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Encyclopedia of Evolution.pdf - Online Reading Center

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0 Devonian period<br />

gene from an invertebrate chordate amphioxus was inserted<br />

into its DNA.<br />

Hox genes do not control the structure <strong>of</strong> the body parts<br />

but makes them develop in their proper places. One example<br />

<strong>of</strong> this is the Hox gene Antp. This gene normally controls the<br />

development <strong>of</strong> legs in the thorax <strong>of</strong> flies. A mutation in this<br />

gene can cause legs to grow out <strong>of</strong> the fly’s head, where the<br />

antennae would have been, a mutation called antennapedia<br />

(antenna, plus Greek pedia for foot). A missing Hox gene<br />

product causes a structure, such as a leg, to misunderstand its<br />

location in the developing animal. The normal gene, like the<br />

mutation, does not affect the structure <strong>of</strong> the legs but their<br />

location.<br />

Homeotic genes other than Hox genes are essential to<br />

the order <strong>of</strong> development in animal bodies. For example, the<br />

sog gene in flies promotes the development <strong>of</strong> a nerve cord<br />

along the bottom or front (ventral) surface, while the chordin<br />

(chd) gene in vertebrates promotes the development <strong>of</strong> a<br />

nerve cord along the top or back (dorsal) surface. Molecular<br />

biologists E. M. DeRobertis and Y. Sasai reported in 1996<br />

that these two genes were derived from a common ancestor.<br />

Experiments have confirmed this. If RNA transcripts from<br />

either a fly sog gene or a frog chd gene are injected into an<br />

embryo, they induce the normal development <strong>of</strong> the nerve<br />

cord: on the ventral surface in flies, on the dorsal surface in<br />

frogs. Thus, it appears, the embryonic development <strong>of</strong> arthropods<br />

is inverted with respect to that <strong>of</strong> vertebrates. This was<br />

the first scientific confirmation <strong>of</strong> the theory presented by<br />

Ge<strong>of</strong>froy St.-Hilaire in the early 19th century that vertebrates<br />

developed as upside-down arthropods (see invertebrates,<br />

evolution <strong>of</strong>; Cuvier, Georges).<br />

Another homeotic gene (not a Hox gene) in flies is the<br />

hedgehog gene. This gene got its name from a mutant form, in<br />

which the fly was covered with prickles. The normal function<br />

<strong>of</strong> the hedgehog gene is to control the front-to-back order <strong>of</strong><br />

development within each <strong>of</strong> the segments defined by the Hox<br />

genes. When this gene was also discovered to be present in<br />

vertebrates, it was given the name sonic hedgehog (shh). The<br />

shh gene is involved in the development <strong>of</strong> vertebrate limbs.<br />

It is expressed for a brief period in the development <strong>of</strong> fish<br />

fins; its prolonged expression allows the development <strong>of</strong> legs<br />

and feet in four-legged vertebrates (see amphibians, evolution<br />

<strong>of</strong>). Within each <strong>of</strong> the vertebrate limbs, the genes are<br />

expressed in order.<br />

Other homeotic genes (not Hox genes) found widely<br />

among animal genomes are the pax and tinman genes. The<br />

first gene stimulates the development <strong>of</strong> eyes. The structure <strong>of</strong><br />

the eye is very different in flies and vertebrates, but both are<br />

stimulated by versions <strong>of</strong> the same pax gene inherited from<br />

an ancestor that may have had simple eye spots. The tinman<br />

gene specifies the development <strong>of</strong> a heart, although the form<br />

