Current Genetics (1983) 7:57-61
Current Genetics @ Springer-Verlag 1983
Analysis of the Effect of Radiation Repair Mutations on the DELl Mutator Region of Saccharomyces cerevisiae K. M. Downs, A. E. Szwast, and S. W. Liebman University of Illinois at Chicago. Department of BiologicalSciences,Box 4348, Chicago, Illinois 60680, USA
Summary. In DELl strains of the yeast, Saccharomyces cerevisiae, the iso-l-cytochrome c (CYC1) region is flanked on either side by Tyl elements in direct orientation which promote cycl deletions of the bracketed DNA in the haploid cell. In this study, we asked which genes might control this event by testing the possibility that the D E L l mutation mechanism requires an enzyme (or enzymes) that is also utilized in the repair of damaged DNA. To this end, we independently coupled eight repair mutations, rad3-2, rad4-4, rad6-1, rad6-3, rad9-1, rev3-1, rad50-1, and rad51-1, to DELl and asked whether D E L l was still functional. We found that none of these rad mutations significantly affects the mutation frequency of 1 0 - 6 - 1 0 -5 established in D E L l strains for the CYC1 locus. Furthermore, we determined that ste7, a temperature-sensitive sterile allele known to alter gene regulation in Ty-mediated mutations, is not required for D E L l function. Finally, DELl is not temperature-sensitive at 23 ° or 37 °C. Key words: Saccharomyces cerevisiae - DELl - rad ste 7
Introduction
In the yeast, Saccharomyces cerevisiae, the DELl phenotype is characterized by a high frequency (10 - 6 - 1 0 -5 ) of deletions occurring during mitotic growth which encompass three genes on the right arm of chromosome X (Liebman et al. 1979): CYC1, the structural gene encoding iso-l-cytochrome c; OSM1, a gene involved in osmotic regulation; and RAD7, one of the requisite genes for excision of UV-induced pyrimidine dimers and Offprint requests to: S. W. Liebman
subsequent repair of this damaged DNA. In normal, dell + strains, deletions of the CYC1 locus generally do not occur even after mutagenic treatment, while other types of c y c l - mutations appear at the low frequency of 1 0 - 9 - 1 0 - 8 (Liebman et al. 1979). Classical genetics suggest that DELl is both cis- and trans- dominant in the diploid cell and segregates as a single Mendelian gene during meiosis. Recent experiments (Liebman et al. 1981) have demonstrated that the DELl phenotype is attributable to Tyl sequences (Cameron et al. 1979) which flank the CYC1 region in direct orientation. All of the DNA between the two Ty i elements is missing in each of the deletions. Some of these deletions differ, however, and are accompanied by any one of the following: loss of one Tyl; retention of most of both Tyl s; loss of most of both Tyls; retention of most of both Tyls with their apparent concomitant inversion; or retention of a single altered Tyl. The types and frequencies of these deletions are inconsistent with the theory that they arise simply by a single recombination event between the two bracketing Tyl elements. Rather, the deletions seem to involve preferential recombination at the directly repeated termini (deltasequences) of the Tyls. Our rationale forthe present study was that these recombination events might necessitate the products of certain known repair genes (designated "rad"), or mating loci (see below). We have already shown (Liebman and Downs 1980) that rad52-1, the gene whose product mediates generalized recombination, is not required for cycl deletion formation. This communication involves a further sampling of various DNA repair genes from the three DNA repair groups of yeast and their effect on D E L l , as well as the effect of the ste7 mutation on this region. The following rad mutations were coupled to DELl in independent experiments and an analysis of subsequent D E L l function in the haploid cells undertaken (see C. Lawrence in: Advances in Genetics 1981 for a
58
K.M. Downs et al.: Radiation Repair Mutation
