genetic mapofsaccharomyces cerevisiae, edition 9 · genetic mapof s. cerevisiae 183 table 1....

Vol. 49, No. 3MICROBIOLOGICAL REVIEWS, Sept. 1985, p. 181-2120146-0749/85/030181-32$02.00/0Copyright C 1985, American Society for Microbiology

Genetic Map of Saccharomyces cerevisiae, Edition 9ROBERT K. MORTIMER' 2* AND DAVID SCHILD2

Department ofBiophysics and Medical Physics, University of California,' and Division ofBiology and Medicine,Lawrence Berkeley Laboratory,2 Berkeley, California 94720

INTRODUCTION .................................................. 181GENETIC MAP, GLOSSARY, AND LIST OF MAPPED GENES... ..... 185COMMENTS ON NEW ADDITIONS TO EACH CHROMOSOME................................................185Chromosome I...................................................185Chromosome II..........................................185Chromosome Im... .....185ChromosomeIV..188Chromosome V...................................................189Chromosome VI........s**...O..se....g....O..O.so............... 189Chromosome VII .................................................. 191Chromosome VIII .................................................. 193Chromosome IX............................ 194Chromosome X....................194Chromosome XI..................194Chromosome XII ..................197Chromosome XII................. 197Chromosome XIV................. 200Chromosome XV................. 201Chromosome XVI..so............. 206Chromosome XVII..........6**"*..206

DISCUSSION................. 207ACKNOWLEDGMENTS................. 208LITERATURE CITED..................208

INTRODUCTIONThe yeast Saccharomyces cerevisiae is currently widely

used for biochemical, molecular, and genetic research. Be-sides its traditional importance in the baking and brewingindustries, it is increasingly being utilized for the industrialproduction of recombinant deoxyribonucleic acid (DNA)products. The ease by which this yeast can be geneticallyand molecularly manipulated, coupled with its extensivegenetic map, have made it particularly suitable for use inboth research and industry.

In this review we present a new edition of the genetic mapof S. cerevisiae, edition 9. Included are all pertinent data ongenes mapped since our publication of the map in 1980 (132)and data on revisions of previously mapped genes. Thisreview, as was our previous one, is based on both publishedmapping data and unpublished data kindly supplied by manyresearchers. We have decided to call the current genetic mapedition 9, since there have been eight previous editionspartially or wholly emanating from this laboratory. Editions1 to 8 appeared between 1960 and 1984 (69, 70, 129-132, 135,136). In addition to these editions, several genetic mnaps werepublished by the Lindegren group (77, 78, 105-107, 165),including the first genetic map of S. cerevisiae (106). Numer-ous other versions of the genetic map with minor changeshaVe been published by us and others.The new molecular biological techniques being used for

studies on S. cerevisiae yield information abotit the genomenot previously available; because of this, questions arise as

*Corresponding author.

181

to what should be placed on the genetic map and what typesof information should be used. The vast majority of genes onthe current map have been identified by the phenotype(s) ofmutations and have been placed on the map by tetradanalysis. In addition, some genes have been placed on themap by using only aneuploid or mitotic recombinationanalyses (see Discussion). Some regions, such as the ribo-somal ribonucleic acid (RNA) coding region (rDNA) (150)and some of the Tyl transposon sequences (92), have beenincluded on the map even though they have not beenmutated; however, these sequences were mapped by tetradanalysis, using either restriction site polymorphisms or inte-grated plasmids containing scorable markers. Regions con-taining open reading frames, even if they have been shown tobe transcribed, have not been included on the map unlessthese regions have been shown, by disruption or deletionanalysis, to have a phenotype. Some Tyl sequences whichhave been genetically mapped (92) have been included,whereas others which have been identified only by recom-binant DNA techniques have not. Since Tyl sequencestranspose and are found in different numbers and positions indifferent strains (18, 48), the map position of a Tyl elementshould be considered tentative. Although to a lesser extent,restriction site (142, 150) and chromosome size (G. F. Carleand M. V. Olson, Proc. Natl. Acad. Sci. U.S.A., in press)polymorphisms have also been found between differentlaboratory strains of yeasts. Therefore, the distances be-tween genes and possibly even the relative position of genesmay vary between different strains. Determination of theorder of closely linked clusters of genes by tetrad analysis isusually very difficult. Recently, the order of some clusters of

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182 MORTIMER AND SCHILD MICROBIOL. REV.

