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<CENTER><B>HAECKEL AND THE VERTEBRATE ARCHETYPE</B> </CENTER></H2>
<P>&nbsp;</P>
<P><FONT face=3DPalatino>A recent paper by Michael Richardson and his =
colleagues=20
(1997) has corrected a major misunderstanding among developmental =
biologists and=20
evolutionary biologists which can be traced back to Ernst Haeckel. =
Haeckel=20
(1874) had claimed that members of all vertebrate classes pass through =
an=20
identical evolutionarily conserved "phylotypic" stage. Until this new =
paper=20
appeared, it was assumed that Haeckel was correct and that there was a=20
particular stage of development that was identical to all vertebrates. =
Only=20
later in development would specific differences appear. Interestingly, =
there was=20
some discussion as to what exactly this stage was (Richardson , 1995). =
This=20
conserved stage was sometimes considered the neurula stage (Wolpert, =
1991), the=20
"pharyngula" stage (characterized by the branchial arches; Ballard, =
1981), the=20
tailbud stage (Slack et al., 1993), or the stages between those of =
headfold and=20
tailbud (Duboule, 1994). Figure 1 shows one of Haeckel's representation =
of=20
vertebrate embryology. The top row shows that all the vertebrate embryos =
(no=20
matter what path they took to reach this stage) have a stage where they =
appear=20
almost identical.</FONT></P>
<P>&nbsp;</P>
<P>
<CENTER><IMG height=3D416 =
src=3D"http://zygote.swarthmore.edu/Richardson1.gif"=20
width=3D482 align=3Dbottom X-SAS-UseImageHeight =
X-SAS-UseImageWidth></CENTER>
<P></P>
<P><B>FIGURE 1. Haeckel's 1874 version of vertebrate embryonic =
development. The=20
top row shows an early stage common to all groups, the second row shows =
a middle=20
stage of development, and the bottom row shows a late stage embryo. =
Groups from=20
left to right are: fish, salamander, turtle, chicken, pig, cow, rabbit, =
and=20
human. (Adapted from Gilbert, 1997.) </B></P>
<P>&nbsp;</P>
<P><FONT face=3DPalatino>But Haeckel's drawings are wrong. Photographing =
actual=20
embryos at these stages, Richardson and colleagues show that Haeckel's =
drawings=20
are oversimplified to the point of obscuring important differences =
between=20
classes of vertebrates. The Richardson et al. paper does not dispute =
that there=20
is a highly conserved embryonic stage among the vertebrate classes. =
Indeed, at=20
the late tailbud stage, vertebrate embryos of most all classes possess =
"somites,=20
neural tube, optic anlagen, notochord, and pharyngeal pouches." However, =
these=20
authors do criticise the notion that this stage is nearly identical in =
all=20
species and that differences beteen the classes can be resolved only =
after=20
subsequent development. Rather, they discover significant differences =
between=20
groups. Size is one distinctive marker. The scorpion fish embryo is 700 =
microns=20
long at the tailbud stage, while the mudpuppy salamander measures some 9 =

millimeters. Heterochrony is another problem. In some species of direct=20
developing frogs, and in monotreme mammals, limb buds are already =
present at the=20
tailbud stage, whereas in other species, these are not seen until =
significantly=20
later. Birds are characterized by their prominent mesencephalon. Whereas =
most=20
amniote embryos have a heart by this stage, the zebrafish does not. =
(Teleosts=20
such as zebrafish are even the exception to the rule mentioned above. =
They=20
eventually possess a notochord, somites, pharyngeal pouches, etc., but =
they do=20
not have the pharyngeal pouches until </FONT><I><FONT=20
face=3DPalatino>after</FONT></I><FONT face=3DPalatino> the tailbud =
stage). Some of=20
these differences are depicted in Figures 2 and 3.</FONT></P>
<P>&nbsp;</P>
<P>
<CENTER><IMG height=3D308 =
src=3D"http://zygote.swarthmore.edu/Richardson2.jpeg"=20
width=3D487 align=3Dbottom X-SAS-UseImageHeight =
X-SAS-UseImageWidth></CENTER>
<P></P>
<P><B></B>&nbsp;</P>
<P><B>FIGURE 2. Selected embryos shown at the "phylotypic" stage. (A) =
Sea=20
lamprey (jawless cyclostome fish, Petroyzon) with nearly identical =
pharyngeal=20
pouches and heart that is caudal to the pharynx; (B) electric ray =
(cartilagenous=20
chondrichthes fish, <I>Torpedo</I>) with nearly identical pharyngeal =
pouches and=20
a pronounced hindbrain/midbrain flexure; (C) sterlet (bony Osteichthyes =
fish,=20
<I>Acipenser</I>) where the pharyngeal pouches have not yet formed; (D) =
frog=20
(direct -developing anuran amphibian <I>Eleutherodactylus</I>) which has =