comprehensive review of tad genes and mutagenesis): rad3 and rad4, loci necessary for the excision of UV-induced pyrimidine dimers (Parry and Parry 1969; Prakhsh 1977); rev3-1, a mutation which is deficient with respect to UV-induced mutagenesis, thereby causing a decrease in the revertibility of the highly UV-revertible ochre allele, ar94-17 (Lemontt 1971); tad6-1, a complex and little understood mutation having pleiotropic effects, among which is a reduction or abolition of induced mutagenesis by UV, X-rays, and alkylating agents (Prakash 1974; Prakash and Prakash 1976; Lawrence and Christensen 1976) and an inability to spomlate (Cox and Game 1974); rad6-3, a mutation showing effects similar to those of rad6-1, but which differs from rad6-1 in meiotic recombination (Montelone et al. 1981); tad9, a mutation which fails into the same epistatic group as rad6 and rev3 (Lawrence and Christensen 1976) and which is deficient in UV-induced mutagenesis; and rad50-1 and rad51-1, mutations which confer ionizing radiation and MMSsensitivity on yeast cells (Game and Mortimer 1974). rad51-1 is defective in the repair of double-strand breaks (Ho 1975; Resnick and Martin 1976; Mowat and Hastings 1979), but it is capable of repairing induced singlestrand breaks (Mowat and Hastings 1979). The ste7 gene has been shown to be a regulatory element affecting Ty-mediated regulation of several genes (Errede et al. 1980). Roeder et ai. (1980) have described other regulatory genes, called spin, which also affect Tyor delta- mediated regulation. One such gene, spm2, also increases the frequency of delta-delta recombination, resulting in the loss of a Ty element in the HIS4 region. We wanted to know whether ste7 might likewise act as a regulatory factor for the Tyl-induced recombination in the CYC1 region of our D E L l strains.
Contopoulou, Curator of the Yeast Stock Center in Berkeley, California, and is from Dr. Leland Hartwell's collection.
We also wanted to know whether the protein or proteins involved in the deletion-forming mechanism were temperature-sensitive. Experiments carried out by Reif and Saedler (1975) showed that the frequency of deletions for the IS1 insertion sequence in the 9al operon o f E. coli K12 is lowered when the temperature is raised. We tested the possibility that D E L l might also be temperature-sensitive by carrying out cycl mutation frequency determinations at 23 °C and 37 °C. In this paper, we report that none of the rad genes examined, or ste7, significantly altered the established 1 0 - 6 _ 1 0 - 5 cycl mutation frequency for D E L l strains, nor did temperatures of 23 °C and 37 °C have any effect on this frequency.
Verification of the Continued Presence of the rad and ste7 Alleles in cycl Deletion Strains. Severalrad3-2, rad4-4, rad9-1, and rev3-1 cycl deletions were complemented with appropriate rad strains and UV-irradiated for verification of the continued presence of the rad or rev alleles. Likewise, rad6-3 deletions were complemented with a rad6-1 tester strain and the diploid colonies were tested for their UV- and MMS-sensitivity.None of these alleles had reverted. Also, all one hundred eight rad50-1 cycl deletions and all fifty-three rad51-1 cycl deletions complemented with appropriate rad tester strains were still MMS-sensitive.All 56 ste7 cycl deletions were capable of mating at 23 °C but were sterile at 37 °C. The rad3-2 CYCI+ DELl, rad4-4 CYC1+DELl, rad6-3 CYC1+ DELl, rad9-1 CYC1+ DELl, and rev3-1 CYC1+ DELl parent strains were also subcloned to complete YPD medium and 30 independent subclones from each rad strain were picked up and complementation tested for their reversion frequency. Except for rev3-1 (in which 10% of the colonies had reverted to REV3+), none of the subclones had reverted. We found that rad6-1, an unstable, amber-suppressible nonsense mutation spontaneously reverts at a frequency of 0.1-0.3 in SL512-4B and SL512-6C when complementation tested for UVand MMS-sensitivity.This reversion frequency does not, however, alter the 10- 6 cycl mutation frequency obtained in the two strains tested, as 71-94% of the chlorolactate-resistant colonies picked up from them were still rad6-1.
