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Gamet yang terbentuk dari individu dihibrid A. Gen tidak berangkai B. Kedua Gen berangkai : Couple Phases, Kedudukan Sis Cara penulisan: (AB) ab C. Kedua Gen berangkai : Repulsion Phases, Kedudukan Trans Cara penulisan: (Ab) aB
Lalat Drosophila gen terangkai: Cu = gen untuk sayap normal cu = gen untuk sayap keriput Sr = gen untuk dada polos (normal) sr = gen untuk dada bergaris-garis
Cu Sr cu sr Cu sr cu Sr
Cu Sr Cu Sr
F1
jantan
cu sr cu sr
Cu Sr cu sr
3:0;0;1
F2 :
Linkage
Eukaryotic chromosomes are long ds DNA molecules Typical chromosome contains thousands of genes (loci) Linkage
loci located on the same chromosome linked loci tend to be transmitted as a unit
Linkage
Because they are a group of genes linked together, chromosomes are functionally linkage groups # of linkage groups = the # of types of chromosomes
Crossing over causes loci that are far apart on the same chromosome to sometimes independently assort
Crossing over (meiotic recombination) Occurs during prophase I of meiosis at the bivalent stage
Figure 5.1
gametes with a combination of alleles NOT found in the original chromosomes as a result of meiotic recombination These are called nonparental or recombinant gametes
Figure 5.1
Example of Linkage
Bateson and Punnett conducted a cross in sweet pea involving two traits
Example of Linkage
x Purple flowers, long pollen (PPLL) F1 offspring Red flowers, round pollen (ppll)
Self-fertilization
F2 offspring phenotypes
P NP NP P
(9:3:3:1)
Purple flowers, long pollen Purple flowers, round pollen Red flowers, long pollen Red flowers, round pollen 296 19 27 85 15.6 1.0 1.4 4.5 240 80 80 27 9 3 3 1
The first direct evidence of linkage came from studies of Thomas Hunt Morgan Morgan investigated several traits that followed an X-linked pattern of inheritance
y w m/y w m
y+ w+ m+ Y
F1 generation contains wild-type females and yellow-bodied, white-eyed, miniature-winged males. ywm/Y
F2 generation
Tan body, red eyes, normal wings Tan body, red eyes, miniature wings Tan body, white eyes, normal wings Tan body, white eyes, miniature wings Yellow body, red eyes, normal wings Yellow body, red eyes, miniature wings Yellow body, white eyes, normal wings Yellow body, white eyes, miniature wings
P Males
P Females
Morgan observed a much higher proportion of the combinations of traits found in the parental generation
Morgans explanation: All three genes are located on the X chromosome Therefore, they tend to be transmitted together as a unit
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Why did the F2 generation have a significant number of nonparental combinations? Why was there a quantitative difference between the various nonparental combinations?
Figure 5.4
Figure 5.4
Incomplete Linkage
Linked loci that are sometimes separated by recombination are inherited in a pattern between linked and independently assorting
The percentage of offspring with the loci linked vs those with the loci separated is a measure of the physical distance separating the loci on the chromosome
An undergraduate in the laboratory of T. H. Morgan Constructed first genetic map in 1911 Sturtevant wrote:
In conversation with Morgan I suddenly realized that the variations in the length of linkage, already attributed by Morgan to differences in the spatial orientation of the genes, offered the possibility of determining sequences [of different genes] in the linear dimension of the chromosome. I went home and spent most of the night (to the neglect of my undergraduate homework) in producing the first chromosome map, which included the sex-linked genes, y, w, v, m, and r, in the order and approximately the relative spacing that they still appear on the standard maps.