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14go10jLLsppT8

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Fpho3-SseclioglgTES TF2 -

Nt8,ssn6 -

~gic gic6 -

;sup 45,47NSUP4S.4mokS5

petii

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it--.SUF2-% SUF16

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1%rod IS-ISUP61, [tupl,cy9,.umr7,fI kltsm5-cdc39nHMRMAL2

SUC5

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pho2-cdc2-tsmO225

-spoT4mor lsir2rnoltcdc7

errg6Ilts4arod57.

>\tsmOQ8OSUP27smm162UP88

rod55maok21SUP80

-nul 3,ste5

-prt 3conltrblLmaklOFSUSUFI9mintICCocI23anpl-cup5uro3Kgcn4 ,

arg99SC3tSM0O39

- chol GAL83I gin 3INHS,LBCOR I3,hisiKSUPt9 20LSUP88morg5,6ilvi arg84trp2met6rod51\sam2(ith2)

pet-ts1402

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oarolhom2/ ...- E

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drl1pt 4

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skilL rod54Mcu2dc43~lys5-Llts 3ro2

|metl3,SUP383

<cyh2I mfo2-mak24

olel1hemlO-rod6

UP54trp5--__-smr1

t;sml Tytrrmeialg?ade6

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SUF4SUP77

tfro2

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tfrolthipi..ode3:-ADE15ser2TUFI5wrad2smesl

LMALsucI

-ade8

;pur S

-cupI4

-rna3-snf I-pho8- ssn2

----Mitotic linkage-Trisomiclinkage

) Sequence not estoblishedwith genes outside porenthesis

Left orm is on top

[mok20

mak7

+spoll

ur3dur4

ti ardl13

4rg4tsmO186tsrnOI5ithrl

+put2

TsupillCUPI

cdc12SUF8pol2pot3

I makl8

mak 19

SUC2

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doa 181dall

Sdai 42

ys

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;inol

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csmOI85met3ilv3rev5

|cdc8- cdci1

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lop4

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sm 8740 -Ipe. i -ut-aT*2i

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tp dep44,cdc60

wpho85isinOIISnbt

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hU21mok6

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m-net?%gcd,tra3 KR81-cdc3t

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GENETIC MAP OF S. CEREVISIAE 183

TABLE 1. New tetrad analysis data for chromosome IPSegregation (no.) Ascus type (no.) Map distance

Interval ReferenceFD SD PD NPD T x' (cM) SE

26 0 40 30.7237 0 64 10.756 12 162 54.666 17 232 56.941 0 19 16.367 11 144 51.078 18 207 57.215 1 11 34.9

43 43 30.339 4 52 43.252 0 2 2.4

23 5

69 1

20142

41284

1522

64

01

01

00

0

9.462 11.98 16.6

373

3.511.0

2 6.08 14.3

0.7

13

548 6

501

77 858 4

500 45

0 45

8.8

0.5

4.2

4.83.34.2

4.2635 57

3.31.26.45.53.46.56.5

18.54.49.42.5

4.01.4

13.0

2.11.4

85858585858585858585R. E. Esposito and

C. W. Waddell,personalcommunication

8585R. E. Esposito and


147

4.3 1475.1 1470.6 R. E. Esposito and


2.5 R. E. Esposito andC. W. Waddell,personalcommunication

0.2 C. Cummins andM. Culbertson,personalcommunication

0.6 C. Cummins andM. Culbertson,personalcommunication

1.7 851.6 1010.6 C. Cummins and

M. Culbertson,personalcommunication

0.5

51 115858

0

67

106

9.35.4

43.6

2.7 1012.0 1015.7 169

a FD, First-division segregation; SD, second-division segregation. These segregations are determined by examination of the segregation of a marker relative tothat of a known centromere-linked marker (131, 134).