Materials and Methods Origin of Strains. rad3-2, rev3-1, rad4-4, rad6-1, rad6.3, rad9-1, rad50-1, and rad51-1, were kindly supplied by Dr. Louise Prakash and Dr. Christopher Lawrence. ste7 was supplied by Rebecca
Strain Construction. Standard yeast genetic procedures of crossing, sporulation, and tetrad analysis were used. Coupling of rad and DELl have been described (Liebman and Downs 1980), as has scoring of cycl mutations (Liebman et al. 1979). Since all the rad genes coupled to DELl are unlinked to DELl, they assort independently during meiosis and were scored as follows: rad3-2, rad4-4, rad6-1, rad6-3, and radg-1 mutants were detected by a lack of growth on complete medium after a UV-irradiation dose of 750 ergs/mm2 (tad3-2, rad4-4, rad6-1, and rad6-3) or 1,250-1,580 ergs/mm2 (rad9-1) using a UV germicidal lamp. rad6-1 and rad6-3 mutants were also detected by a concomitant lack of growth on 0.009% MMS medium (Liebman and Downs 1980). rad50-1 and rad51-1 were detected by lack of growth on 0.009% MMS.rev3-1 mutants were detected by spotting segregants on arginineless medium (Lemontt 1971), UV-irradiating at doses of 140 and 280 ergs/mm2, and screening for revertant colonies (Lemontt 1971). Segregants which gave rise to fewer revertant colonies were scored as rev-3-1 mutants, ste7 was scored by its ability to mate at 23 °C but not at 37 °C. In one of the coupling experiments (rad50-1 DELl), the resultant rad DELl strain was put through a second mating with a RAD +DELl strain in order to clarify an unfavorable genetic background for the isolation of cycl mutants. Mutation Frequency. Mutation frequencies were determined as previously described (Liebman et al. 1979; Liebman and Downs 1980). The rad DELl experiments were carried out at 30 °C, while that for ste7 DELl was carried out at 37 °C. To test for the temperature-sensitivity of DELl, SL392-1B (CYC1 + DELl) and SL392-2B (CYC1 + DELl) were subcloned on synthetic lactate media plus amino acids to select for CYC1+ (Sherman et al. 1974) at 23 °C, 30 °C, or 37 °C and plated to chlorolactate media (Sherman et al. 1974), on which the ceils were then incubated at 23 °C, 30 °C, or 37 °C until resistant colonies of all sizesappeared (7 30 days). Resistant colonies were tested for the presence of cycl mutations at 30 °C.
K. M. Downs et al.: Radiation Repair Mutation
59
Table 1. Frequency of cycl deletions in various rad DELl, rev DELl, and ste DELl strains Strain No.
Expt. No. a
Relevant Genotype
No. cells per plate after residual growth b
Av. no. resistant mutants per plate
% of resistant mutants that are cycl deletions c
Freq. of cycl deletions
AS1-2B
1
rad3-2DEL1
1.2 (-+0.4) x 107
244 -+ 22
54.1
1.3 (-+0.5) x 10 - 5
AS5-6C
1
rad4-4DEL1
4.5 (-+0.6) x 107
203 -+ 22
44.3
2.1 (-+0.5) x 10 - 6
i
rad6-1 DELl rad6-1 DELl
5.6 (±1.0) x 107 1.2 (-+0.9) x 108
343 227
1.0 3.7
6.3 (-+1.1) x 10 - 8 1.6 (-+1.2) x 10 - 7
rad6-1 DELl tad6-1 DELl
2.4 (-+0.4) x 107 9.7 (-+1.2) x 106
131 ± 7 209 -+ 19
5.2 1.6
2.9 (-+0.6) x 10 - 7 3.6 (+-0.8) x 10 - 7
2
rad6-1 DELl rad6-1 DELl
4.5 (-+0.4) x 107 3.3 (-+0.2) x 107
75 ± 8 160 +- 24
0.9 2.6
1.6 (-+0.3) x 10 - 8 1.3 (-+0.3) x 10 - 7
SL512-4C
1
rad6-1 DELl
4.0 (-+0.3) x 107
218 ± 11
11.5
6.3 (-+0.8) x 10 - 7
SL512-5B
1
rad6-1 DELl
7.1 (-+1.3) x 107
290 -+ 57
0.9
4.0 (-+1.5) x 10 - 8
SL512-4B
1
rad6-1 DELl rad6-1 DELl
2.4 (+-0.4) x 107 2.2 (-+0.1) x 107
198 -+ 29 193 -+ 17
29.8 40.9
2.6 (-+0.7) x 10 - 6 3.6 (-+0.5) x 10 - 6
2
rad6-1 DELl rad6-1 DELl
2.6 (±0.9) x 107 2.9 (±0.5) x 107
203 ± 8 162 ± 16
54.4 63.2
4.9 (-+1.9) x 10 - 6 3.7 (-+1.0) x 10 - 6
SL512-7A
1
rad6-1 DELl
