I’m going to introduce you to either a fascinating question or a throbbing headache in evolution, depending on how interested you are in peculiar details of arthropod anatomy (Mrs Tilton may have just perked up, but the rest of you may resume napping). The issue is tagmosis.
I just learned (via John Lynch) about a paper on cetacean limbs that combines developmental biology and paleontology, and makes a lovely argument about the mechanisms behind the evolution of whale morphology. It is an analysis of the molecular determinants of limb formation in modern dolphins, coupled to a comparison of fossil whale limbs, and a reasonable inference about the pattern of change that was responsible for their evolution.
One important point I’d like to make is that even though what we see in the morphology is a pattern of loss—whale hindlimbs show a historical progression over tens of millions of years of steady loss, followed by a near-complete disappearance—the molecular story is very different. The main players in limb formation, the genes Sonic hedgehog (Shh), the Fgfs, and the transcription factor Hand2, are all still present and fully functional in these animals. What has happened, though, is that there have been novel changes to their regulation. Even loss of structures is a consequence of changes and additions to regulatory pathways.
I’m going to briefly summarize an interesting new article on cnidarian Hox genes…unfortunately, it requires a bit of background to put it in context, so bear with me for a moment.
First you need to understand what Hox genes are. They are transcription factors that use a particular DNA binding motif (called a homeobox), and they are found in clusters and expressed colinearly. What that means is that you find the Hox genes that are essential for specifying positional information along the length of the body in a group on a chromosome, and they are organized in order on the chromosome in the same order that they are turned on from front to back along the body axis. Hox genes are not the only genes that are important in this process, of course; animals also use another class of regulatory genes, the Wnt genes, to regulate development, for instance.
A gene can only be called a Hox gene sensu stricto if it has a homeobox sequence, is homologous to other known Hox genes, and is organized in a colinear cluster. If such a gene is not in a cluster, it is demoted and called simply a Hox-like gene.
Hox genes originated early in animal evolution. Genes containing a homeobox are older still, and are found in plants and animals, but the particular genes of the Hox system are unique to multicellular animals, and that key organization arrangement of the set of Hox genes in a cluster is more unique still. The question is exactly when the clusters arose, shortly after or sometime before the diversification of animals.
If you take a look at animal phylogeny, an important group are the diploblastic phyla, the cnidarians and ctenophores. They branched off early from the metazoan lineage, and they possess some sophisticated patterns of differentiation along the body axis. We know they have homeobox containing genes that are related to the ones used in patterning the bodies of us vertebrates, but are they organized in the same way? Did the cnidaria have Hox clusters, suggesting that the clustered Hox genes were a very early event in evolution, or do they lack them and therefore evolved an independent set of mechanisms for specifying positional information along the body axis?
Four of my favorite things are development, evolution, and breasts, and now I have an article that ties them all together in one pretty package. It’s a speculative story at this point, but the weight of the evidence marshaled in support of the premise is impressive: the mammalian breast first evolved as an immunoprotective gland that produced bacteriocidal secretions to protect the skin and secondarily eggs and infants, and that lactation is a highly derived kind of inflammation response. That mammary glands may have had their origin as inflamed glands suppurating mucus may not be the most romantic image to arise in a scientific study, but really—they got better and better over the years.
I’m going to link to a post on Uncommon Descent. I try to avoid that, because I think it is a vile harbor of malign idiocy, but Dembski has just put up something that I think is merely sincerely ignorant. That’s worth correcting. It also highlights the deficiencies of Dembski’s understanding of biology.
Dembski makes a strange argument for ID on the basis of a certain class of experiments in developmental biology.
Crap. Coturnix tagged me with this beautiful bird meme, and I am the wrong person to ask. I don’t get out much, preferring to sit in the lab or the library, so my favorite birds are all in pieces and dead. But OK, since he asked…
Bird digits
Bird teeth
Bird brains
Jurassic bird brains
Bird lungs
Oviraptor pelves (does that count?)
