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Actually, any prokaryote or virus should suf ce, but Benzer made use of other unique screening properties of the phage that made it possible to recognize one particular mutant in about a billion phages Benzer used rII mutants of T4These mutants produce large, smooth-edged plaques on E coli, whereas the wild-type produces smaller plaques whose edges are not as smooth (see g 77) The screening system that Benzer employed made use of the fact that rII mutants do not grow on E coli strain K12, whereas the wild-type canThe normal host strain, E coli B, allows growth of both the wild-type and rII mutants Thus, various mutants can be crossed by mixed infection of E coli B cells, and Benzer could screen for wildtype recombinants by plating the resultant progeny phages on E coli K12 ( g 12.

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Equation 631 is of the form H (j ) = |H |ej H , with |H (j )| = 1+ and H (j ) = 90 arctan 1 arctan 2 1 A

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6), on which only a wildtype recombinant produces a plaque It is possible to detect about one recombinant in a billion phages, all in an afternoon s work This ability to detect recombinants occurring at such a low level of frequency allowed Benzer to see recombinational events occurring very close together on the DNA, events that would normally occur at a frequency too low to detect in fruit ies or corn Benzer sought to map the number of sites subject to recombination and mutation within the rII region of T4 He began by isolating independently derived rII mutants and crossing them among themselves The rst thing he found was that the rII region was composed of two cistrons; almost all of the mutations belonged to one of two complementation groups The A-cistron mutations would not complement each other but would complement the mutations of the B cistron.

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The magnitude and phase plots for the frequency response of the band-pass lter of Figure 619 are shown in Figure 620 These plots have been normalized to have the lter pass-band centered at the frequency = 1 rad/s The frequency response plots of Figure 620 suggest that, in some sense, the band-pass lter acts as a combination of a high-pass and a low-pass lter As illustrated in the previous cases, it should be evident that one can adjust the lter response as desired simply by selecting appropriate values for L, C, and R The expression for the frequency response of a second-order band-pass lter (equation 629) can also be rearranged to illustrate two important features of this circuit: the quality factor, Q, and the resonant frequency, 0 Let 1 0 = LC Then we can write CR = 0 CR =Q 0 0 0 and Q = 0 CR = R 0 L (633)

(634)

The exceptions were mutations that seemed to belong to both cistrons These mutations were soon found to be deletions in which part of each cistron was missing (table 123) Deletion Mapping As the number of independently isolated mutations of the A and B cistrons increased, it became obvious that to make every possible pairwise cross would entail millions of crosses To overcome this prob-.

In equation 634, the resonant frequency, 0 , corresponds to the center frequency of the lter, while Q, the quality factor, indicates the sharpness of the resonance,

recombination at the rII locus of phage T4. Two rII mutants are crossed by infecting the same B-strain bacteria with both phages. The offspring are plated on a lawn of K12 bacteria in which only wild-type phages can grow. The technique thus selects only wild-type recombinants.

Band-pass filter amplitude response 1 08 Amplitude 06 04 02 0 _ 10 3

(47)

lem, Benzer isolated mutants that had partial or complete deletions of each cistron. Deletion mutations were easy to discover because they acted like structural alleles to alleles that were not themselves structurally allelic. In other words, if mutations a, b, and c are functional but not

0 _50

_3 _2

The McGraw Hill Companies, 2001

10 1 100 101 Radian frequency (logarithmic scale)

that is, how narrow or wide the shape of the pass-band of the lter is The width of the pass-band is also referred to as the bandwidth, and it can easily be shown that the bandwidth of the lter is given by the expression B= 0 Q (635)

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convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp

convert tiff to pdf c# itextsharp