella, unlikeextant polyplacophoans and monoplacophoans,obviously displays a seial epetition patten in thefoot sole (Figue 4). The halwaxiid Orthrozanclus ,

possibly a moe deived fom of stem-mollusc, had onepominent shell, a head-like convex at the anteio end,and blade-like, slightly cuved spines in the epipodialegion (Figue 4). Given the simila topogaphy,

these chaactes may pemit pecise compaisonswith Naut ilus and coleoid embyonic body plansto econstuct homological elationships. The mostsubstantial diffeences in the Nautilus embyonic

body plan with othe stem-molluscs ae the following;(1) the educed shell field, (2) the specialiation ofthe hood, colla, and funnel in the epipodial egion,(3) the segmental fou o moe compatments fotentacles in the foot egion, and (4) the enlagement

of the cephalic complex including lage eye pats. Thesimilaity with the ealy embyonic body plans doesnot necessaily mean a similaity with the adult planof the eal cephalopod ancesto. We have actually noinfomation whethe the common cephalopod ancestowent though a so-called laval and/o benthic juvenilestage duing the ontogeny. Howeve, in any case, theshaed concentic o oval entie pattens in fossil andextant molluscs, seem to also provide a rm geometric

basis fo undestanding the stuctual constaints and theevolutionay plasticity in molluscan foms.

Evolutionary transition of phylotypes

What kinds of events occued duing the longevolutionay pocess fom a bilateian ancesto tocephalopods? To addess this question, a simple scenaiomay be helpful fo a compaison among the phylotypicstages of key animals (Figue 5). The phylotype o

phylotypic stage is a stage of development at which allmembes in the taxon look essentially the same (Slack,

Figure 3. A-C. The embyonic development ofNautilus. D. The ealy body plans ofNautilus display a simila geometic concenticcicle patten with the fom of the pimitive gastopod (Lottia gigantea) and the coleoid cephalopod (Idiosepius paradoxus). The embyosofNautilus pompilius, lateal view. A. 3 months afte oviposition, DAPI whole mount staining. B. 4 month old embyo, and C. 6 month oldembyo (the shell was emoved). Details fo the all theNautilus embyos ae given in Shigeno et al. (2008). D. The dosal and lateal viewsof an adult ofLottia gigantea, and schematic gures of the dorsal views ofNautilus and a squidIdiosepius embryo (modied from Shigenoet al. 2008). ceph, cephalic compatment o head pat. I-V, am buds o a pat of tentacle buds. mus, shell muscle.


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2003, i.e., the stage at which vaious adaptive pessuesfo changes ae minimied, and theefoe, the stage most

likely to etain the featues of the common ancesto). Incephalopod embryology, the rst version of phylotype,suggested by Boletky (2003, 2006) based on coleoidembryos, was composed of ve pairs of arm rudiments,

the am base elongated fom each udiment, the anteiohead with eyes, the ostal mouth, and the mantle

pat with concentic o oval pattens of the oveallembyonic body. In the second vesion (Shigeno et al.,2008), diffeent intepetations wee given fo the colla,funnel, head pat, details of tentacles buds, and nevoussystems fom the data ofNautilus embyos. Based onthe Nautilus embyos, intepetation fo the oiginsof some ogans was modified. The phylotypes, in any

vesion, have to be tested by futhe mophological andmolecula analysis.

The last common bilateian ancesto has beenconsideed to be a ceatue like cnidaian planulalavae, o acoelomophs, o annelid tochophoa lavae,although thee ae still many contovesial issues(e.g. Aendt et al., 2001; Bagua and riutot, 2001;Holland, 2003; Ewin, 2006; Hejnol and Matindale,2008). In any event, it may be enough to suppose that

the ealiest condition aleady has a polaity fo the APand DV axes, and some segegated neuons and sensoy

ciliay cells might have been pesent in the anteioegion (Figue 5) (Holland, 2003; raikova et al., 2004;Hejnol and Matindale, 2008).

Fo the common molluscan ancesto, a benthic adultfom has been suggested as the molluscan pototype(Salvini-Plawen, 1972; Haspuna, 1992), but the

phylotype may be econstucted by a developmentals ta te such as the t ochophoa lava, s ince a l lextant membes othe than cephalopods includingaplacophoans, polyplacophoans, and some othecown classes shae a simila developmental pattensof the lava (Naef, 1928; raven, 1958; Okusu, 2002;Fiedich et al., 2002; Nielsen, 2004; Nielsen et al.,

2007). Notably, in the phylotype, the anteio head isgured with an apical tuft, the ventral foot region may

be epesented as a medial goove like an aplacophoanlava (Okusu, 2002), and the nevous system consists ofthe ceebal ganglia, pedal, and pleual cods as shaed

by all membes of molluscs (Haspuna, 1992). Atthis stage in the gastopods, the foot sole is paticulalydistinct. The mantle including the shell field, and thevisceal mass develop dosally. The tipatite neual

Figure 4. Topogaphical pattens in the fossil molluscsKimberella and Orthozanclus with a pimitive cephalopodNautilus embyoto show the simila body plans; the segmental natue o spines developed in the foot egion along the anteio-posteio axis (see inboxesfo details), the anteio head pat (except foKimberella), and the oval shell mophology at the dosal side. These views ae seen fomthe lateal (above) and dosal (below) side. The 3D models ae epoduced foKimberella (Fedonkin and Waggone, 1997), Orthozanclus (Conway Mois and Caon, 2007), andNautilus (Shigeno et al., 2008) by the thee dimensional digital model softwae, Amophium ve.3,EI Technology Goup LLC. ceph, cephalic compatment o head pat.