FIG. 1. Genetic tnap of S. cerevisiae, based on the data presented in Tables 1 through 17, in our earlier reivew (132), and in the text.

Centromeres are represented as circles, and the left arm of each chromosome has arbitrarily been drawn above the centromere. Solid lines

are drawn to scale and represent linkage distances established by tetrad analysis. The dashed and dotted lines represent linkages established

by mitotic and aneuploid analysis, respectively. They have arbitrarily been assigned a minimum distance of 100 cM (see text); these intervals

are not drawn to scale. When the orientation of two or more genes relative to outside markers has not been determined, these genes are

enclosed within parentheses.

cdc24-mak16cdc24-pyklcdc24-cyslcdc24-adelpykl-makl6pykl-cysipyki-adelmakl6-cyslmakl6-cenlmakl6-adelcysl-spo7

cysi-cenicysi-adelcysl-adel

cysl-adelTotal

cysl-cys3cys3-adelspo7-cenl

spo7-adel

trnl-cenl

trnl-adel

ceni-adelceni-adelcenl-adel

Total

SUP56-cenlSUP56-adelFLOI-adel

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184 MORTIMER AND SCHILD

TABLE 2. New tetrad analysis data for chromosome ii"

Segregation (no.) Ascus type (no.) Map distanceInterval

FD SD PD NPD T x' (cM) SE Reference

cly2-AMY2 43 1 59 31.7 3.8 199cly2-cen2 111 133 34.0 2.8 130, 199cly2-gall 25 13 65 115.5 58.7 130AMY2-cen2 60 21 14.0 2.9 199AMY2-gal7 62 0 30 16.6 2.6 199spoT8-pho5 31 0 13 15.4 4.0 191spoT8-lys2 49 1 38 25.3 4.5 191SUP87-gal7 31 1 64 36.6 3.8 68tsm7269-gall 8 0 23 38.0 4.8 72SUP87- 26 0 12 16.6 4.5 68SUP72

SUP87-lys2 33 1 40 31.4 5.1 68SUP72-lys2 43 0 15 13.4 3.3 68tsm7269-lys2 32 0 42 28.7 3.1 72secl8-lys2 111 0 40 13.4 1.9 C. Fields, and R.

Schekman,personalcommunication

secl8-tyri 42 3 110 41.7 3.9 C. Fields and R.Schekman,personalcommunication

algl-lys2 45 0 2 2.2 1.5 33ssn6-lys2 12 0 0 0 21TEF2-lys2 2 162glc6-lys2 117 0 16 6.2 1.6 J. Pringle, personal

communicationSUP46-tyrl 685 1 140 8.8 0.7 143sup47-lys2 74 1 59 24.4 3.1 143syp47-tryl 177 1 69 15.2 1.9 143DUR80-tyrl 197 3 221 28.5 1.7 235DUR80-durl 390 0 31 3.8 0.7 235DUR80-met8 404 0 17 2.1 .5 235

a See footnote a, Table 1, for abbreviations.

genes has been determined by cloning studies; two examplesare the orders of leu2-cen3-cdclO (chromosome III) (28) andcdc9-cdc39-ste7 (chromosome IV) (14, 149).The genetic map is based mainly on data from tetrad

analysis of the four meiotic products (ascospores) fromindividual diploid cells. For any given pair of heterozygousmarkers in a cross (AB x ab), tetrad dissection of thesporulated diploid will result in three types of asci: parentalditype (PD) (AB, AB, ab, ab), nonparental ditype (NPD)(Ab, Ab; aB, aB), and tetratype (T) (AB, Ab, aB, ab). Mapdistances are calculated by using the relative frequencies ofthese classes. In Tables 1 through 17 the map distances x'have been calculated with equations developed by R. Snow(173), using a maximum likelihood analysis. These equationsgive the most accurate determination of map distances sincethey take into account all classes of crossovers between twogenes. These equations are very complex and require acomputer to solve them. Although less accurate, most pub-lished map distances are calculated by using the equationderived by Perkins (7): Xp = (100/2) x [(T + 6NPD)/(PD +NPD + T)].