hindlimb buds at the tailbud stage and only two pairs of aortic arches; =
(E) pond=20
turtle (chelonian reptile, <I>Emys</I>) where there is no craniocaudal =
rotation=20
(torsion) to the embryo, whereas the same stage of the chick (avian) =
embryo (F)=20
shows pronounced torsion; (G) brush-tailed possum (marsupial mammal,=20
<I>Trichosurus</I>) where there is (in contrast to the cartilagenous =
fishes)=20
large maxillary and mandibular processes. In ther cat (eutherian mammal, =

<I>Felis</I>) there is a similar size difference between the anterior =
and=20
posterior pouches, although their appearance differs from that of the=20
marsupials. (After Richardson, 1997.)</B></P>
<P><B></B>&nbsp;</P>
<P><B></B>&nbsp;</P>
<P><B><IMG height=3D432 =
src=3D"http://zygote.swarthmore.edu/Richardson3.gif"=20
width=3D480 align=3Dbottom X-SAS-UseImageHeight =
X-SAS-UseImageWidth>FIGURE 3.=20
Photographs of a fish, amphibian, reptile, mammal, and bird embryo at =
the=20
"phylotypic" stage. Photographs courtesy of M. Richardson.</B> </P>
<P><FONT face=3DPalatino>Interestingly, this knowledge appears to be =
"old hat"=20
among German biologists. Haeckel's drawings were not trusted (see =
Goldschmidt,=20
1956), and Haeckel was accused of scientific fraud by a university court =
in=20
Jena, where he worked and by other embryologists, as well (see Hamblin, =
1997;=20
Richardson et al., 1997b). Yet, the idea that early vertebrate embryos =
are=20
essentially identical has survived. I think there were two reasons for =
the=20
survival. First, Haeckels' illustration was reproduced in Romanes'=20
(1901)</FONT><I><FONT face=3DPalatino> Darwin and After =
Darwin</FONT></I><FONT=20
face=3DPalatino>. From here, the illustration entered Anglophone =
biology,=20
"sanitized" from Haeckel. Second, the picture can be used (as it has =
been in=20
several developmental biology books, including my own [Gilbert, 1997, p. =
254])=20
to illustrate von Baer's principles rather than Haeckel's biogenetic =
law. K. E.=20
von Baer had noted that the general features of a large group of animals =
appear=20
earlier in the embryo than do the specialized features. Indeed, von Baer =

wrote:</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<BLOCKQUOTE>
  <P><FONT face=3DPalatino>"The embryo of the mammal, bird, lizard, and =
snake and=20
  probably also the turtle, are in their early stages so uncommonly =
similar to=20
  one another that one can distinguish them only according to their =
size. I=20
  posess two small embryos in spirits of wine, embryos whose name I =
neglected to=20
  note down, and I am now in no position to determine the classes to =
which they=20
  belong. They could be lizards, small birds, or even very young=20
  mammals."</FONT></P></BLOCKQUOTE>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Darwin quoted this (it was Thomas Huxley's =
translation)=20
although he misattributed the story to Agassiz rather than to von Baer =
(see=20
Richards, 1992 for an analysis). The notion of development proposed by =
von Baer=20
is still used as a general approximation of certain developmental =
phenomena.=20
Certainly, the early embryos of certain vertebrate classes are very =
similar (for=20
instance, the chick and mouse twenty-somite embryos), and the processes =
of=20
somitogenesis, limb formation, axis generation, etc. are probably =
conserved=20
throughout the vertebrate groups).</FONT></P>
<P><B><FONT face=3DPalatino></FONT></B>&nbsp;</P>
<P><B><FONT face=3DPalatino>Similarities and differences</FONT></B> </P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>What the Richardson et al. paper tells us is =
that we have=20
to be attentive to the differences as well as to the similarities. A =
bird embryo=20
is not the same as a mammalian embryo. We should not assume identity, =
even at a=20
stage where all vertebrate embryos seem to pass. The Richardson et al. =
paper can=20
be read as a critique of the recent priority of similarities over=20
differences.</FONT></P>
<P><FONT face=3DPalatino>The concept of homology enables one to =
celebrate the=20
differences or the similarities between two structures. Whether one =
emphasizes=20
the similarities between our forelimb and a bird's wing or the =
differences=20
between them depends on what you are describing. Comparative =
anatomy--with its=20
Aristotelian and Cuverian interest in relating structure to =
function--usually=20
emphasizes the differences. Morphology--with its Platonic and Geoffroyan =