Estimating the relative distances between linked genes, based on the amount of recombination occuring between them allows us to generate genetic maps
If the genes are far apart many recombinant offspring If the genes are close very few recombinant offspring
Number of recombinant offspring X 100 Total number of offspring The units of distance are called map units (mu) They are also referred to as centiMorgans (cM) One map unit is equivalent to 1% recombination frequency
Map distance =
Genetic mapping experiments are typically accomplished by carrying out a testcross Example of a two-point mapping cross
Cross of two linked genes affecting bristle length and body color in fruit flies
One parent double recessive (homozygous recessive at both loci) s/s ; e/e Other parent is heterozygous at both loci (+/s ; +/e)
Chromosomes are the product of a crossover during meiosis in the heterozygous parent
Figure 5.9
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The phenotype data are used to estimate the distance between the two loci
X 100
Therefore, the s and e genes are 12.3 map units apart from each other along the same chromosome
Trihybrid Crosses
Data from trihybrid crosses can also yield information about map distance and gene order The following experiment outlines a common strategy for using trihybrid crosses to map genes In this example, we will consider fruit flies that differ in body color, eye color and wing shape
b = black body color b+ = grey body color pr = purple eye color pr+ = red eye color vg = vestigial wings vg+ = normal wings
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The goal in this step is to obtain F1 individuals that are heterozygous for all three genes
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Step 2: Perform a testcross by mating F1 female heterozygotes to homozygous recessive, male flies
During gametogenesis in the heterozygous female F1 flies, crossovers may produce new combinations of the 3 alleles
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411 61 2 30 28 1 60
412
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Analysis of the F2 generation flies will allow us to map the three genes
The three genes exist as two alleles each Therefore, there are 23 = 8 possible combinations of offspring If the genes assorted independently, all eight combinations would occur in equal proportions It is obvious that they are far from equal
Parental phenotypes occur most frequently Double crossover phenotypes occur least frequently Single crossover phenotypes occur with intermediate frequency
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The combination of traits in the double crossover tells us which gene is in the middle A double crossover separates the gene in the middle from the other two genes at either end
In the double crossover categories, the recessive purple eye color is separated from the other two recessive alleles Thus, the gene for eye color lies between the genes for body color and wing shape
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Step 4: Calculate the map distance between pairs of genes To do this, one strategy is to group the data according to pairs of phenotypes resulting from non-crossovers, single crossovers, & double crossovers
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Phenotype
Gray body, purple eyes, vestigial wings Black body, red eyes, normal wings Gray body, red eyes, vestigial wings Black body, purple eyes, normal wings Gray body, purple eyes, normal wings Black body, red eyes, vestigial wings
28
30 + 28
= 0.058
1,005
61
60
61 + 60
= 0.120 1,005
1+2
= 0.003 1,005
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To determine the map distance between the genes, we need to consider both single and double crossovers
To calculate the distance between b and pr Map distance = (0.058 + 0.003) X 100 = 6.1 mu To calculate the distance between pr and vg Map distance = (0.120 + 0.003) X 100 = 12.3 mu To calculate the distance between b and vg
The double crossover frequency needs to be multiplied by two Because both crossovers are occurring between b and vg
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Based on the map unit calculation the body color and wing shape genes are farthest apart The eye color gene is in the middle
The data is also consistent with the map being drawn as vg pr b (from left to right) In detailed genetic maps, the locations of genes are mapped relative to the centromere
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Alternatively, the distance between b and vg can be obtained by simply adding the map distances between b and pr, and between pr and vg Map distance = 6.1 + 12.3 = 18.4 mu
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Interference
The product rule allows us to predict the likelihood of a double crossover from the individual probabilities of each single crossover
Interference
Therefore, we would expect seven or eight offspring to be produced as a result of a double crossover However, the observed number was only three!
Two with gray bodies, purple eyes, and normal sings One with black body, red eyes, and vestigial wings
This lower-than-expected value is due to a common genetic phenomenon, termed positive interference The first crossover decreases the probability that a second crossover will occur nearby
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7.5
I = 1 C = 1 0.4 = 0.6 or 60% This means that 60% of the expected number of crossovers did not occur
Rarely, the outcome of a testcross yields a negative value for interference This suggests that a first crossover enhances the rate of a second crossover
The molecular mechanisms that cause interference are not completely understood However, most organisms regulate the number of crossovers so that very few occur per chromosome
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Fusion of human cell and rodent cell Eventually, most human sequences lost
Lines carrying sub-sets of human chromosomes are compared to determine which express the given gene product
Process of elimination determines on which chromosome (region of chromosome) a gene is located Hybrid Panel
Cytogenetic Mapping
normal
loss of an entire chromosome lethal in all cases for autosomes XO Turners syndrome is only viable monosomy
Mapping Strategies
Deletion Mapping
Cross recessive heterozygote to deletion heterozygote lines appearance of recessive phenotype localizes recessive gene within deletion interval
Complementation
Complementation Analysis
Complementation Analysis
Complementation groups
Complementation Analysis
+ bm + pr
+ pr
+ bm + bm
+ bm + pr
+ pr
+ bm + bm