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Monday, August 01, 2005

Bat development

It always gives a fellow a warm feeling to see an old comrade-in-arms publish a good paper. Chris Cretekos was a graduate student working on the molecular genetics of zebrafish at the University of Utah when I was a post-doc there, and he's a good guy I remember well…so I was glad to see his paper in Developmental Dynamics. But then I notice it wasn't on zebrafish—Apostate! Heretic!

Except…it's on bats. How cool is that? And it's on the embryonic development of bats. Even cooler! I must graciously forgive his defection from the zebrafish universe since he is working on an organism that is weird and fascinating and important.

Here's the abstract:

There are approximately 4,800 extant species of mammals that exhibit tremendous morphological, physiological, and developmental diversity. Yet embryonic development has been studied in only a few mammalian species. Among mammals, bats are second only to rodents with regard to species number and habitat range and are the most abundant mammals in undisturbed tropical regions. Bat development, though, remains relatively unstudied. Here, we describe and illustrate a staging series of embryonic development for the short-tailed fruit bat, Carollia perspicillata, based on embryos collected at timed intervals after captive matings. As Carollia can be readily maintained and propagated in captivity and is extremely abundant in the wild, it offers an attractive choice as a chiropteran model organism. This staging system provides a framework for studying Carollia embryogenesis and should prove useful as a guide for embryological studies of other bat species and for comparisons with other orders of mammals.

What this paper is is meat-and-potatoes embryology—it's a staging series, the kind of paper that documents the pattern of normal development for an organism as a reference. The zebrafish staging series is online, so you can see what one is like; it's a collection of photos and descriptions coupled to a timeline so that everyone has a standard reference point for future studies.

I can't say much about the paper. It consists of tables of stages and dry Latin words and lots and lots of photos of embryos, and it's the kind of droolworthy thing where you just want to look at the pretty pictures. So, here, a few of the figures from the paper.

Of course I have to show you a bat pharyngula. Early development isn't very surprising—it's generic mammalian stuff—but here you can see that common appearance all of us vertebrates have early on, looking like a segmented worm with an odd cluster of protrusions at the head end. The glossopharyngeal arch (ga) is marked, as is the otic vesicle (otv) and forelimb bud (fl).

bat embryo

You can start to see some dramatic differences from the human pattern of development at 60-90 days of gestation. Look at those forelimbs; you can see our five-fingered layout there, but there is extensive webbing and the fingers just grow and grow. I love those first couple of shots. The little guy looks so shy, hiding behind those big hands.

bat embryobat embryobat embryobat embryobat embryo

These are some closer shots of the developing limbs in 54-70 day old embryos.

bat embryo
a, autopod; aer, apical ectodermal ridge; ca, calcar; chp, chiropatagium; cl, claw primordium; dc, digit condensation; fp, foot plate; hp, hand plate; id, interdigit; mc, metacarpal; pl, phalange; plp, plagiopatagium; prp, region of the propatagium primordium; s, stylopod; tm, thumb; urp, uropatagium; z, zeugopod. All panels show the dorsal surface of the right limb with anterior toward top and the proximal at left, views are not to scale.

A lovely piece of work like this makes me want to open up a few pregnant bats…but on the other hand, I also like to keep our local bats happy and thriving and eating mosquitos. At the very least, though, I hope there is more work on bat embryology coming up soon.


Cretekos CJ, Weatherbee SD, Chen C-H, Badwaik NK, Niswander L, Behringer RR, Rasweiler JJ (2005) Embryonic staging system for the short-tailed fruit bat, Carollia perspicillata, a model organism for the mammalian order Chiroptera, based upon timed pregnancies in captive-bred animals. Developmental Dynamics 233(3):721-738.


Trackback url: http://pharyngula.org/index/trackback/2660/jlYtw7Pk/

Comments:
#33359: — 08/01  at  10:16 AM
Bat Boy: The Musical.



#33361: — 08/01  at  10:23 AM
That is totally cool! I can't wait to see these on a poster for some metal band.
One of the morphological clues for staging midgestation mouse embryos is the loss of the webbing between the forelimb digits (compare figures L and N for the hindlimb). I guess bats, somewhere in their evolution, skipped that developmental step.



#33371: bitchphd — 08/01  at  11:23 AM
Cool! I'll have to show PK these pictures.



#33373: dr. dave — 08/01  at  11:26 AM
Crap, somebody beat me to the bat-boy comment. Either way... I can't wait until the Weekly World News gets hold of THESE pics.



#33380: GrrlScientist — 08/01  at  12:20 PM
They are so cute, they look rather like little people.



#33387: — 08/01  at  01:13 PM
Very cute. Though the last of the group of 5 does look rather like it's turned itself into a crumpled bag of bones.