1.7 (-+0.3) x 107
231 +- 38
50.7
7.3 (±2.4) x 10 - 6
SL512-8A
1
rad6-1 DELl
2.8 (-+0.4) x 107
1,015 -+ 92
83.9
3.1 (-+0.7) x 10 - 5
AS4-11C
2 SL512-1A
1
2 SL512-3D
1
2 SL512-6C
1
SL603-1D
1
rad6-3DEL1
8.5 (+-0.5) x 106
724 -+ 77
1.1
9.8 (-+1.6) x 10 - 7
AS3-2B
1
rev3-1 DELl
1.8 (±0.4) x 107
401 +- 12
49.6
1.2 (-+0.3) x 10 - 5
AS6-1C
1
rad9-1 DELl
9.4 (+-4.9) x 106
183
13.0
3.5 (±1.8) x 10 - 6
SL559-1D
1
rad50-1 DELl
1.6 (-+0.2) x 107
117 ± 16
28.3
2.2 (-+0.5) x 10 - 6
SL628-2B
1
rad51-1 DELl
9.0 (±1.7) x 106
203 ± 24
48.4
1.2 (-+0.4) x 10 - 5
SL637-4C
1
ste7DEL1
1.7 (+-0.3) x 107
280 +- 48
59.4
1.0 (+-0.4) x 10 - 5
D311-3A
dell +
Current Genetics @ Springer-Verlag 1983
Analysis of the Effect of Radiation Repair Mutations on the DELl Mutator Region of Saccharomyces cerevisiae K. M. Downs, A. E. Szwast, and S. W. Liebman University of Illinois at Chicago. Department of BiologicalSciences,Box 4348, Chicago, Illinois 60680, USA
Summary. In DELl strains of the yeast, Saccharomyces cerevisiae, the iso-l-cytochrome c (CYC1) region is flanked on either side by Tyl elements in direct orientation which promote cycl deletions of the bracketed DNA in the haploid cell. In this study, we asked which genes might control this event by testing the possibility that the D E L l mutation mechanism requires an enzyme (or enzymes) that is also utilized in the repair of damaged DNA. To this end, we independently coupled eight repair mutations, rad3-2, rad4-4, rad6-1, rad6-3, rad9-1, rev3-1, rad50-1, and rad51-1, to DELl and asked whether D E L l was still functional. We found that none of these rad mutations significantly affects the mutation frequency of 1 0 - 6 - 1 0 -5 established in D E L l strains for the CYC1 locus. Furthermore, we determined that ste7, a temperature-sensitive sterile allele known to alter gene regulation in Ty-mediated mutations, is not required for D E L l function. Finally, DELl is not temperature-sensitive at 23 ° or 37 °C. Key words: Saccharomyces cerevisiae - DELl - rad ste 7
Introduction
In the yeast, Saccharomyces cerevisiae, the DELl phenotype is characterized by a high frequency (10 - 6 - 1 0 -5 ) of deletions occurring during mitotic growth which encompass three genes on the right arm of chromosome X (Liebman et al. 1979): CYC1, the structural gene encoding iso-l-cytochrome c; OSM1, a gene involved in osmotic regulation; and RAD7, one of the requisite genes for excision of UV-induced pyrimidine dimers and Offprint requests to: S. W. Liebman
subsequent repair of this damaged DNA. In normal, dell + strains, deletions of the CYC1 locus generally do not occur even after mutagenic treatment, while other types of c y c l - mutations appear at the low frequency of 1 0 - 9 - 1 0 - 8 (Liebman et al. 1979). Classical genetics suggest that DELl is both cis- and trans- dominant in the diploid cell and segregates as a single Mendelian gene during meiosis. Recent experiments (Liebman et al. 1981) have demonstrated that the DELl phenotype is attributable to Tyl sequences (Cameron et al. 1979) which flank the CYC1 region in direct orientation. All of the DNA between the two Ty i elements is missing in each of the deletions. Some of these deletions differ, however, and are accompanied by any one of the following: loss of one Tyl; retention of most of both Tyl s; loss of most of both Tyls; retention of most of both Tyls with their apparent concomitant inversion; or retention of a single altered Tyl. The types and frequencies of these deletions are inconsistent with the theory that they arise simply by a single recombination event between the two bracketing Tyl