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can be accomplished by the formation of a cross-shaped gure ( g. 8.10). Such a gure is diagnostic of a reciprocal translocation. A single crossover in a reciprocal translocation heterozygote will not produce chromatids that are further imbalanced, as it does in an inversion heterozygote. However, reciprocal translocation heterozygotes do produce nonviable progeny. Problems
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can arise when centromeres separate at the rst meiotic division.
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Segregation After Translocation
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Since two homologous pairs of chromosomes are involved, we have to keep track of the independent
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Tamarin: Principles of Genetics, Seventh Edition
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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
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Eight
Cytogenetics
BOX 8.1 CONTINUED
because D. ananassae is commensal with people. Some of Futch s results are shown in gures 1-4, which diagram the left and right arms of the y s second chromosome, as well as the synaptic patterns. We can see vividly the sequence of change in which one inversion occurs after a previous inversion has already taken place. In gures 2 and 4, the standard (a) gave rise to (b), which then gave rise independently to (c) and (d ). The standard is from Majuro in the Marshall Islands and is believed to be in the ancestral group of the species. Ponape is the home of (b), (c) is from Tutuila (eastern Samoa), and (d ) is from New Guinea. Thus, the sequence is Majuro to Ponape, and from there the same stock was transferred to Tutuila and New Guinea. This type of analysis has been useful in the Drosophila group throughout its range but especially in the Paci c Island populations and in the southwestern United States.
(a) Figure 4
31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1
31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 3 4 5 6 7 8 9 10 11 12 13 2 1
14 15 16 17 18 19 20 21 9 10 11 12 13
31 30 29 28 27 26 25 24 23 22 8 7 6 5 4
20 21 22 23 24
31 30 29 28 27 26 25
3 4 5 6 2
19 18 17 16 15 14
3 2 1
13 12 11 10 9 8 7 1
Chromosomal maps of 2R. (a) Standard gene sequence. (b) Ponape: breakpoints of 2RA are indicated and the segment is shown inverted. (c) Tutuila light: breakpoints of 2RB are indicated. 2RA is inverted and 2RB, which overlaps it, is also shown inverted. (d ) New Guinea: breakpoints of 2RC and 2RD are indicated. 2RA is inverted; 2RC, which overlaps 2RA, and 2RD are shown inverted. (From David G. Futch, A study of
speciation in South Paci c populations of Drosophila ananassae, in Marshall R. Wheeler, ed., Studies in Genetics, no. 6615 [Austin: University of Texas Press, 1966]. Reproduced by permission.)
segregation of the centromeres of the two tetrads. There are two common possibilities and one that occurs less often ( g. 8.11). The rst, called alternate segregation, occurs when the rst centromere assorts with the fourth centromere, leaving the second and third centromeres to go to the opposite pole. The result will be balanced
gametes, one with normal chromosomes and the other with a reciprocal translocation. Also likely is the adjacent-1 type of segregation, in which the rst and third centromeres segregate together in the opposite direction from the second and fourth centromeres. Here, both types of gametes are unbalanced, carrying duplica-
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
8. Cytogenetics
The McGraw Hill Companies, 2001
Variation in Chromosomal Structure
f e e a b c d g h i j g
a a a j
b b b i
c c c h
d d d g
e e g f f
g g e
h d h i
i c i j
j b j a
Nonrecombinant chromosome Imbalanced chromosome Nonrecombinant inversion chromosome Imbalanced chromosome
Consequences of a crossover in the loop region of a pericentric inversion heterozygote.
tions and de ciencies that are usually lethal. Since adjacent-1 segregation occurs at a relatively high frequency, a signi cant amount of sterility results from the translocation (as much as 50%). An adjacent-2 type of segregation ( g. 8.11), in which homologous centromeres go to the same pole ( rst with second, third with fourth), is a third possibility.This can result when the cross-shaped double tetrad opens into a circle in late prophase I. In the German cockroach, adjacent2 patterns have been observed in 10 to 25% of meioses, depending upon which chromosomes are involved. In summary, then, reciprocal translocations result in new linkage arrangements, variegated position effects, a cross-shaped gure during synapsis, and semisterility.
markedly different chromosome numbers without any signi cant difference in the quantity of their genetic material.Therefore, cytologists frequently count the number of chromosomal arms rather than the number of chromosomes to get a more accurate picture of species af nities. The number of arms is referred to as the fundamental number, or NF (French: nombre fondamentale). In a similar fashion, centromeric ssion increases the chromosome number without changing the fundamental number.
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