This equation is reasonably accurate for distances up toabout 35 centimorgans (cM), but underestimates longerdistances (for discussion of this, see references 111, 136,173). An empirical formula has been developed which can beused to convert Xp (map distance calculated from Perkin'sequation) into an accurate approximation of x' (111): Xe =

[(80.7 Xp - 0.883 Xp2)/(83.3 - Xp)]. Xe has been shown to beapplicable for distances >35 cM and can be easily calcu-lated. These mapping equations are used to determinewhether two genes are linked and are applicable for dis-tances up to approximately 100 cM.There are many ways in which the general location of a

gene can first be determined, and most of these techniqueshave recently been reviewed (131, 134). In the last few yeprsadditional techniques have been devised, including two newmethods useful for mapping mutations. The spoll mnappingmethod (89) is based on the analysis of the rare viablemeiotic products resulting from meiosis of diploids homozy-gous for a recombination-deficient mutation. The rad52mapping method (D. Schild and R. K. Mortimer, Genetics,in press) is based on the high frequency of chromosome lossobserved in diploids homozypous for the rad52 mutation(128). This technique has also been modified to make it moresuitable for mapping temperature-conditional mutations(P. J. Hanic-Joyce, Genetics, in press). Two new methodshave also been developed to map cloned genes. The 2 ,ummapping method (49) is based on the instability of a chro-mosomne which contains an integrated plasmid containingpart or all of 2 ,um DNA (51). It is also now possible to mapgenes by using Southern hybridization to individual yeastchromosomes which are first separated with the new orthog-onal-field-alternating gel electrophoresis (OFAGE) tech-nique (19, 163; Carle and Olson, in press).

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TABLE 3. New tetrad analysis data for chromosome IIIa

Segregation (no.) Ascus type (no.) Map distanceInterval Reference

FD SD PD NPD T x' (cM) SE

SUP53-leu2 154 0 1 0.3 0.3 52SUP53-leu2 90 1 3 4.8b 101

Total 244 1 4 1.2 0.9

SUP53-cen3 75 19 10.7 2.4 101Tyl-MAT 32 0 5 6.8 2.9 92Tyl-thr4 23 0 23 25.1 3.7 92secS5-MAT 67 2 100 33.3 3.1 C. Fields and R.

Schekman, personalcommunication

sec55-thr4 139 0 26 7.9 1.4 C. Fields and R.Schekman, personalcommunication

SUFl6- 285 0 25 4.1 0.8 55SUF2

SUFl6-pgkl 233 0 4 0.9 0.4 55SUFl6-cen3 69 6 4.1 1.6 55SUFl6-petl8 56 0 6 4.9 1.9 55SUFl6-MAT 183 2 122 21.9 2.0 55flkl-MAT 10 0 48 41.8 2.9 175flkl-thr4 9 0 11 27.6 5.7 175flkl-MAL2 40 0 16 14.8 3.4 175

a See footnote a, Table 1.b X' could not be determined; calculated by Perkins' (7) equation for Xp.

GENETIC MAP, GLOSSARY, AND LIST OF MAPPEDGENES

Edition 9 of the genetic map of S. cerevisiae is presentedin Fig. 1. It is based on the data in Tables 1 through 17 andon data included in our previous review (132). A generaldiscussion of the genetic map is included at the end of thisarticle. The glossary (Table 18) briefly describes the pheno-types of different groups of genes. The chromosomal arm onwhich each mapped gene is located is included in the list ofmapped genes (Table 19), as are one or more referenceswhich include data on the map position of the gene. Morecomplete descriptions of each gene and the enzyme encodedby them, if known, have been presented by Plischke et al.(151) and Broach (15).

COMMENTS ON NEW ADDITIONS TO EACHCHROMOSOME

In this section, additions and changes to each of thechromosomes since our last major review (132) are dis-cussed. Ambiguities in gene locations or orders are alsodiscussed.