interest in the underlying unities of structure--usually focuses on the=20
similarities.</FONT></P>
<P><FONT face=3DPalatino>As Joseph Needham noted, embryology has swung =
between=20
these two poles during different stages of its morphogenesis. In the =
late 1800s,=20
the morphological tradition prevailed, and the similarities between=20
developmental stages in different organisms constituted some of the best =

evidence for classification (see Nyhart, 1995; Bowler, 1996). Thus, the=20
discovery of the Nauplius stage of the barnacle showed that it was a =
modified=20
crustacean, and the notochord-containing tadpole of the tunicate =
demonstrated=20
its affinities with the chordates. However, after the 1920s, embryology =
was no=20
longer a major support for evolutionary biology, and the comparative =
anatomy=20
tradition came to predominate. Until the 1980s, embryology was extremely =

descriptive. Each organ was seen to develop differently from any other =
organ,=20
and each species was seen to develop differently from any other species. =
Indeed,=20
embryology was defined (by one of its practitioners, E. G. Conklin) as a =

"lawless science" because generalities could not be made from the =
observations=20
of animal development. When embryology underwent its anagenic transition =
to=20
become developmental biology, the similarities among organisms were =
again=20
emphasized. The similarities were now posited on the molecular rather =
than the=20
morphological or cell lineage level. The 1990s has seen a remarkable =
celebration=20
of the similarity of molecular processes throughout the animal kingdom.=20
Homologous genes abound (the </FONT><I><FONT =
face=3DPalatino>Hox</FONT></I><FONT=20
face=3DPalatino> genes, </FONT><I><FONT =
face=3DPalatino>fringe</FONT></I><FONT=20
face=3DPalatino> , </FONT><I><FONT =
face=3DPalatino>tinman</FONT></I><FONT=20
face=3DPalatino>, and </FONT><I><FONT =
face=3DPalatino>Pax6</FONT></I><FONT=20
face=3DPalatino> being seen to specify the anterior-posterior axis, the =
limb, the=20
heart, and the eye, respectively, of organisms as diverse as insects and =
flies).=20
Even signalling pathways are seen as being homologous both within a =
developing=20
organism and between organisms. Thus, the neural tube in vertebrates and =
insects=20
are seen as being formed through the same interactions of the "same" =
proteins,=20
even though one neural tube is dorsal and the other ventral. The =
Richardson et=20
al paper reminds us that despite these similarities, differences are =
also=20
important, especially if one is thinking in terms of the relationships =
between=20
development and evolution.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>The Richardson et al paper is a also plea for =
comparative=20
anatomy and less reliance on "model systems." This critique of model =
systems=20
echoes a recent plea from Leo Buss, Rudy Raff, Jessica Bolker, and James =
Hanken=20
who observe that model systems converge on several factors (see Bolker =
and Raff,=20
1997; Hanken, 1993): they all can develop in the laboratory and are thus =

relatively free of environmental factors effecting their morphogenesis; =
they are=20
all organisms which segregate their germ line extremely early; they =
develop=20
rapidly, are small as adults, and they have short generation =
times.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>The Richardson et al. paper does a great =
service to=20
developmental biology. As they mention, there is renewed interest in=20
evolutionary developmental biology, and in view of the conservation of=20
developmental mechanisms, we need to re-examine the extent of variation =
in=20
vertebrate embryos. There is more variation than had been assumed, and =
these=20
variations foreshadow important differences in the adult bodies. This is =