<strong>of</strong> the heart is different in each animal.<br />

Most <strong>of</strong> the studies <strong>of</strong> MADS genes in plants have<br />

involved the development <strong>of</strong> flowers from buds. Plants<br />

retain embryonic tissues in their buds, which are free to<br />

develop in different ways, like the stem cells <strong>of</strong> animals. A<br />

bud resembles an embryo in its developmental possibilities.<br />

Different MADS genes influence the development <strong>of</strong> differ-<br />

ent flower parts: the sepals on the outside, the petals inside<br />

<strong>of</strong> them, then the male stamens, and the female carpels<br />

in the middle (see angiosperms, evolution <strong>of</strong>). Mutations<br />

in the MADS genes can cause unusual developmental<br />

patterns, such as the development <strong>of</strong> sepals and petals<br />

where stamens and carpels would normally develop, or vice<br />

versa. A single mutation <strong>of</strong> a homeotic gene in a flowering<br />

plant can change the entire shape <strong>of</strong> the flower, which will<br />

change its relationship with pollinators (see coevolution).<br />

For example, a single mutation changes snapdragon-like<br />

flowers (Linaria vulgaris), which are bilaterally symmetrical,<br />

into the Peloria version, which is radially symmetrical.<br />

Genes similar to the MADS genes can be found in plants<br />

related to the ancestors <strong>of</strong> the angiosperms (see gymnosperms,<br />

evolution <strong>of</strong>), even though these plants do not<br />

have flowers.<br />

Some biologists, such as the botanist Sonia Sultan, have<br />

pointed out that it is not only necessary to understand the<br />

genetic processes <strong>of</strong> development but also to understand the<br />

ecological consequences <strong>of</strong> development (such as phenotypic<br />

plasticity; see adaptation), in order to understand the process<br />

<strong>of</strong> evolution. This approach has been called “eco-devo.”<br />

Further <strong>Reading</strong><br />

Amundson, Ron. The Changing Role <strong>of</strong> the Embryo in <strong>Evolution</strong>ary<br />

Thought: The Roots <strong>of</strong> Evo-Devo. Cambridge, U.K.: Cambridge<br />

University Press, 2005.<br />

Carroll, Sean B. Endless Forms Most Beautiful: The New Science <strong>of</strong><br />

Evo Devo and the Making <strong>of</strong> the Animal Kingdom. New York:<br />

Norton, 2005.<br />

Carroll, Sean B. The Making <strong>of</strong> the Fittest: DNA and the Ultimate<br />

Forensic Record <strong>of</strong> <strong>Evolution</strong>. New York: Norton, 2006.<br />

Coen, Enrico. The Art <strong>of</strong> Genes: How Organisms Make Themselves.<br />

New York: Oxford University Press, 1999.<br />

Davidson, Eric H., and Douglas H. Erwin. “Gene regulatory networks<br />

and the evolution <strong>of</strong> animal body plans.” Science 311<br />

(2006): 796–800.<br />

DeRobertis, E. M., and Y. Sasai. “A common plan for dorsiventral<br />

patterning in bilateria.” Nature 380 (1996): 37–40.<br />

Freeman, Scott, and John C. Herron. “Development and evolution.”<br />

Chap. 18 in <strong>Evolution</strong>ary Analysis. 3rd ed. Upper Saddle River,<br />

N.J.: Pearson Prentice Hall, 2004.<br />

Maynard Smith, John. Shaping Life: Genes, Embryos, and <strong>Evolution</strong>.<br />

New Haven, Conn.: Yale University Press, 1998.<br />

Shubin, Neal, C. Tabin, and Sean B. Carroll. “Fossils, genes, and the<br />

evolution <strong>of</strong> animal limbs.” Nature 388 (1997): 639–648.<br />

Zimmer, Carl. “A fin is a limb is a wing: How evolution fashioned its<br />

masterworks.” National Geographic, November 2006, 110–135.<br />

Devonian period The Devonian period (410 to 360 million<br />

years ago) was the fourth period <strong>of</strong> the Paleozoic era<br />

(see geological time scale). The first significant advances<br />

<strong>of</strong> life onto land had occurred during the preceding Silurian<br />

period. During the Devonian period, plants filled the wet<br />

areas <strong>of</strong> the Earth, making large areas <strong>of</strong> the Earth’s surface<br />

green for the first time.<br />

Climate. The climate in the equatorial regions was warm<br />

and wet. Portions <strong>of</strong> the southern continent, over the South

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