elements. Rather, the deletions seem to involve preferential recombination at the directly repeated termini (deltasequences) of the Tyls. Our rationale forthe present study was that these recombination events might necessitate the products of certain known repair genes (designated "rad"), or mating loci (see below). We have already shown (Liebman and Downs 1980) that rad52-1, the gene whose product mediates generalized recombination, is not required for cycl deletion formation. This communication involves a further sampling of various DNA repair genes from the three DNA repair groups of yeast and their effect on D E L l , as well as the effect of the ste7 mutation on this region. The following rad mutations were coupled to DELl in independent experiments and an analysis of subsequent D E L l function in the haploid cells undertaken (see C. Lawrence in: Advances in Genetics 1981 for a
58
K.M. Downs et al.: Radiation Repair Mutation
comprehensive review of tad genes and mutagenesis): rad3 and rad4, loci necessary for the excision of UV-induced pyrimidine dimers (Parry and Parry 1969; Prakhsh 1977); rev3-1, a mutation which is deficient with respect to UV-induced mutagenesis, thereby causing a decrease in the revertibility of the highly UV-revertible ochre allele, ar94-17 (Lemontt 1971); tad6-1, a complex and little understood mutation having pleiotropic effects, among which is a reduction or abolition of induced mutagenesis by UV, X-rays, and alkylating agents (Prakash 1974; Prakash and Prakash 1976; Lawrence and Christensen 1976) and an inability to spomlate (Cox and Game 1974); rad6-3, a mutation showing effects similar to those of rad6-1, but which differs from rad6-1 in meiotic recombination (Montelone et al. 1981); tad9, a mutation which fails into the same epistatic group as rad6 and rev3 (Lawrence and Christensen 1976) and which is deficient in UV-induced mutagenesis; and rad50-1 and rad51-1, mutations which confer ionizing radiation and MMSsensitivity on yeast cells (Game and Mortimer 1974). rad51-1 is defective in the repair of double-strand breaks (Ho 1975; Resnick and Martin 1976; Mowat and Hastings 1979), but it is capable of repairing induced singlestrand breaks (Mowat and Hastings 1979). The ste7 gene has been shown to be a regulatory element affecting Ty-mediated regulation of several genes (Errede et al. 1980). Roeder et ai. (1980) have described other regulatory genes, called spin, which also affect Tyor delta- mediated regulation. One such gene, spm2, also increases the frequency of delta-delta recombination, resulting in the loss of a Ty element in the HIS4 region. We wanted to know whether ste7 might likewise act as a regulatory factor for the Tyl-induced recombination in the CYC1 region of our D E L l strains.
Contopoulou, Curator of the Yeast Stock Center in Berkeley, California, and is from Dr. Leland Hartwell's collection.
We also wanted to know whether the protein or proteins involved in the deletion-forming mechanism were temperature-sensitive. Experiments carried out by Reif and Saedler (1975) showed that the frequency of deletions for the IS1 insertion sequence in the 9al operon o f E. coli K12 is lowered when the temperature is raised. We tested the possibility that D E L l might also be temperature-sensitive by carrying out cycl mutation frequency determinations at 23 °C and 37 °C. In this paper, we report that none of the rad genes examined, or ste7, significantly altered the established 1 0 - 6 _ 1 0 - 5 cycl mutation frequency for D E L l strains, nor did temperatures of 23 °C and 37 °C have any effect on this frequency.