Chromosome I

FLO4 is an allele of FLO] and so has been removed fromthe map. The marker osm, which was placed on chromo-some I by random spore analysis (176), has also beenremoved from the map due to lack of tetrad data. Themutation cdc24 has been mapped on chromosome I distal tocyc3 (85); it has also been shown to be allelic to tsll, whichhas accordingly been removed from the map. Other genesadded to chromosome I are trn) (G. Knapp, C. M. Cum-mins, and M. Culbertson, personal communication), spo7(R. E. Esposito and C. Waddell, personal communication),and SUP56 (101). Chromosome I has been shown byOFAGE to have a size of 260 kilobases (kb) (Carle and

Olson, in press). Its total genetic length is approximately 100cM, which leads to a ratio of 2.6 kb/cM. By both criteria.,chromosome I is the smallest yeast chromosome, with thepossible exception of chromosome XVII (see Discussion).

Chromosome IIChromosome II is one of the most densely mapped chro-

mosomes, with 39 genes now distributed over its length of250 cM. Since our last compilation (132), 15 genes have beenadded to chromosome II. AMY2 and cly2 have been shownto be on the left arm of this chromosome, with AMY2 distal(199). The sporulation-deficient gene spoT8 is on the rightarm and distal to the gal7,10,1 cluster (191). SUP87, a UGAsuppressor (68), and tsm7269 (72) map close to tsm134, andthe order of these three genes and pho3,5 relative to thecentromere and each other is unknown. secl8 (C. Fields andR. Schekman, personal communication), and SUP72 (68)map distal to this cluster. The genes algl (33) and TEF2 (162)are very close together with the probable order cen2-algl-lys2-TEF2. ssn6 has been shown to be an allele of cyc8 (21).sup47 maps in the same region as the omnipotent suppressorsup45 and these two genes are probably allelic; SUP46,another omnipotent suppressor, maps distal to these twogenes (143). dur2 and DUR80 have been shown to be veryclose to durl (25), and the distance of this cluster from met8has been shown to be much shorter than previously re-ported.

Chromosome IllSeveral changes have been made on chromosome III, the

mating type chromosome. SUP53 is now placed distal,rather than proximal, to leu2 (52, 101) and cdclO is locatedon the right arm near the centromere rather than on the leftarm (28). Two genes, SUF16 and secS5, have been added tothe map of this chromosome. The frameshift suppressorSUF16 is 1 map unit distal to pgkl (55). The secretorymutant secS5 maps 7.9 + 1.4 cM distal to thr4 (Fields and

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TABLE 4. New tetrad analysis data for chromosome IVaSegregation (no.) Ascus type (no.) Map distanceInterval Reference1FD SD

SUF25-cdc9SUF25-trplSUF25-HOcdc9-HOcdc9-trplspoT4-trplspoT4-cdc7PHSI-trplerg6-trpltsmOO80-cdc7

tsmOO80-trpl

tsmOO80-radS7

tsmOO80-tsm5162

tsmS162-cdc7

tsmS162-trpl

cdc7-trpl

trpl-radS7

SUP88-trplSUP88-SUP80SUP80-trplSUP80-nul3steS-arolst'S-hom2sup36-arolsecl-hom2

secl-secS

sec5-cdc37

sufl2-hom2sufl2-arolcdc37-arolcdc37-hom2cdc37-ste5adri-arol

PD NPD T x' (cM) SE

181 6 230 32.2 2.2 55, 5642 22 114 110.2 38.6 55176 0 66 13.7 1.5 5680 14 148 53.7 7.7 5643 19 117 88.8 22.6 5533 2 53 37.7 5.9 19124 0 26 26.8 4.1 191149 0 0 0 17923 1 1 14 0b 12247 0 8 7.3 2.4 10; F. Boutelet and F.

Hilger, personalcommunication

48 0 7 6.4 2.3 10; F. Boutelet and F.Hilger, personalcommunication







37 0 17 15.8 3.2 6820 0 34 32.0 3.6 6827 6 95 54.0 7.6 6822 2 41 42.1 8.3 686 0 3 17.0 8.5 1664 0 9 34.8 6.7 166

17 1 19 35.1 10.8 143112 0 6 2.6 1.0 C. Fields and R.