important because we need to know how conservative the early embryonic =
stages=20
actually are.</FONT> </P>
<P><B><FONT face=3DPalatino></FONT></B>&nbsp;</P>
<P><B><FONT face=3DPalatino>References: </FONT></B></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Ballard, W. B. 1981. Morphogenetic movements =
and fate=20
maps of vertebrates. </FONT><I><FONT face=3DPalatino>Amer. =
Zool.</FONT></I><FONT=20
face=3DPalatino> 21: 391 - 399.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Bolker, J. A. and Raff, R. A. 1997. =
</FONT><I><FONT=20
face=3DPalatino>J. N. I. H. Res</FONT></I><FONT face=3DPalatino>. 9: 35 =
-=20
39.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Bowler, P. J. 1996.</FONT><I><FONT =
face=3DPalatino> Life's=20
Splendid Drama</FONT></I><FONT face=3DPalatino>. U. of Chicago Press,=20
Chicago.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Duboule. D. 1994. Temporal colinearity and the=20
phylogenetic progression: a basis for the stability of the vertebrate =
Bauplan=20
and the evolution of morphologies through heterochrony. </FONT><I><FONT=20
face=3DPalatino>Development</FONT></I><FONT face=3DPalatino> (suppl.) =
1994: 135 -=20
142.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Gilbert, S. F. 1997. </FONT><I><FONT=20
face=3DPalatino>Developmental Biology</FONT></I><FONT face=3DPalatino>. =
Fifth ed.=20
Sinauer Associates, Sunderland, MA.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Gould, S. J. 1977. </FONT><I><FONT =
face=3DPalatino>Ontogeny=20
and Phylogeny</FONT></I><FONT face=3DPalatino>. Belknap Press, =
Cambridge,=20
MA</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Haeckel, E. 1874. </FONT><I><FONT=20
face=3DPalatino>Anthropogenie oder Entwickelungsgeschichte des=20
Menschen</FONT></I><FONT face=3DPalatino>. Engelmann, Leipzig.</FONT> =
</P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Hambin, T. J. 1997. Haeckel's drawings. =
</FONT><I><FONT=20
face=3DPalatino>Times</FONT></I><FONT face=3DPalatino> (London), 18 Aug. =

1997.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Hanken, J. 1993. Model systems versus =
outgroups:=20
alternative approaches to the study of head development and evolution.=20
</FONT><I><FONT face=3DPalatino>Amer. Zool</FONT></I><FONT =
face=3DPalatino>. 33: 448=20
- 456.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Nyhart, L K. 1995. </FONT><I><FONT =
face=3DPalatino>Biology=20
takes Form: Animal Morphology and the German Universities=20
1800-1900</FONT></I><FONT face=3DPalatino>. University of Chicago Press, =

Chicago.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Richards, R. J. 1992.</FONT><I><FONT =
face=3DPalatino>The=20
Meaning of Evolution</FONT></I><FONT face=3DPalatino>. U. of Chicago =
Press,=20
Chicago.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Richardson, M. K. 1995. Heterochrony and the =
phylotypic=20
period.</FONT><I><FONT face=3DPalatino> Dev. Biol.</FONT></I><FONT =
face=3DPalatino>=20
172: 412 - 421.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Richardson, M. K., Hanken, J., Gooneratne, M. =
J. Pieau,=20
C., Raynaud, A., Selwood, L., and Wright, G. M. 1997. There is no highly =

conserved embryonic stage in the vertebrates: implications for current =
theories=20
of evolution and development.</FONT><I><FONT face=3DPalatino>=20
Anat.Embryol.</FONT></I><FONT face=3DPalatino> 196: 91 - 106.</FONT> =
</P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Richardson, M. K., Hanken, J., Selwood, L., =
Wright, G.=20
M., Richards, R. J., Pieau, C., and Raynaud, A. 1997b) Haeckel, embryos, =
and=20
evolution. </FONT><I><FONT face=3DPalatino>Science</FONT></I><FONT =
face=3DPalatino>=20
280: 983 -984.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Romanes, G. J. 1901. </FONT><I><FONT =
face=3DPalatino>Darwin=20
and After Darwin</FONT></I><FONT face=3DPalatino>. Open Court, =
london.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Slack, J. M. W., Holland, P. W. H., and Graham, =
C. F.=20
1993. The zootype and the phylotypic stage. </FONT><I><FONT=20
face=3DPalatino>Nature</FONT></I><FONT face=3DPalatino> 361: 490 - =
492.</FONT></P>
<P><FONT face=3DPalatino></FONT>&nbsp;</P>
<P><FONT face=3DPalatino>Wolpert, L. 1991. </FONT><I><FONT =
face=3DPalatino>The=20
Triumph of the Embryo.</FONT></I><FONT face=3DPalatino> Oxford =
University Press,=20
Oxford.</FONT></P>
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