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recent papers on digit homology



 don't recall seeing these papers being cited on the DML (might be wrong 
though, especially about the last one, which is from the january issue).

J Exp Zoolog B Mol Dev Evol. 2005 May 15;304(3)

"Hox genes, digit identities and the theropod/bird transition" 
Frietson Galis 1 *, Martin Kundrát 2, Johan A.J. Metz 1 3 
ABSTRACT
Vargas and Fallon (2005. J Exp Zool (Mol Dev Evol) 304B:86-90) propose that Hox 
gene expression patterns indicate that the most anterior digit in bird wings is 
homologous to digit 1 rather than to digit 2 in other amniotes. This 
interpretation is based on the presence of Hoxd13 expression in combination 
with the absence of Hoxd12 expression in the second digit condensation from 
which this digit develops (the first condensation is transiently present). This 
is a pattern that is similar to that in the developing digit 1 of the chicken 
foot and the mouse hand and foot. They have tested this new hypothesis by 
analysing Hoxd12 and Hoxd13 expression patterns in two polydactylous chicken 
mutants, Silkie and talpid2. They conclude that the data support the notion 
that the most anterior remaining digit of the bird wing is homologous to digit 
1 in other amniotes either in a standard phylogenetic sense, or alternatively 
in a (limited) developmental sense in agreement with the Frameshift
 Hypothesis of Wagner and Gautier (1999, i.e., that the developmental pathway 
is homologous to the one that leads to a digit 1 identity in other amniotes, 
although it occurs in the second instead of the first digit condensation). We 
argue that the Hoxd12 and Hoxd13 expression patterns found for these and other 
limb mutants do not allow distinguishing between the hypothesis of Vargas and 
Fallon (2005. J Exp Zool (Mol Dev Evol) 304B:86-90) and the alternative one, 
i.e., the most anterior digit in bird wings is homologous to digit 2 in other 
amniotes, in a phylogenetic or developmental sense. Therefore, at the moment 
the data on limb mutants does not present a challenge to the hypothesis, based 
on other developmental data (Holmgren, 1955. Acta Zool 36:243-328; Hinchliffe, 
1984. In: Hecht M, Ostrom JH, Viohl G, Wellnhofer P, editors. The beginnings of 
birds. Eichstätt: Freunde des Jura-Museum. p 141-147; Burke and Feduccia, 1997. 
Science 278:666-668; Kundrát et al., 2002. J Exp Zool (Mol
 Dev Evol) 294B:151-159; Larsson and Wagner, 2002. !
J Exp Zool (Mol Dev Evol) 294B:146-151; Feduccia and Nowicki, 2002. 
Naturwissenschaften 89:391-393), that the digits of bird wings are homologous 
to digits 2,3,4 in amniotes. We recommend further testing of the hypothesis by 
comparing Hoxd expression patterns in different taxa. J. Exp. Zool. (Mol. Dev. 
Evol.) 304B:198-205. 2005


"The digits of the wing of birds are 1, 2, and 3. a review "
Alexander O. Vargas 1 2 *, John F. Fallon 3 
ABSTRACT
Fossil evidence documenting the evolutionary transition from theropod dinosaurs 
to birds indicates unambiguously that the digits of the wing of birds are 
digits 1, 2, and 3. However, some embryological evidence suggests that these 
digits are 2, 3, and 4. This apparent lack of correspondence has been described 
as the greatest challenge to the widely accepted theropod-bird link (Zhou 2004. 
Naturwissenschaften 91:455-471). Here we review the pertinent literature 
regarding the debate on the origin of birds and wing digital identity and the 
evidence in favor of a 1, 2, 3 identity of the wing digits. Recent molecular 
evidence shows that the expression of Hoxd12 and Hoxd13 in the developing wing 
supports the theropod-bird link. In the chicken foot and in the mouse hand and 
foot, digit 1 is the only digit to combine the expression of Hoxd13 with the 
absence of expression of Hoxd12. The same is observed in the anterior digit of 
the wing, suggesting it is a digit 1, as expected for a theropod.
 Nevertheless, Galis et al. (2005. J Exp Zool (Mol Dev Evol) in press), argue 
that Hoxd12 and Hoxd13 expression patterns in mutant limbs do not allow 
distinguishing the most anterior digit in the bird wing from digit 2. They also 
argue that constraints to the evolution of limb development support the 2, 3, 4 
identity of the wing digits. However, the case put forward by Galis et al. is 
biased and flawed with regard to interpretation of mutant limbs, developmental 
mechanisms, stages observed, and the description of the evolutionary variation 
of limb development. Importantly, Galis et al. do not present evidence from 
wild-type limbs that counters the conclusions of Vargas and Fallon (2005. J Exp 
Zool (Mol Dev Evol) 304B(1):85-89), and fail to provide molecular evidence to 
specifically support the hypothesis that the wing digits are 2, 3, and 4. The 
expression of Hoxd12 and Hoxd13 in the developing wing is consistent with the 
hypothesis that birds are living dinosaurs; this view can lead
 to a greater understanding of the actual l!
imits to the evolutionary variation of limb development. J. Exp. Zool. (Mol. 
Dev. Evol.) 304B:206-219, 2005

J Exp Zoolog B Mol Dev Evol. 2005 Jan 15;304(1)

"Birds have dinosaur wings: The molecular evidence.
Vargas AO, Fallon JF".

ABSTRACT
Within developmental biology, the digits of the wing of birds are considered on 
embryological grounds to be digits 2, 3 and 4. In contrast, within 
paleontology, wing digits are named 1, 2, 3 as a result of phylogenetic 
analysis of fossil taxa indicating that birds descended from theropod dinosaurs 
that had lost digits 4 and 5. It has been argued that the development of the 
wing does not support the conclusion that birds are theropods, and that birds 
must have descended from ancestors that had lost digits 1 and 5. Here we use 
highly conserved gene expression patterns in the developing limbs of mouse and 
chicken, including the chicken talpid(2)mutant and polydactylous Silkie breed 
(Silkie mutant), to aid the assessment of digital identity in the wing. Digit 1 
in developing limbs does not express Hoxd12, but expresses Hoxd13. All other 
digits express both Hoxd12and Hoxd13. We found this signature expression 
pattern identifies the anteriormost digit of the wing as digit 1, in accordance
 with the hypothesis these digits are 1, 2 and 3, as in theropod dinosaurs. Our 
evidence contradicts the long-standing argument that the development of the 
wing does not support the hypothesis that birds are living dinosaurs.



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