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Figure 5. A simplied evolutionary scheme for the possible transition from the common bilaterian ancestor to cephalopod body

plans. The focus is paticulaly on the simila geometic pattens aanged along the conseved anteio-posteio and doso-vental axes withspecied neural characters and novel external organs such as the shell, mantle, and the foot. Cephalopods have the largest brain in inverte -

bates with many diffeentiated lobes, though the tipatite neual cod plan of the common molluscan ancesto is still conseved (Budelmannet al., 1997). Each gure was constructed by data combined from various species to show developmentally conserved patterns. Source of g-ues: the nautiloid cephalopod (Shigeno et al., 2008), the late tochophoa lava of pimitive and deived gastopods (Page, 1992; Nedebagtet al., 2002a; Hejnol et al., 2007), the putative laval plan of a molluscan ancesto simila to the tochophoa lava of aplacophoans (Okusu,2002; Nielsen et al. 2007) with polyplacophoans (Voonehskaya et al., 2002; Fiedich et al., 2002; Heny et al., 2004), and a bilateianancesto like an acoelomoph (raikova et al., 2004; Hejnol and Matindale, 2008).


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components (the ceebal, pedal, and pleual codso ganglia) ae always associated with the apical tuftof the pe-tocheal egion, the foot, and the mantle/visceal mass, espectively. It is not clea whethe thegastopods and extinct basal cephalopods shaed avelige-like lava that was adapted to planktonic life.The common ancesto of gastopods and cephalopodshad aleady established a topogaphical aangementof ogan systems that ae commonly identifiablealong the AP and DV axes (Naef, 1928) as seen in the

phylotype of cephalopods (Boletky, 2003). Basedon the position of the foegut and mantle, the head

pat at the whole anteio egion of the pototoch of

gastopods may coespond to the cephalic compatmentincluding eyes and the ceebal cod of pimitivecephalopods. Theefoe, the position of the apical tuftmay additionally be compaed to the labial pat incephalopods. The ceebal and pleual components ofthe nevous systems ae etained as cod-like featues(Shigeno et al., 2008).

If the supeficial elationships ae pesent in eachorgan as stated above, how is the similarity reected inthe eal events of oganogenesis? One similaity is seenin the compaative mophogenetic tissue movement ofthe tochophoa lava and cephalopod embyo (Figue6). In the compaable egions of the vental side of both

Figure 6. Compaison of the vental side of the tochophoe lava and the octopus embyo (without the oute yolk sac) to show thesimilar inward ow of morphogenetic movement. The digestive organ formation is focused on this gure (black). The head part, ventral/

pedal neual teitoies, and inne yolk sac ae epesented by light shading. Left: a mode of amphistomy in the annelid tochophoe lava(Nielsen, C., 2001; fo gastopods, see also Fiooni, 1980; van Biggelaa et al., 2002). The vental lateal sides of amphistome ae to fuseleaving two openings of the anteio apetue fo the mouth, and a posteio opening becomes anus. Vental neual teitoies ae also migat-ed togethe in the lava (Denes, et al., 2007). Right: in the ealy octopus embyo, the half cicle midgut pimodia centalie to fuse a singletube of digestive tact (lage aows) duing the dastic centaliation of whole bodies including neual tissues (Fuchs, 1973; Shigeno et al.,unpublished). In tun, the ecto-mesodemal am bases elongate dosally to cove the whole head pat (small aows).


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Figure 7. Compaative scheme to chaacteie the shaed o unique body plans of vetebates, insects, cephalopods, and cnida-ians. The neual stuctue is shown by light shading o black (anteio non-Hox genes ootd/otx gene expessed teitoies). The foegut o

esophagus is situated at the anteio pat of each animal body. TheHox gene expession code is simply epesented by black bas along theAP axis. The expession ofdecapentaplegic (dpp/BMP2/4) othologs is similaly expessed dosal o vental side with a gadual o discetemanne. Though the expession ofdpp in cephalopods has not been epoted yet, but that of a patellogastopod appea at the dosal side of

petochal head egion and mantle of the lava (Nedebagt et al., 2002a). The data was used fom the multipleAnthox, dpp, and otx genesof sea anemoneNematostella (Finnety et al., 2004; Matus et al., 2006; Maa et al., 2007), the orthodenticle/otx gene,Hox code, and dpp/

BMP2-4 of yDrosophila and a epesentative vetebate (reichet and Simeone, 2001; Holland, 2003), and theHox genes of the squidEu-prymna (Lee et al., 2003).