#33394: — 08/01  at  01:40 PM
I'm not sure what's the best place to look for comparative embryology, but I'd collected a lot of pages in this article. (the ' in "Haeckel's" disappears from this URL when I try to access it; you may have to put it in by hand)

In particular, note human vs. mouse embryology; the two embryos look very simular until the hindlimb buds start to grow, at which point the human one's tail starts to go away and the mouse one's tail starts to lengthen -- and its face starts to grow a snout. The bat embryos discussed here have a similar pattern of similarity and divergence, it would seem.



#33399: — 08/01  at  01:59 PM
Those pictures are cool beyond words! In between being some horrific little mammalian lamprey and a distended sack of wing, the little chap's adorable. I can't wait until this work starts getting into the nitty gritty of bat evolution.

-Schmitt.



#33410: arensb — 08/01  at  03:25 PM
Let's hear it for Hox clusters!



#33434: — 08/01  at  06:08 PM
Loren, Good page and links, but an odd word or two, as 'forgery'. Not quite a synonym for 'inaccurate'.
Should the topic be presented without citing


Haeckel's ABC of evolution and development
MICHAEL K. RICHARDSON and GERHARD KEUCK

Biol. Rev. (2002), 77, pp. 495-528 ?


V. CONCLUSIONS
(1) Zoologist Ernst Haeckel is widely known for
his in¯uential popular science books, magni®cent
zoological monographs, and scienti®c innovations.
He attempted to integrate taxonomy and embryology
into the Darwinian framework and to use the
data for phylogeny reconstruction. His work is
historically and scienti®cally important, and has
in¯uenced modern thinking in evolutionary developmental
biology and phylogenetics.
(2) Haeckel's Biogenetic Law makes predictions
about the rank of primitive and advanced characters
in developmental sequences. To Haeckel, heterochrony
was a change in developmental sequence.
Some modern methods in comparative embryology,
such as event-pairing, also exploit sequence heterochronies.
Haeckel's `alphabetical analogy' shows

522 Michael K. Richardson and Gerhard Keuck

characters or stages (it is not always clear which)
dropping out of a sequence, being added terminally,
or undergoing transformation. Signi®cantly,
Haeckel's caenogenetic changes were all adaptations
to embryonic life ; he does not acknowledge the
possibility that embryos could be affected by
selection acting on adult characters. Haeckel believed
that primitive characters alone are important
in phylogeny reconstruction, in contrast to modern
cladistic methodologies which employ synapomorphies.
Haeckel is often accused of advocating absurd
recapitulatory scenarios ± such as ®sh gills in human
embryos. However, we ®nd that he explicitly rejected
this scenario in some of his writings.
(3) Modern views on Haeckel are, typically,
ambivalent. His early work is praised, but there is
confusion about the Biogenetic Law and recapitulation
(the latter being very often confused with
embryonic resemblance). Some of this confusion can
be blamed on ambiguities and logical ¯aws in
Haeckel's writing. Several modern studies support
the Biogenetic Law in the case of single character
transformations. However, there is no evidence from
vertebrates that entire stages are recapitulated.
Haeckelian and von Baerian models both make the
same prediction: that plesiomorphies are transformed
into apomorphies during ontogeny. The
principle differences between the two models are
that Haeckel's scheme involves heterochrony as one
of its mechanisms, and leads to a series of conserved
stages.
(4) Haeckel's embryo drawings are important
as phylogenetic hypotheses, teaching aids ± even
scienti®c evidence. They re¯ect a phenetic, nonquantitative,
` portrait gallery ' approach to comparative
embryology. This approach is still common
today, although quantitative comparative methodologies
are gaining ground. The drawings illustrate
embryonic similarity, recapitulation, and phenotypic
divergence. They have been criticized ± not always
fairly ± on the grounds of inaccuracy and tendentiousness.
We have identi®ed potential sources for
several of the drawings, and ®nd some evidence of
doctoring. In opposition to Haeckel and his drawings,
Wilhelm His proposed a rational, morphometric
approach for comparing embryos. His
failed, however, because he overlooked the importance
of rigorous developmental staging.

/Conclusions

needlefishes



Trackback: Bat Begins Tracked on: PhaWRONGula (72.9.234.70) at 2005 08 01 21:43:06
I wanted to ask Chris Cretekos about his bat development paper, but I must have selected the wrong number in my phone book. I got Chris Nolan's response instead...



#33457: John Wilkins — 08/01  at  10:23 PM
One reason why this is important is that one in four mammal species is a bat. For too long our verspertilian comrades have suffered under the indignities of the ungulate oppressors!

John S. Wilkins : evolvethought.blogspot.com



#33460: — 08/01  at  10:54 PM
Scott Weatherbee just presented a talk on some of this work at the Society for Developmental Biology meeting this past weekend. Though, actually, it was more about the developmental basis allowing the maintenance of webbing between the digits of forewings (which is normally eliminated by programmed cell death). Very cool stuff!



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