Bilateral symmetry in a sea anemone



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.

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A brief overview of Hox genes


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)”.

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Paul Nelson has been twittering about ORFans for some time now—he seems to precede his talks by threatening to make us evolutionists tremble in our boots by bringing them up, but he never seems to follow through. Ian Musgrave got tired of waiting for him to give us a coherent creationist argument about them, and has gone ahead and cut him off at the knees by explaining the place of ORFans in evolution.

In case you’re baffled by the jargon, “ORF” is an Open Reading Frame, or a stretch of DNA bracketed by a start and stop codon; it’s a kind of bare minimum criterion for recognizing an actual gene within a DNA sequence. An ORFan is an orphan ORF sequence, or one that doesn’t have a known function or affinity to other known genes. It is not surprising that genes exist that do not have an easily recognized homology with other genes—novel genes have to arise sometime, and we do not have a complete understanding of all sequences of all organisms.

The short answer is that Nelson is deluded, and ORFans do not conflict with evolution at all…but read Ian’s post for all the details.

Evo-Devo in NYR Books!

This really is an excellent review of three books in the field of evo-devo

From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design (amzn/b&n/abe/pwll),

Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom (amzn/b&n/abe/pwll), and

The Plausibility of Life:Resolving Darwin’s Dilemma (amzn/b&n/abe/pwll)—all highly recommended by me and the NY Times. The nice thing about this review, too, is that it gives a short summary of the field and its growing importance.

Evolving spots, again and again

a–c, The wing spots on male flies of the Drosophila genus. Drosophila tristis (a) and D. elegans (b) have wing spots that have arisen during convergent evolution. Drosophila gunungcola (c) instead evolved from a spotted ancestor. d, Males wave their wings to display the spots during elaborate courtship dances.

It’s all about style. When you’re out and about looking for mates, what tends to draw the eye first are general signals—health and vigor, symmetry, absence of blemishes or injuries, that sort of thing—but then we also look for that special something, that je ne sais quoi, that dash of character and fashionable uniqueness. In humans, we see the pursuit of that elusive element in shifting fashions: hairstyles, clothing, and makeup change season by season in our efforts to stand out and catch the eye in subtle ways that do not distract from the more important signals of beauty and health.

Flies do the same thing, exhibiting genetic traits that draw the attention of the opposite sex, and while nowhere near as flighty as the foibles of human fashion, they do exhibit considerable variability. Changes in body pigmentation, courtship rituals, and pheromones are all affected by sexual selection, but one odd feature in particular is the presence of spots on the wing. Flies flash and vibrate their wings at prospective mates, so the presence or absence of wing spots can be a distinctive species-specific element in their evolution. One curious thing is that wing spots seem to be easy to lose and gain in a fly lineage, and species independently generate very similar pigment spots. What is it about these patterns that makes them simultaneously labile and frequently re-expressed?

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Evolving spots


Here’s what seems to be a relatively simple problem in evolution. Within the Drosophila genus (and in diverse insects in general), species have evolved patterned spots on their wings, which seem to be important in species-specific courtship. Gompel et al. have been exploring in depth one particular problem, illustrated below: how did a spot-free ancestral fly species acquire that distinctive dark patch near the front tip of the wing in Drosophila biarmipes? Their answer involves dissecting the molecular regulators of pattern in the fly wing, doing comparative sequence analyses and identifying the specific stretches of DNA involved in turning on the pigment pattern, and testing their models experimentally by expressing novel gene constructs in different species of flies.

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How to make a bat

The relative length of bat forelimb digits has not changed in 50
million years. (a) Icaronycteris index, which is a 50-million-year-old bat fossil. (b) Extant adult
bat skeleton. The metacarpals (red arrows) of the first fossil bats are already
elongated and closely resemble modern bats. This observation is confirmed by
morphometric analysis of bat forelimb skeletal elements.

or•gan•ic | ôr’ganik | adjective. denoting a relation between elements of something such that they fit together harmoniously as necessary parts of a whole; characterized by continuous or natural development.

One of the wonderful things about how development works is that organisms function as wholes, and changes in one property trivially induce concordant changes in other properties. Tug on one element, changing it’s orientation or size, and during embryogenesis any adjacent elements make compensatory adjustments, so that the resultant form flows, fits, and looks organic. This isn’t that surprising a feature of development, though, unless you have the mistaken idea that the genome encodes a blueprint of morphology. It doesn’t; what it contains is a description of interacting agents that work together in a process to produce a complex result. Changes in genes and regulatory elements can essentially produce changes in rules of development, rather than crudely specifying blocks of morphology.