115 0 3 1.3 0.8 C. Fields and R.Schekman, personalcommunication


93 0 7 3.5 1.3 3583 0 17 8.5 1.9 3544 0 17 14.0 2.9 166L00 0 4 2.0 1.0 1668 0 11 29.1 5.8 166

12 2 25 51.5 16.2 J. Wood and C. L.Denis, personalcommunication

6 0 11 32.5 6.0 J.WoodandC.L.Denis, personalcommunication

34 0 2 2.8 2.0 J. Wood and C. L.Denis, personalcommunication


20 7 55 72.0 20.7 J. M. Ivy, personalcommunication

I

1

Continued on following page

adrl-hom2

adrl-petl4

sec7-cdc37

sir4-hom2

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TABLE 4-ContinuedSegregation (no.) Ascus type (no.) Map distance

Interval ReferenceFD SP PD NPP T x' (cM) SE

56 0 26 16.2

19 10 54 104.9

22 0 9 14.6

24 0 6 10.1

19 0 10 17.3

16 6 51 70.6

67 0 89 28.7115 0 44 14.054 8 104 49.4109 0 56 17.132 19 110 100.155 2 42 28.120 1 49 39.556 11 112 56.1

138 0 5 1.8

29 5 59 52.5

pep7-lys4

pep7-trp4

pep7-ade8

SUF3-trp4SUF3-ade8spt3-SUF3spt3-trp4spt3-ade8cupl4-ade8

cupl4-trp4

snfl-rna3snfl-ade8pho8-ade8pho8-rna3snfl-ssn2

a See footnote a, Table 1.b See footnote b, Table 3.

15 2 41 47.2

83 0 33 14.5

46 5 60 44.3

124 0 53 15.187 0 69 22.349 0 1 1.014 0 9 19.753 0 37 20.968 3 73 32.4

42 8 106 53.2

87 0 11 5.632 2 59 38.8

218 13 157 33.1213 1 35 8.315 0 6 14.4

2.8 J. M. Ivy, personalcommunication

48.8 J. M. Ivy, personalcommunication

4.2 E. L. Dowling, Ph.D.thesis


4.5 E. L. Dowing, Ph.D.thesis


2.1 551.9 557.5 551.9 55

28.7 555.7 735.0 739.4 Y. Kassir, M. Kupiec,

A. Shalom, and G.Simchen, personalcommunication

0.8 Y. Kassir, M. Kupies,A. Shalom, and G.Simchen, personalcommunication

11.1 Y. Kassir, M. Kupiec,A. Shalom, and G.Simchen, personalcommunication

9.0 E. Jones and M.Kolodny, personalcommunication



1.8 562.1 561.0 2055.2 2052.8 2054.6 J. Welch and S. Fogel,

personalcommunication

7.8 J. Welch and S. Fogel,personalcommunication

1.6 225.5 224.4 861.8 865.1 21

Schekman, personal communication), which places it veryclose to makl4 (8.9 ± 2.9 cM distal to thr4). Two Tylsequences have been located on chromosome III, one distalto leu2 and the other just distal to pet]$ (92). Finally, themarker SAD] was shown to be due to a structural change

and not to a point mutation (87) and therefore has beenremoved from the map.A circular chromosome resulting from an exchange be-

tween HML on the left arm and MAT] on the right arm hasbeen isolated (178). This section has a genetic length of 81

sir4-lys4

hom2-lys4

rad9-petl4

rad9-lys4

petl4-lys4

rad9-ade8

SUF24-petl4SUF24-lys4SUF24-trp4petl4lys4pet14-trp4gcn2-lys4gcn2-petl4cdc4O-lys4

cdc4O-trp4

cdc4O-ade8

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TABLE 5. New tetrad analysis data for chromosome V"

Segregation Ascus type Map distanceInterval (no.) (no.) Reference

FD SD PD NPD T x' (CM) SE

162 0 36 9.2 1.4141 0 11 3.6 1.1209 0 5 1.2 0.545 0 11 9.9 2.7

213 0 2 0.5 0.4157 0 3 1.0 0.620 0 1 2.5 2.8138 1 44 13.8 2.456 3 100 37.6 3.9130 0 36 11.0 1.7

