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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.

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