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Autopolyploidy and allopolyploidy. If A and B are the haploid genomes of species A and B, respectively, then autopolyploidy produces a species with an AAAA karyotype, and allopolyploidy (with chromosome doubling) produces a species with AABB karyotype. If A represents seven chromosomes, then an AA diploid has fourteen chromosomes and an AAAA tetraploid has twenty-eight chromosomes. If B represents ve chromosomes, then a BB diploid has ten chromosomes and an AABB allotetraploid has twenty-four chromosomes.
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Although polyploids in the animal kingdom are known (in some species of lizards, fish, invertebrates, and a
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II. Mendelism and the Chromosomal Theory
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8. Cytogenetics
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The McGraw Hill Companies, 2001
Summary
Radish (Raphanus, 2n = 18) Gametes (n = 9)
Cabbage (Brassica, 2n = 18) Gametes (n = 9)
F1 hybrid (sterile, 2n = 18)
Chromosome doubling
Raphanobrassica (fertile, 4n = 36)
tetraploid mammal, the red viscacha rat), polyploidy as a successful evolutionary strategy is primarily a plant phenomenon. There are several reasons for this. To begin with, many more animals than plants have chromosomal sex-determining mechanisms. Polyploidy severely disrupts these mechanisms. For example, Bridges discovered a tetraploid female fruit y, but it has not been possible to produce a tetraploid male. The tetraploid female s progeny were triploids and intersexes. A second reason why polyploidy is more common in plants is because plants can generally avoid the meiotic problems of polyploidy longer than most animals. Some plants can exist vegetatively, allowing more time for the rare somatic doubling event to occur that will produce an amphidiploid; animal life spans are more precisely de ned, allowing less time for a somatic doubling. And third, many plants depend on the wind or insect pollinators to fertilize them and thus have more of an opportunity for hybridization. Many animals have relatively elaborate courting rituals that tend to restrict hybridization. Polyploidy has been used in agriculture to produce seedless as well as jumbo varieties of crops. Seedless watermelon, for example, is a triploid. Its seeds are mostly sterile and do not develop. It is produced by growing seeds from the cross between a tetraploid variety and a diploid variety. Jumbo Macintosh apples are tetraploid.
Figure 8.30 Hybridization of cabbage and radish, showing the
resulting hybrid fruiting structures.
S U M M A R Y
STUDY OBJECTIVE 1: To observe the nature and consequences of chromosomal breakage and reunion 178 190 Variation can occur in the structure and number of chromosomes in the cells of an organism. When chromosomes break, the ends become sticky ; they tend to reunite with other broken ends. A single break can lead to deletions or the formation of acentric or dicentric chromosomes. Dicentrics tend to go through breakage-fusion-bridge cycles, which result in duplications and de ciencies. Two breaks in the same chromosome can yield deletions and inversions. Variegation position effects, as well as new linkage arrangements, can result. Inversion heterozygotes produce loop gures during synapsis, which can form either at meiosis or in polytene chromosomes. Heterozygosity for an inversion suppresses crossovers; organisms that are heterozygotes are semisterile. Reciprocal translocations can result from single breaks in nonhomologous chromosomes. These produce crossshaped gures at synapsis and result in semisterility. The Bar eye phenotype of Drosophila is an example of a duplication that causes a position effect. STUDY OBJECTIVE 2: To observe the nature and conse-
quences of variation in chromosome numbers in human and nonhuman organisms 190 199
Changes in chromosome number can involve whole sets (euploidy) or partial sets (aneuploidy) of chromosomes. Aneuploidy usually results from nondisjunction or chromosomal lagging. Several medical syndromes, such as Down, Turner, and Klinefelter syndromes, and the XYY karyotype are caused by aneuploidy.
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
8. Cytogenetics
The McGraw Hill Companies, 2001
Eight
Cytogenetics
Polyploidy leads to dif culties in chromosomal sexdetermining mechanisms, general chromosomal imbalance, and problems during meiotic segregation. It has been more successful in plants than in animals because plants generally lack chromosomal sex-determining mechanisms. Plants can also avoid meiotic problems by propagating vegetatively. In both animals and plants, even-numbered poly-
ploids do better than odd-numbered polyploids because they have a better chance of producing balanced gametes during meiosis. Somatic doubling provides each chromosome in a hybrid organism with a homologue, and thus makes possible tetrad formation at meiosis. New species have arisen by polyploidy.
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