04a origen metazoa-coanoflagelados

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    HemichordataChordata

    Z

    A -B -

    C -D -E -F -G -H-I -K -L -

    M -N -O -P -Q -R -S -T -U -V -

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    AB

    C

    D

    E

    F

    G

    H

    I

    KK

    L

    M

    N

    O

    P

    Q

    R

    SU

    V

    X Y

    AnimaliaEuradiculataProepitheliozoaEumetazoaNeuraliaTriploblastica

    BilateriaAcoelomorphaEubilateriaProtostomiaSpiraliaSchizocoeliaGnathiferaPolyzoaBrachiozoaEcdysozoa

    Panarthropoda

    CycloneuraliaNematoideaScalidophoraDeuterostomiaAmbulacrariaHemichordataChordata

    Z

    A -B -C -D -

    E -F -G -H-I -K -L -M -N -O -

    P -Q -R -

    S -T -U -V -X -Y -

    Z -

    T

    T ..

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

    11/29

    !"#$#!%

    !!

    < 43.?o/7 .?6230 ,34/]./?6.7

    5(3 /0:374?/* ,34/]./0 C/7 8?.;/;*)

    / (>#&*#9)&38&"& K`62D RD%M9 C(6:(

    (/E E3F3*.83E 7. ()#3" (")) (#*8&(83

    4(/4 0+4?63047 :.+*E ;3 3N:(/023E

    ;34C330 :3**7G 4(67 ,/E3

    783:6/*6]/I.0 .> :3**7 8.776;*39

    ;3:/+73 0.4 /** :3**7 (/E 4. >33ED

    X4 8?.;/;*) (/E / *6>3 :):*3 C64(

    3"@/&) $"A$#5/(B#* &*5 >&A)#+5

    "CC3 &*5 3A"$D9 :(/?/:43?67I: .> 4(3

    *6F602 ,34/]./07D

    Z3N+/* ?38?.E+:I.0 .?6260/43E /4 /0

    3/?*) 7438 60 4(3 3F.*+I.0 .>

    3+J/?).4379 ;+4 64 :.+*E .> :.+?73

    (/F3 ;330 *.74 60 4(3

    :(./0.1/23**/437D

    Choanoblastaea Advancedchoanoblastaea

    Choanoflagellate

    A

    occluding junction

    circumblastoporalnerve

    sensory cellgerm cells

    cadherins

    mitosis

    basementmembrane

    nervewater current

    < 43.?o/7 .?6230 ,34/]./?6.7

    5(3 03C +0E3?74/0E602 .> 4(3 3/?*)

    ,34/]./0 ?/E6/I.0 C64( /

    8/?/8()*3I: '.?6>3?/ 6,8*637 4(/4 4(3

    *6>3 :):*3 >.+0E 60 4(3 3A#*C" C$&5"9

    C64( / :6*6/43E >?33@7C6,,602 */?F/

    /0E / 73776*3 /E+*4 >.?, C64( / C/43?@

    :/0/* 7)743, /0E :(./0.:)43

    :(/,;3?79 C/7 /0:374?/*D 5(3

    3F.*+I.0 .> 4(67 4)83 .> .?2/06]/I.0

    67 F67+/*6]3E 60 `62D RD%Dll

    l ll

    Ancestral sponge

    Primitive sponge

    l

    l li i l

    l i

    < 43.?o/7 .?6230 ,34/]./?6.7

    [ :(./0.;*/74/3/ *603/23 E3F3*.83E

    / 0+,;3? .> +*8"$*&)+0"5 ("))3K4(3

    A/EF/0:3E :(./0.;*/74/3/BM9 /0E

    "38&9)+3>"5 & A#)&$+8< ;+8> 3A"(+&)

    ("))3 ;+8>#/8 (#))&$3 &8 8>" &*8"$+#$

    A#)"?Choanoblastaea Advanced

    choanoblastaea

    Choanoflagellate

    A

    occluding junction

    circumblastoporalnerve

    sensory cellgerm cells

    cadherins

    mitosis

    basementmembrane

    nervewater current

    .. . . .

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

    12/29

    !"#$#!%

    ! 6043?0/*6]3E :3**7 K4(3

    A/EF/0:3E :(./0.;*/74/3/BM9 /0E

    374/;*67(3E / 8.*/?64) C64( 783:6/*

    :3**7 C64(.+4 :.**/?7 /4 4(3 /043?6.?