79 29 14.6 2.533 8 127 54.9 6.6

92 3 1.6 0.945 0 9 8.4 2.647 0 15 12.6 3.186 0 12 6.4 1.9

97 7 3.4 1.322 3 69 47.5 6.652 0 9 7.4 2.3

61 0 061 0 062 0 27 15.2 2.580 0 9 5.1 1.624 3 38 46.7 12.048 0 11 9.8 3.037 0 15 14.5 3.222 0 30 28.9 3.565 23 133 88.3 23.2610 1 9 1.4 1.0562 2 41 4.5 1.0158 12 234 39.3 3.3

8 0 1 5.8 6.017 0 0 010 0 0 0

32 0 13 14.5 3.441 0 4 4.5 2.218 3 48 50.4 10.0

197 0 1 0.3 0.3

555555

123123123123123J. Welch and S. Fogel, personal communicationJ. Welch and S. Fogel, personal communicationJ. Welch and S. Fogel, personal communicationJ. Welch and S. Fogel, personal communicationJ. Pringle, personal communication7368C. Fields and R. Schekman, personal communicationC. Fields and R. Schekman, personal communicationC. Fields and R. Schekman, personal communication10; F. Boutelet and F. Hilger, personal communication10; F. Boutelet and F. Hilger, personal communication10; F. Boutelet and F. Hilger, personal communication120120127127127127180180180180686879

Schild and Mortimer, in pressSchild and Mortimer, in press116205

a See footnote a, Table 1.

cM. The isolated circular chromosome has a contour lengthof 62.6 ,um, which corresponds to 180 kb. The ratio ofphysical size to genetic map length of this region of chromo-some III is 2.2 kb/cM.The whole of chromosome III has been resolved by

OFAGE. The size of this chromosome is approximately 370kb (19, 163; Carle and Olson, in press) and its total maplength is 140 cM. Therefore, the ratio of physical size togenetic map length for all of this chromosome is 2.5 kb/cM.

Chromosome IV

Chromosome IV is the longest chromosome of yeast asdetermined by genetic mapping procedures (137). It is alsoone of the largest chromosomes as identified by OFAGE (19,163; Carle and Olson, in press).

Since our last major review (137), 22 genes have beenlocated on chromosome IV and several other changes havebeen made. The most distal gene on the left arm of thischromosome is a SUC gene (G. Kawasaki, Ph.D. thesis,University of Washington, Seattle, 1979) which has sincebeen shown to be SUC5 (20). SUF25 has been locatedbetween HO and cdc9 (55, 56), whereas the order, proximal

to distal, of cdc9, ste7, and cdc39 has been changed tocdc9-cdc39-ste7 (14, 149). The sporulation-deficient genespoT4 maps distal to marl on the left arm (94). Two genes,PHS1 and erg6, map very close to trpl near the centomere ofthis chromosome (122, 179).On the right arm, two temperature-sensitive lethals,

tsmOO80 and tsmS162, and a UGA suppressor, SUP88, mapdistal but close to trpl (10, 68; F. Boutelet and F. Hiuger,personal communication). Another UGA suppressor,SUP80, is further out on the right arm near radS5 (68). Thesterile mutation steS maps 16.7 cM proximal to arol (166)and has been shown to be allelic to nul3 (J. Shuster, personalcommunication), which maps in the same location. Thegenes secl, secS, cdc37, and sec7 are located in that orderbetween 2.5 and 6.7 cM distal to hom2 (166; Fields andSchekman, personal communication). sufl2 (55), as well assup35 (129) and arg82 (72), map in the same region but thesegenes have not been tested for linkage against each other oragainst the secl-secS-cdc37-sec7 group. The gene adri isnear petl4 but the order of these two genes relative to thecentromere is unknown (J. Wood and C. Denis, personalcommunication). rad5O and foll, which previously had beenmapped to chromosome IV, have been found to map on

