Encyclopedia of Evolution.pdf - Online Reading Center
Encyclopedia of Evolution.pdf - Online Reading Center
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