    8.*3D

    5(373 :3**7 C3?3 +73E 60 73L*6029 /0E

    4(3 (>#&*#( /*?3/E) 8?.:3773E

    C/43?D

    Advancedchoanoblastaea

    Ancestral sponge

    circumblastoporalnerve

    sensory cell

    nerve

    Primitive sponge

    . ,, . . . , .

    < 43.?o/7 .?6230 ,34/]./?6.7

    5(67 A/0:374?/* 78.023B ?34/603E 4(3

    83*/26: 74/23 /7 / E6783?7/* */?F/*

    74/239 /0E / 83*/2.;304(6: *6>3 :):*3

    (/E ;3:.,3 374/;*67(3ED

    5(3 C$##E"C64( 4(3 F+*03?/;*3

    :(./0.:)437 9"(&D"A$#8"(8"5;)

    .F3?/?:(602 >?., 4(3 */43?/* :3**7 7.

    4(/4 / 4+;+*/? A:(./0.:)43 :(/,;3?B

    C/7 >.?,3E9 >+?4(3? 30(/0:602 4(3

    C/43? :+??3047 KA4(3 8?6,6IF3

    78.023BMD

    l

    l a

    l ll

    Ancestral sponge

    Primitive sponge

    l

    l li i l

    l i

    I8.>3('+

    < 43.?o/7 .?6230 ,34/]./?6.7

    5(67 67 /0 .?2/06]/I.0 ?373,;*602 4(3

    *6F602 :/*:/?3/07 .> 4(3>3('+ @4)83

    K733 -(/843? W M9 3N:384 4(/4 64 (/E 0.

    7J3*34.0D

    !>+3 =#$D (#/)5 C$#; 8# & )&$C"$ 3+0"

    ;) #$C&*+0+*C 8>" )&&D9"$39 /0E 4(3 */?23?76]3 ,/E3 64 8.776;*3 4. 60F374 ,.?3

    ).*J 60 4(3 32279 7. 4(/4 E3F3*.8,304

    :.+*E ;3:.,3 *3:64(.4?.8(6:D

    ll a

    l ll

    Ancestral sponge

    Primitive sponge

    l li i l

    >3('+

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

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    !"#$#!%

    !%

    < 43.?o/7 .?6230 ,34/]./?6.7

    5(373 *#*,=""5+*C */?F/3 /;/0E.03E

    4(3 3N43?0/* :(./0.:)437 C64( 4(3

    +0E+*/4.?) :6*6+,9 ;+4 5"E")#A"5 &

    *"; 8 (+)+&(/7 / *.029 ,.?3 .?

    *377 .?2/06]3E ?..49 /0E 3>#;3

    (##$5+*&8"5 D#E"D"*83 9"8;""*

    *"+C>9#/$+*C ("))39 :?3/I02

    :(/?/:43?67I: D"8&(>$#*&) ;&E"3G

    K`62D 3 :):*3 7330 60

    ,.74 *6F602 A78.0237BD

    w /EF/0:3 /0E ?3:.F3?) :):*37

    Choanoblastaea Advanced

    choanoblastaea

    Choanoflagellate

    Ancestralspongeoccludingjunction

    circumblastoporalnerve

    sensorycellgerm cells

    cadherins

    mitosis

    basementmembrane

    nervewater current

    Primitivesponge

    . h l ut i n r n r i t t . h t r u rr n t r i n i t it h l u r r , th n t r n ir , n th n r u ll r n . h u l l l f th tr h n itt f r l rit . i ur . , .

    Modern

    homoscleromorph

    B

    Homoscleromorph-like

    ancestor withgastrulalarvaand dissogony

    Gastraea

    Neurogastraea

    Thisgradeis

    represented

    by Trichoplax

    Thisgradeis

    represented

    byCnidaria

    >3('+

    < 43.?o/7 .?6230 ,34/]./?6.7

    !;# )+*"&C"3 :/0 ;3 ?3:.206]3E >?.,

    4(3 &*("38$&) 3A#*C"d #*"*603/23

    *3/E602 4. 4(3 7+)+("&9 C(6:(

    E3F3*.83E / 3+)+("#/33.")"8#*

    K-(/843? SMG /0E &*#8>"$ )+*"&C"

    *3/E602 4. 4(3 H/$&5+(/)&8& K-/*:/?3/

    x '?.3864(3*6.]./G -(/843? V M 4(/4

    5"E")#A"5 ($#33,38$+&8"5 (+)+&$< $##83

    60 4(3 3864(3*6+,@*6J3 .+43? :3** */)3?