Cretaceous bird embryos
Four-winged birds
Waimanu
And Archaeopteryx, of course.
The Wnt genes produce signalling proteins that play important roles in early development, regulating cell proliferation, differentiation and migration. It’s hugely important, used in everything from early axis specification in the embryo to fine-tuning axon pathfinding in the nervous system. The way they work is that the Wnt proteins are secreted by cells, and they then bind to receptors on other cells (one receptor is named Frizzled, and others are LRP-5 and 6), which then, by a chain of cytoplasmic signalling events, removes β-catenin from a degradation pathway and promotes its import into the nucleus, where it can modify patterns of gene expression. This cascade can also interact with the cytoskeleton and trigger changes in cell migration and cell adhesion. The diagram below illustrates the molecular aspects of its function.
One of the hallmark characters of animals is the presence of a specific cluster of genes that are responsible for staking out the spatial domains of the body plan along the longitudinal axis. These are the Hox genes; they are recognizable by virtue of the presence of a 60 amino acid long DNA binding region called the homeodomain, by similarities in sequence, by their role as regulatory genes expressed early in development, by the restriction of their expression to bands of tissue, by their clustering in the genome to a single location, and by the remarkable collinearity of their organization on the chromosome to their pattern of expression: the order of the gene’s position in the cluster is related to their region of expression along the length of the animal. That order has been retained in most animals (there are interesting exceptions), and has been conserved for about a billion years.
Think about that. While gene sequences have steadily changed, while chromosomes have been fractured and fused repeatedly, while differences accumulated to create forms as different as people and fruit flies and squid and sea urchins, while continents have ping-ponged about the globe and meteors have smashed into the earth and glaciers have advanced and retreated, these properties of this set of genes have remained constant. They are fundamental and crucial to basic elements of our body plan, so basic that we take them completely for granted. They determine that we can have different regions of our bodies with different organs and organization. Where did they come from and what forces constrain them to maintain their specific organization on the chromosome? Are there other genes that are comparably central to our organization?
There are quite a few genes that are known to be highly conserved in both sequence and function in animals. Among these are the various Hox genes, which are expressed in an ordered pattern along the length of the organism and which define positional information along the anterior-posterior axis; and another is decapentaplegic (dpp) which is one of several conserved genes that define the dorsal-ventral axis. Together, these sets of genes establish the front-back and top-bottom axes of the animal, which in turn establishes bilaterality—this specifically laid out three-dimensional organization is a hallmark of the lineage Bilateria, to which we and 99% of all the other modern animal species belong.
There are some animals that don’t belong to the Bilateria, though: members of the phylum Cnidaria, the jellyfish, hydra, sea anemones, and corals, which are typically radially symmetric. A few cnidarian species exhibit bilateral symmetry, though, and Finnerty et al. (2004) ask a simple question: have those few species secondarily reinvented a mechanism for generating bilateral symmetry (so that this would be an example of convergent evolution), or do they use homologous mechanisms, that is, the combination of Hox genes for A-P patterning and dpp for D-V patterning? The answer is that this is almost certainly an example of homology—the same genes are being used.
In previous articles about fly development, I’d gone from the maternal gradient to genes that are expressed in alternating stripes (pair-rule genes), and mentioned some genes (the segment polarity genes) that are expressed in every segment. The end result is the development of a segmented animal: one made up of a repeated series of morphological modules, all the same.
Building an animal with repeated elements like that is a wonderfully versatile strategy for making an organism larger without making it too much more complicated, but it’s not the whole story. Just repeating the same bits over and over again is a way to make a generic wormlike thing—a tapeworm, for instance—but even tapeworms may need to specialize certain individual segments for specific functions. At its simplest, it may be necessary to modify one end for feeding, and the opposite end for mating. So now, in addition to staking out the tissues of the embryo as belonging to discrete segments, we also need a mechanism that says “build mouthparts here (and not everywhere)”, and “put genitalia here (not over there)”.
