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 hybridization<br />
Three species <strong>of</strong> mustard (Brassica carinata, B. juncea, B. napus) have<br />
arisen as hybrids between B. oleracea, B. nigra, and B. rapa. Boxes<br />
represent original species; arrows represent hybrid species.<br />
• The sunflowers Helianthus deserticola and H. anomalus<br />
have been shown to have hybrid origins. Botanist Loren<br />
Rieseberg has experimentally replicated the hybrid origin<br />
<strong>of</strong> H. anomalus in the greenhouse. By crossing H. annuus<br />
with H. petiolaris, Rieseberg produced three independent<br />
hybrid lines. Only four generations were required to select<br />
vigorous hybrids. The genetic markers (see quantitative<br />
trait loci) <strong>of</strong> the greenhouse hybrids closely resemble<br />
those in the wild populations <strong>of</strong> H. anomalus.<br />
In plants, intergeneric hybrids (while still rare) are more<br />
common than in animals. Russian botanist Georgi Karpechenko<br />
crossed the radish Raphanus sativus with the mustard<br />
Brassica oleracea, producing a Raphanobrassica hybrid,<br />
in 1927. Unfortunately, the hybrid had the leaves <strong>of</strong> radish<br />
and the roots <strong>of</strong> cabbage. Wild Raphanobrassica hybrids<br />
have been reported from California. Triticale, a hybrid <strong>of</strong><br />
wheat and rye, is reported to have the high yield <strong>of</strong> wheat<br />
and the hardiness <strong>of</strong> rye but is rarely grown on a commercial<br />
scale. Hybrids between plant genera are known from<br />
orchids and cypresses as well (see table on page 203). Among<br />
orchids, hybrids are possible among three or even more genera,<br />
although this almost always occurs under artificial conditions.<br />
In natural habitats, pollinator specificity usually<br />
prevents extensive crossings among orchids. For example,<br />
Brassolaeliocattleya is a cross among the genera Brassavola,<br />
Laelia, and Cattleya. When a scientific name is assigned to an<br />
intergeneric hybrid, an × may precede the generic name.<br />
Natural hybrids tend to be rare, but hybrid plants<br />
produced for horticultural purposes can be vigorous. The<br />
reduced reproductive capacity <strong>of</strong> the horticultural hybrid is<br />
not a drawback because artificial propagation is available.<br />
Perhaps the plant hybridizations that are most famous in<br />
the history <strong>of</strong> the world are the crosses that produced modern<br />
wheat. The wild wheat Triticum boeoticum had chromosomes<br />
in pairs (diploid, 14 chromosomes) and was bred by ancient<br />
farmers into the cultivated einkorn wheat Triticum monococcum.<br />
However, T. boeoticum accidentally cross-pollinated<br />
with the wild goatgrass Aegilops speltoides. The chromosomes<br />
from the two parents were incompatible, but chromosome<br />
doubling produced a fertile hybrid with chromosomes<br />
in groups <strong>of</strong> four (tetraploid, with 28 chromosomes), the wild<br />
T. dicoccoides. This was bred by ancient farmers into the<br />
emmer wheat T. dicoccum and into durum (T. durum) and<br />
other wheats. T. dicoccum also accidentally cross-pollinated<br />
with a wild goatgrass, this time A. squarrosa. Chromosome<br />
doubling turned a sterile hybrid into a species <strong>of</strong> wheat with<br />
chromosomes in groups <strong>of</strong> six (hexaploid, 42 chromosomes),<br />
which are today’s major wheat species T. spelta (spelt) and<br />
T. aestivum (bread wheat) (see table on page 203). One <strong>of</strong><br />
the distinguishing features <strong>of</strong> modern bread wheat is the gluten<br />
protein, which makes the flour sticky, allowing it to hold<br />
in bubbles <strong>of</strong> carbon dioxide during leavening. The gene for<br />
gluten apparently came not from the Triticum ancestor but<br />
from the Aegilops squarrosa weed with which emmer wheat<br />
accidentally hybridized.<br />
Hybridization, aside from producing new plant species,<br />
can also facilitate the transfer <strong>of</strong> genes from one species<br />
<strong>of</strong> plant to another. Consider two plant species that<br />
hybridize and produce a new hybrid species. The two original<br />
plant species seldom hybridize with one another, but<br />
may be able to crossbreed more <strong>of</strong>ten with the intermediate<br />
hybrid species. Genes from the first plant species can enter<br />
into the hybrid population by cross-pollination and can<br />
then enter into the population <strong>of</strong> the second plant species,<br />
also by cross-pollination. The hybrid species has thus acted<br />
as a bridge over which genes have crossed, or introgressed,<br />
from one species into another. This concept <strong>of</strong> introgressive<br />
hybridization or introgression was first suggested by Edgar<br />
Anderson, an early 20th-century expert on the genetic history<br />
<strong>of</strong> crop species.<br />
The relative rarity <strong>of</strong> hybridization between species suggests<br />
that a new species, once it evolves, has a coordinated<br />
team <strong>of</strong> genes that would be disrupted by mixing with a<br />
different set <strong>of</strong> genes. This may explain why evolutionary<br />
changes may occur so rapidly when a species first forms,<br />
followed by a period <strong>of</strong> relative stability (see punctuated<br />
equilibria).<br />
Further <strong>Reading</strong><br />
Gompert, Zachariah, et al. “Homoploid hybrid speciation in an<br />
extreme habitat.” Science 314 (2006): 1,923–1,925.<br />
Schwarzbach, Andrea, Lisa A. Donovan, and Loren H. Rieseberg.<br />
“Transgressive character expression in a hybrid sunflower species.”<br />
American Journal <strong>of</strong> Botany 88 (2001): 270–277.