    .> 4(3 */?F/3 K`62D VD!MD 5(3

    -/*:/?3/07 E3F3*.83E (&)(&$"#/3

    3A+(/)"3K -(/843? W MD

    Choanoblastaea Advancedchoanoblastaea

    Choanoflagellate

    Ancestralspongeoccludingjunction

    circumblastoporal

    nerve

    sensorycellgerm cells

    cadherins

    mitosis

    basement

    membrane

    nervewater current

    Primitivesponge

    . h l ut i n r n r i t t . h t r u rr n t r i n i t it h l u r r , th n t r n ir , n th n r u ll r n . h u l l l f th tr h n itt f r l rit . i ur . , .

    Modernhomoscleromorph

    B

    Homoscleromorph-like

    ancestor withgastrulalarvaand dissogony

    Gastraea

    Neurogastraea

    Thisgradeis

    represented

    by Trichoplax

    Thisgradeis

    represented

    byCnidaria

    >3('+

    < 43.?o/7 .?6230 ,34/]./?6.7

    5(3 I$#"A+8>")+#0#&

    Ke.,.7:*3?.,.?8(/ x c+,34/]./M

    5"E")#A"5 (#))&C"* JK +* 8>" 9&3&)

    D"D9$&*"9 /0E 4(3 P?74 74/237 .>

    #(()/5+*CL/*(B#*3;3:/,3

    374/;*67(3ED

    M*" )+*"&C" *3E 4. 4(3

    -#D#3()"$#D#$A>&K-(/843? "M9C(6:( ?3,/603E /4 4(67 74/23D 5(3

    .4(3? *603/23 *3E 4. 4(3 H/D"8&0#&9

    C(6:( C6** ;3 E67:+773E 60 -(/843?

    != D

    Choanoblastaea Advanced

    choanoblastaea

    Choanoflagellate

    Ancestralspongeoccludingjunction

    circumblastoporalnerve

    sensorycellgerm cells

    cadherins

    mitosis

    basementmembrane

    nervewater current

    Primitivesponge

    . h l ut i n r n r i t t . h t r u rr n t r i n i t it h l u r r , th n t r n ir , n th n r u ll r n . h u l l l f th tr h n itt f r l rit . i ur . , .

    Modernhomoscleromorph

    B

    Homoscleromorph-likeancestor withgastrula

    larvaand dissogony

    Gastraea

    Neurogastraea

    Thisgradeis

    represented

    by Trichoplax

    Thisgradeis

    represented

    byCnidaria

    >3('+

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

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    !"#$#!%

    !R

    k37:30E3,.7 E3 :./0.1/23*/E.7

    :.*.06/*37

    k3P06:6p0 E3 [06,/*

    N/8$+B#*&) O#5"

    [06,/*7 E6\3? >?., ;.4( 8*/047 /0E >+026 604(36? ,.E3 .> 0+4?6I.0D

    I)&*83 &$" &/8#8$#A>+( 3+J/?).437 :/8/;*3 .>2303?/I02 .?2/06: ,.*3:+*37 4(?.+2(A>#8#3"3+3D

    P/*C+ &$" >"8"$#8$#A>3 4(/4 2?.C .0 .? 03/?4(36? >..E /0E 4(/4 >33E ;) &93#$AB#*K.y30/y3? 4(3) (/F3 ?3*3/73E 30]),37 4(/4 E623744(3 >..E .+476E3 4(36? ;.E637MD

    g0*6J3 8*/0479 /06,/*7 :/00.4 :.074?+:4 /** .>4(36? .C0 .?2/06: ,.*3:+*37 /0E 7.9 60 ,.74:/737D

    5(3) +*C"38 4(3,@364(3? ;) "&B*C #8>"$)+E+*C #$C&*+3D3 #$ 9< "&B*C *#*)+E+*C#$C&*+( D&8"$+&)D l+4 +0*6J3 >+0269 ,.74/06,/*7 E. 0.4 >33E ;) /;7.?8I.0G 60743/E9/06,/*7 602374 4(36? >..E /0E 4(30 +7330]),37 4. E62374 64 C64(60 4(36? ;.E637D