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embyos, the simila inwad o convegent movementcan be identied to form two openings (mouth and anus)and a single midgut as the main pat of the digestiveogans.

Molecular signature: how are the gene regularity

networks shared?

A supising esult of ecent yeas is that theexpession pattens of developmental contol genes aehighly conseved acoss animal phyla (e.g. Buce andShankland 1998; Aendt and Nble-Jung, 1999; Loweet al., 2003; Finnety et al., 2004; Denes et al., 2007).Some key tansciption factos elated to the embyonic

pat ten fomation have been studied to chaacteiethe cephalopod body plans at the molecula level;

pax-6 (Tomaev et al., 1997; Hatmann et al., 2003),Hox (Lee et al., 2003), and engrailedgene (Baatte

et al., 2007; Shigeno et al., 2008). In these studies,evidence has been shown to suppot a conseved anddeived molecula signatue fo developing embyos incephalopods and othe bilateians (Figue 7).

Fist, Hox genes of the squid exhibit a oughlycolinea expession code in the am pimodiumalong the AP axis as seen in the segments with neualstuctues of insects and vetebates (paticulaly,Scr, Antp, Scr/Hox5) (Aendt and Nble-Jung, 1999;reichet and Simeone, 2001). The expession of

poste ioHox gene Abd-B/Post-2 was detected inthe most posteio teitoy in a developmental stageof gastopods (Hinman et al., 2003), squids (Lee et

al. , 2003), and annelids (Pudhomme et al., 2003).Inteestingly, no Hox genes ae eve expessed in alage pat of the anteio head pat, wheeas otx/otdothologous ae expessed in the teitoies as seen inthose of insects, vetebates, annelids, and gastopods tochaacteie a compaable teitoy fo the shaed head

pats (Aendt and Nble-Jung, 1999; Nedebagt et al.,2002b; Hinman et al., 2003; Kulakova et al., 2007).

Second, a paied domain containing tansciptionfactopax-6/ey eless of the squid is expessed atthe ealy eye pimodium like those of insects andvetebates. The gene is usually expessed in the etinalcells in vetebate and insects, howeve it is not seen insuch cells of squids (Tomaev et al., 1997), suggestinga deived featue fo photoecepto developmentof cephalopods in a conseved manne of the eyeevolution.

Thid, a homeodomain containing tansciption factoEngrailedof the decabachian cephalopods is expessedin vaious ogan pimodia such as ams, funnel, collaand the mantle (Baatte et al., 2007; Shigeno et al.,

2008). Unlike the pattens of insects and vetebates, thefeatue did not exhibit a segmental patten as a polaitygene along the AP axis. The cephalopod engrailedalso may not be involved in the neual patteningas epoted in that of gastopods (Nedebagt et al.,2002a). Inteestingly, the cephalopod and gastopodengrailedgenes ae highly expessed at stuctues suchas the shell field and the mantle magin, suggestinga novel ole in shaping such ogans along the DVaxis. It is still an open question whethe cephalopodembyos have a shaed genetic pogam to fom a DVaxis (Figue 7). Howeve, evidence fo the expessionofdecapentaplegic(dpp)/BMP2/4 at the dosal side ofvaious bilateians including gastopods (Nedebagt etal., 2002a), and the involvement ofBrachyury gene tothe vental side of foegut development (Latillot et al.,2002) may indicate that a shaed molecula signatuealong the body axes had aleady been established at a

pe-bilateian animal (Finnety et al., 2004; Matus etal., 2006).Finally and moe speculatively, if the dosal side of

vetebates coesponds to the vental side of molluscs,it may be possible to popose that the common inwadmovement on the vental side of tochophoe lavaand cephalopod embyo (as seen in Figue 6) can becompaed to the medial convegent movement to fomthe neual tube of vetebates (as patly suggested byvan den Biggelaaet al., 2002).

Conclusion

In this eview, we have attempted to descibe anevolutionay linkage between cephalopod body plansand othe bilateians with special focus on ealytopogaphic patten fomation. The pattens of ogansystems have obviously displayed a conseved signatueas well as evolutionay novelty. Ou knowledge istoo limited to elucidate the complete undestandingof the oigins of cephalopods; howeve, the emegingidea and new intepetation of ecently obtained datasuppot the conclusion that cephalopods exhibit anenigmatic evolutionay histoy compaed to those ofwell investigated model animals such as insects andvetebates.

Acknowledgements

We thank Takeya Moitaki, Toba Aquaium inJapan, fo the collection of nautiloids. This wok wassuppoted by the postdoctoal fellowship fo eseachaboad fom the Japan Society fo Pomotion ofScience, and Foundation of reseach Institute of Maine


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Invetebates to S.S. This study was also suppoted bya Gant-in-Aid fom Japan Society of the Pomotion ofScience (No. 20540445) to T.S.

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