Verification of the Continued Presence of the rad and ste7 Alleles in cycl Deletion Strains. Severalrad3-2, rad4-4, rad9-1, and rev3-1 cycl deletions were complemented with appropriate rad strains and UV-irradiated for verification of the continued presence of the rad or rev alleles. Likewise, rad6-3 deletions were complemented with a rad6-1 tester strain and the diploid colonies were tested for their UV- and MMS-sensitivity.None of these alleles had reverted. Also, all one hundred eight rad50-1 cycl deletions and all fifty-three rad51-1 cycl deletions complemented with appropriate rad tester strains were still MMS-sensitive.All 56 ste7 cycl deletions were capable of mating at 23 °C but were sterile at 37 °C. The rad3-2 CYCI+ DELl, rad4-4 CYC1+DELl, rad6-3 CYC1+ DELl, rad9-1 CYC1+ DELl, and rev3-1 CYC1+ DELl parent strains were also subcloned to complete YPD medium and 30 independent subclones from each rad strain were picked up and complementation tested for their reversion frequency. Except for rev3-1 (in which 10% of the colonies had reverted to REV3+), none of the subclones had reverted. We found that rad6-1, an unstable, amber-suppressible nonsense mutation spontaneously reverts at a frequency of 0.1-0.3 in SL512-4B and SL512-6C when complementation tested for UVand MMS-sensitivity.This reversion frequency does not, however, alter the 10- 6 cycl mutation frequency obtained in the two strains tested, as 71-94% of the chlorolactate-resistant colonies picked up from them were still rad6-1.
Materials and Methods Origin of Strains. rad3-2, rev3-1, rad4-4, rad6-1, rad6.3, rad9-1, rad50-1, and rad51-1, were kindly supplied by Dr. Louise Prakash and Dr. Christopher Lawrence. ste7 was supplied by Rebecca
Strain Construction. Standard yeast genetic procedures of crossing, sporulation, and tetrad analysis were used. Coupling of rad and DELl have been described (Liebman and Downs 1980), as has scoring of cycl mutations (Liebman et al. 1979). Since all the rad genes coupled to DELl are unlinked to DELl, they assort independently during meiosis and were scored as follows: rad3-2, rad4-4, rad6-1, rad6-3, and radg-1 mutants were detected by a lack of growth on complete medium after a UV-irradiation dose of 750 ergs/mm2 (tad3-2, rad4-4, rad6-1, and rad6-3) or 1,250-1,580 ergs/mm2 (rad9-1) using a UV germicidal lamp. rad6-1 and rad6-3 mutants were also detected by a concomitant lack of growth on 0.009% MMS medium (Liebman and Downs 1980). rad50-1 and rad51-1 were detected by lack of growth on 0.009% MMS.rev3-1 mutants were detected by spotting segregants on arginineless medium (Lemontt 1971), UV-irradiating at doses of 140 and 280 ergs/mm2, and screening for revertant colonies (Lemontt 1971). Segregants which gave rise to fewer revertant colonies were scored as rev-3-1 mutants, ste7 was scored by its ability to mate at 23 °C but not at 37 °C. In one of the coupling experiments (rad50-1 DELl), the resultant rad DELl strain was put through a second mating with a RAD +DELl strain in order to clarify an unfavorable genetic background for the isolation of cycl mutants. Mutation Frequency. Mutation frequencies were determined as previously described (Liebman et al. 1979; Liebman and Downs 1980). The rad DELl experiments were carried out at 30 °C, while that for ste7 DELl was carried out at 37 °C. To test for the temperature-sensitivity of DELl, SL392-1B (CYC1 + DELl) and SL392-2B (CYC1 + DELl) were subcloned on synthetic lactate media plus amino acids to select for CYC1+ (Sherman et al. 1974) at 23 °C, 30 °C, or 37 °C and plated to chlorolactate media (Sherman et al. 1974), on which the ceils were then incubated at 23 °C, 30 °C, or 37 °C until resistant colonies of all sizesappeared (7 30 days). Resistant colonies were tested for the presence of cycl mutations at 30 °C.