    Q")) 78$/(8/$" &*5 7A"(+&)+0&B#*

    R*+D&)3 &$" "/.&$+0269 (.C3F3?9&*+D&)3 )&(. 8>" 38$/(8/$&)3/AA#$8 #= (")) ;&))3D J*38"&56&*+D&) ("))3 &$" >")5 8#C"8>"$9< 38$/(8/$&) A$#8"+*36 8>"D#38 &9/*5&*8 9"+*C (#))&C"*K733 62+?3 VD%=M9 ;>+(> +3=#/*5 #*)< +* &*+D&)3D

    k3P06:6p0 E3 [06,/*

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

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    !"#$#!%

    !S

    k3P06:6p0 E3 [06,/*

    S"A$#5/(B#* &*5 T"E")#AD"*8

    O#38 &*+D&)3 $"A$#5/(" 3"@/&))" )+="(783:6/*6]3E :3**7 0.4 7330 60 .4(3?,+*I:3**+*/? .?2/067,7d D/3()"("))3/0E *"$E"("))3D X0 ,.74/06,/*79 4(373 :3**7 /?3 .?2/06]3E604. ,+7:*3 I77+3 /0E 03?F.+7I77+39 ?3783:IF3*)9 /0E &$"$"3A#*3+9)" =#$ D#E+*C 8>" 9#5&8 5+F"$"*B&8"&*+D&)3 =$#D A)&*83 &*5 =/*C+6D&.+*C D/3()" &*5 *"$E" ("))3("*8$&) 8# ;>&8 +8 D"&*3 8# 9" &*&*+D&)?

    k3P06:6p0 E3 [06,/*

    k3P06:6p0 E3 [06,/*

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

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    !"#$#!%

    !V

    k3P06:6p0 E3 [06,/*

    k3P06:6p0 E3 [06,/*

    . Generalized cell lineage of an annelid.(Based on Podarkeand Arenicola; Child ; Treadwell ).See Figure .,.

    {

    1a1

    1a2

    1a

    2a11

    2a122a1

    2a2

    2a

    3a

    3A

    2A

    1A

    A

    1a12

    1a111a111

    1a121

    1a122

    1a112

    B

    C

    D

    1d

    1d11d11

    1d111

    1d112

    1d12

    1d2

    2d2

    2D3d

    4d

    4D3D

    apical cell

    1D

    prostomial ectoderm

    prostomial ectoderm

    accessory trochoblast

    AB

    Z

    primary trochoblast

    ectomesoderm

    as A, except that the right cerebral ganglion develops from 1c 112,

    there is no ectomesoderm from the B-quadrant, and that theectomesoderm originates from 3c21in the C-quadrant

    prostomial ectoderm

    primary trochoblast

    CD

    endoderm

    apical cell

    1a1222

    1a1221 prostomial ectoderm

    2a111

    2a112

    2a121

    2a122

    ectoderm

    secondary trochoblast

    secondary trochoblast

    ectoderm

    ectoderm

    3a1 ectoderm

    3a23a21

    3a223a221

    3a222stomodaeum

    stomodaeum

    endoderm

    prostomial ectoderm

    2d111 ectoderm

    2d112 somatoblast2d11

    2d12 ectoderm, telotroch

    2d1

    2d21 ectoderm, telotroch

    2d22 ectoderm, proctodaeum, ectomesoderm

    2d

    ectoderm, proctodaeum, ectomesoderm

    mesoderm, endoderm

    {

    {{

    {{

    {

    {

    { {

    {

    {{

    {{ { {{ {

    {

    {

    {{

    {{

    {

    apical ganglion

    apical ganglion

    cerebral ganglia

    circumblastoporalnerve cord, incl.posterior part ofbrain

    {

    {

    {

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

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    !"#$#!%

    !W

    k3P06:6p0 E3 [06,/*

    S"A$#5/(B#* &*5 T"E")#AD"*8

    [*4(.+2( /E+*4 &*+D&)3F/?) C6E3*) 60,.?8(.*.2)9 4(3 +0E3?*)602 C"*"B(*"8;#$. 8>&8 (#*8$#)3 &*+D&)5"E")#AD"*8 +3 3+D+)&$ &($#33 & 9$#&5$&*C" #= 8&@&?[** 3+J/?).437 (/F323037 4(/4 ?32+*/43 4(3 3N8?3776.0 .>.4(3? 230379 /0E ,/0) .> 4(373?32+*/4.?) 23037 :.04/60 :.,,.0 7347.> kh[ 73O+30:37 :/**3E #'5%'6'1%3K733 -(/843?