What does this mean for evolution? It means that subtle changes to the rules of development can be caused by small changes to genes (and especially, to regulatory regions of genes), and that the resulting morphological changes may be dramatic, but are still integrated organically into the form of the organism as a whole. Our understanding of how development works is making it clear that large scale macroevolutionary change may be much easier than we had thought.

Here’s an example where this insight is clarifying the evolution of an organism: the fossil record of bats shows an abrupt appearance of fairly sophisticated creatures with elongated digits, clearly capable of gliding or powered flight, with no known intermediates. We expect there were less fully flight-ready predecessors, but fossil preservation is not kind to small, delicate boned animals. It’s also possible that the transitional period was fairly brief; it looks like turning a paw into a long-fingered membranous wing may be a fairly simple change on a molecular level.

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Chance and regularity in the development of the fly eye


What has always attracted me to developmental biology is the ability to see the unfolding of pattern—simplicity becomes complexity in a process made up of small steps, comprehensible physical and chemical interactions that build a series of states leading to a mostly robust conclusion. It’s a bit like Conway’s Game of Life in reverse, where we see the patterns and can manipulate them to some degree, but we don’t know the underlying rules, and that’s our job—to puzzle out how it all works.


Another fascinating aspect of development is that all the intricate, precise steps are carried out without agency: everything is explained and explainable in terms of local, autonomous interactions. Genes are switched on in response to activation by proteins not conscious action, domains of expression are refined without an interfering hand nudging them along towards a defined goal. It’s teleonomy, not teleology. We see gorgeously regular structures like the insect compound eye to the right arise out of a smear of cells, and there is no magic involved—it’s wonderfully empowering. We don’t throw up our hands and declare a miracle, but instead science gives us the tools to look deeper and work out (with much effort, admittedly) how seeming miracles occur.

One more compelling aspect of development: it’s reliable, but not rigid. Rather than being simply deterministic, development is built up on stochastic processes—ultimately, it’s all chemistry, and cells changing their states are simply ping-ponging through a field of potential interactions to arrive at an equilibrium state probabilistically. When I’d peel open a grasshopper embryo and look at its ganglia, I’d have an excellent idea of what cells I’d find there, and what they’d be doing…but the fine details would vary every time. I can watch a string of neural crest cells in a zebrafish crawl out of the dorsal midline and stream over generally predictable paths to their destinations, but the actions of an individual melanocyte, for instance, are variable and beautiful to see. We developmental biologists get the best of all situations, a generally predictable pattern coupled to and generated by diversity and variation.

One of the best known examples of chance and regularity in development is the compound eye of insects, shown above, which is as lovely and crystalline as a snowflake, yet is visibly assembled from an apparently homogenous field of cells in the embryo. And looking closer, we discover a combination of very tight precision sprinkled with random variation.

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Lamprey skeletons

Bone is a sophisticated substance, much more than just a rock-like mineral in an interesting shape. It’s a living tissue, invested with cells dedicated to continually remodeling the mineral matrix. That matrix is also an intricate material, threaded with fibers of a protein, type II collagen, that give it a much greater toughness—it’s like fiberglass, a relatively brittle substance given resilience and strength with tough threads woven within it. Bone is also significantly linked to cartilage, both in development and evolution, with earlier forms having a cartilaginous skeleton that is replaced by bone. In us vertebrates, cartilage also contains threads of collagen running through it.

These three elements—collagen, cartilage, and bone—present an interesting evolutionary puzzle. Collagen is common to the matrices of both vertebrate cartilage and bone, yet the most primitive fishes, the jawless lampreys and hagfish, have been reported to lack that particular form of collagen, suggesting that the collagen fibers are a derived innovation in chordate history. New work, though, has shown that there’s a mistaken assumption in there: lampreys do have type II collagen! This discovery clarifies our understanding of the evolution of the chordate skeleton.

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Chicken, archosaur…same difference


My daughter is learning about evolution in high school right now, and the problem isn’t with the instructor, who is fine, but her peers, who complain that they don’t see the connections. She mentioned specifically yesterday that the teacher had shown a cladogram of the relationships between crocodilians, birds, and mammals, and that a number of students insisted that there was no similarity between a bird and an alligator.

I may have to send this news article to school with her: investigators have found that a mutation in chickens causes them to develop teeth—and the teeth resemble those of the common ancestor of alligators and chickens, an archosaur.

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