K. M. Downs et al.: Radiation Repair Mutation
59
Table 1. Frequency of cycl deletions in various rad DELl, rev DELl, and ste DELl strains Strain No.
Expt. No. a
Relevant Genotype
No. cells per plate after residual growth b
Av. no. resistant mutants per plate
% of resistant mutants that are cycl deletions c
Freq. of cycl deletions
AS1-2B
1
rad3-2DEL1
1.2 (-+0.4) x 107
244 -+ 22
54.1
1.3 (-+0.5) x 10 - 5
AS5-6C
1
rad4-4DEL1
4.5 (-+0.6) x 107
203 -+ 22
44.3
2.1 (-+0.5) x 10 - 6
i
rad6-1 DELl rad6-1 DELl
5.6 (±1.0) x 107 1.2 (-+0.9) x 108
343 227
1.0 3.7
6.3 (-+1.1) x 10 - 8 1.6 (-+1.2) x 10 - 7
rad6-1 DELl tad6-1 DELl
2.4 (-+0.4) x 107 9.7 (-+1.2) x 106
131 ± 7 209 -+ 19
5.2 1.6
2.9 (-+0.6) x 10 - 7 3.6 (+-0.8) x 10 - 7
2
rad6-1 DELl rad6-1 DELl
4.5 (-+0.4) x 107 3.3 (-+0.2) x 107
75 ± 8 160 +- 24
0.9 2.6
1.6 (-+0.3) x 10 - 8 1.3 (-+0.3) x 10 - 7
SL512-4C
1
rad6-1 DELl
4.0 (-+0.3) x 107
218 ± 11
11.5
6.3 (-+0.8) x 10 - 7
SL512-5B
1
rad6-1 DELl
7.1 (-+1.3) x 107
290 -+ 57
0.9
4.0 (-+1.5) x 10 - 8
SL512-4B
1
rad6-1 DELl rad6-1 DELl
2.4 (+-0.4) x 107 2.2 (-+0.1) x 107
198 -+ 29 193 -+ 17
29.8 40.9
2.6 (-+0.7) x 10 - 6 3.6 (-+0.5) x 10 - 6
2
rad6-1 DELl rad6-1 DELl
2.6 (±0.9) x 107 2.9 (±0.5) x 107
203 ± 8 162 ± 16
54.4 63.2
4.9 (-+1.9) x 10 - 6 3.7 (-+1.0) x 10 - 6
SL512-7A
1
rad6-1 DELl
1.7 (-+0.3) x 107
231 +- 38
50.7
7.3 (±2.4) x 10 - 6
SL512-8A
1
rad6-1 DELl
2.8 (-+0.4) x 107
1,015 -+ 92
83.9
3.1 (-+0.7) x 10 - 5
AS4-11C
2 SL512-1A
1
2 SL512-3D
1
2 SL512-6C
1
SL603-1D
1
rad6-3DEL1
8.5 (+-0.5) x 106
724 -+ 77
1.1
9.8 (-+1.6) x 10 - 7
AS3-2B
1
rev3-1 DELl
1.8 (±0.4) x 107
401 +- 12
49.6
1.2 (-+0.3) x 10 - 5
AS6-1C
1
rad9-1 DELl
9.4 (+-4.9) x 106
183
13.0
3.5 (±1.8) x 10 - 6
SL559-1D
1
rad50-1 DELl
1.6 (-+0.2) x 107
117 ± 16
28.3
2.2 (-+0.5) x 10 - 6
SL628-2B
1
rad51-1 DELl
9.0 (±1.7) x 106
203 ± 24
48.4
1.2 (-+0.4) x 10 - 5
SL637-4C
1
ste7DEL1
1.7 (+-0.3) x 107
280 +- 48
59.4
1.0 (+-0.4) x 10 - 5
D311-3A
dell +