    R*+D&)3 3>&$" & /*+U/" >#D"#9#@,(#*8&+*+*C =&D+)< #= C"*"36 .*#;* &3!"# %&'&(/ e.N 23037 8*/) 6,8.?4/04?.*37 60 4(3 E3F3*.8,304 .> /06,/*3,;?).79 :.04?.**602 4(3 "@A$"33+#* #=5#0"*3 #$ "E"* >/*5$"53 #= #8>"$C"*"3 8>&8 +*V/"*(" &*+D&)D#$A>#)#C< K733 -(/843? ,/4+?/I.09 ;+4 4(3*6>3 :):*37 .> ,/0) /06,/*7 /*7. 60:*+E3/4 *3/74 .03 */?F/* 74/23D

    [ )&$E&67 / 73N+/**) 6,,/4+?3 >.?, .>/0 /06,/* 4(/4 67 D#$A>#)#C+(&))" &5/*49 +7+/**) "&835+F"$"*8 =##59 /0E ,/) 3F30 (/F3 /5+F"$"*8 >&9+8&8 4(/0 4(3 /E+*49 /7 604(3 :/73 .> 4(3 /O+/I: */?F/ .> /,.7O+64. .? / E?/2.01)D

    [06,/* */?F/3 3F304+/**) +0E3?2.D"8&D#$A>#3+39 / 5"E")#AD"*8&) 8$&*3=#$D&B#*4(/4 4+?07 4(3 /06,/*604. / H+F306*39 C(6:( ?373,;*37 /0/E+*4 ;+4 67 0.4 )34 73N+/**) ,/4+?3D

    r r r r r r , rr r . r r r r r

    r . r r r r r , r r r , rr r , r r r r , . r . r

    , rr , r , r ,, , r , ,

    r , , r , r, . r r r r r

    r . r . , .

    Branchiostoma

    Saccoglossus

    Nematostella

    Symsagittifera

    Xenoturbella

    Capitella

    Tribolium

    Priapulus

    lab pb Hox3 Dfd Scr

    Lox2 Lox4 Post1 Post2

    Pecten

    Spiralia

    Platyhelminthes

    Lingula

    1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

    Strongylocentrotus

    Ambulacraria9/10 11/13 abc

    Eubilateria Deuterostomia Chordata

    Mus

    ? ? ? ? ?

    EcdysozoaUbx abd-A abd-B

    ProtostomiaLox5 Antp

    k3P06:6p0 E3 [06,/*

    S"A$#5/(B#* &*5 T"E")#AD"*8

    Z8.02379 C(6:( /?3 /,.02 4(3 76,8*3743N4/04 /06,/*79 (/F3 @'1 .?,/I.0 .> C/43? :(/003*7 60 4(3 ;.E) C/**9/ J3) >3/4+?3 .> 78.023 ,.?8(.*.2) K733-(/843? %%MD

    X0 4(3 /0:374.?7 .> ,.?3 :.,8*3N /06,/*794(3 @'1 /,6*) +0E3?C304 >+?4(3?

    E+8*6:/I.079 )63*E602 / ,.?3 F3?7/I*3z4..*J64z >.? ?32+*/I02 E3F3*.8,304D X0F3?43;?/4379 6073:479 /0E ,.74 .4(3? /06,/*79

    @'1 4(3/[email protected]?6.? K>?.04@4.@;/:JM /N679 /7 C3**/7 .4(3? /783:47 .> E3F3*.8,304D 5(3 7/,3:.073?F3E 2303I: 034C.?J 2.F3?07 4(3E3F3*.8,304 .> ;.4( / 1) /0E / (+,/09E378643 4(36? .;F6.+7 E6\3?30:37 /0E(+0E?3E7 .> ,6**6.07 .> )3/?7 .> E6F3?23043F.*+I.0D

  • 7/24/2019 04a ORIGEN METAZOA-Coanoflagelados

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    "@B*(8 #*"3D KZ.,3 8/*3.04.*.26747

    (/F3 37I,/43E 4(/4 WW{ .> /** /06,/*

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    E6F3?764) (/7 647 .?62607 60

    3F.*+I.0/?) :(/0237 4(/4 .::+??3E

    E+?602 4(3 */74 ;6**6.0 )3/?7D

    `.? 3N/,8*39 7.,3 37I,/437 ;/73E

    .0 ,.*3:+*/? :*.:J7 7+22374 4(/4 4(3

    /0:374.?7 .> /06,/*7 E6F3?23E >?.,

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    8>&8 8>" (#DD#* &*("38#$ #= )+E+*C

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    ()#3"38 )+E+*C $")&BE"3 #= &*+D&)3D

    l/73E .0 7+:( ?37+*479 ?373/?:(3?7

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    (.C /06,/*7 3F.*F3E >?., 4(36?

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