SUFl9-canlSUF19-maklOSUF19-minlrad23-ura3rad23-cyc7rad23-anplanpl-cyc7anpl-ura3cupS-canicupS-ura3cupS-cen5cupS-hom3glc3-cen5gcn4-ura3SUP71-ura3sec3-ura3sec3-cen5sec3-hisltsmOO39-ura3tsmOO39-cen5tsmOO39-arg9GAL83-ura3GAL83-arg9gln3-gcn4gln3-hislgln3-arg6gln3-trp2NHSI-arg9NHSI-hom3NHSI-hislNHSI-trp2SUP85-hislSUP85-SUP20arg84-arg6met6-ilvlmet6-trp2pet-tsl402-ilvlspt2-rad4

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TABLE 6. New tetrad analysis data for chromosome VI"



actl-cdc4 15 3 49 52.6 10.1 49actl-tub2 22 0 0 0 J. Thomas, S. Falco, and D. Botstein, personal commu-

nicationtub2-cdc4 16 2 59 46.9 6.3 J. Thomas, S. Falco, and D. Botstein, personal commu-

nicationtsmOO70-cdc4 19 0 13 20.4 4.4 10; F. Boutelet and F. Hilger, personal communcationtsmOO70-cen6 16 16 30.3 7.1 10; F. Boutelet and F. Hilger, personal communicationtsmOO70-his2 13 2 17 53.6 26.4 10; F. Boutelet and F. Hilger, personal communicationsec4-cdc4 157 0 4 1.3 0.6 C. Fields and R. Schekman, personal communicationsec4-cen6 150 10 3.2 1.0 C. Fields and R. Schekman, personal communicationTyl-his2 25 1 24 30.7 7.8 92Tyl-cdc4 33 0 2 2.9 2.1 92SUF20-cen6 252 24 4.5 0.9 55SUF20-SUP1 265 0 7 1.3 0.5 55SUF20-cly3 234 0 40 7.4 1.1 55SUF20-SUF18 14 0 3 9.0 4.9 55SUF18-cen6 165 67 15.8 1.8 55SUF18-SUP11 41 0 9 9.1 2.8 55SUF18-his2 31 0 15 16.4 3.5 55SUF18-cly3 222 0 5 1.1 0.5 55SUP57-his2 31 0 15 16.4 3.5 101SUP57-metlO 46 0 0 0 101SUP57-SUP6 7 0 2 11.4 7.6 101


chromosome XIV (93; J. Game, personal communication).The rad gene that was mapped on chromosome IV appearedspontaneously in a cross in which radSO was segregating andwas incorrectly identified as this gene. foll had previouslybeen shown to be linked to rad5O (J. Game and J. Little,personal communication). rad9 and sir4 map in the regionbetween petl4 and lys4 (E. L. Dowling, Ph.D. thesis,University of California, Berkeley, 1982; J. M. Ivy, personalcommunication) but their relative order is unknown. Distalto lys4 are SUF24 (55) and gcn2 (73). The former gene is aframeshift suppressor, whereas gcn2 is involved in thecontrol of general amino acid biosynthesis. pep7 maps 14.5cM proximal to trp4 (E. Jones and M. Kolodny, personalcommunication), and close, but distal, to trp4 is the cellcycle mutation cdc40 (Y. Kassir, M. Kupiec, A. Shalom,and G. Simchen, personal communication), whereas furtherout on the right arm are SUF3 (56) and spt3 (205). The orderof these two genes relative to trp4 is unknown. Near the rightterminus of this chromosome have been added cupl4 (J.Welch and S. Fogel, personal communication), snfl (22),pho8 (86), and ssn2 (21). The former gene is proximal torna3; the other three are distal.

Chromosome VChromosome V was originally identified by the

centromere-linked gene ura3. Since 1980, 13 additionalgenes have been located on this chromosome. SUFJ9 mapsbetween min] and maklO (55), but the position of SUSJ,which maps in the same region, relative to these three geneshas not been determined. rad23 and anpi map close to cyc7(123). cyc7 is one of two structural genes for cytochrome csynthesis and, interestingly, it has been pointed out (123)that the other gene, cycl, which is on chromosome X, hasclose to it rad7, another excision-defective gene similar torad23. The gene cupS maps between anpi and ura3 (Welchand Fogel, personal communication). The following three

genes have been mapped near the centromere of chromo-some V: glc3 (J. Pringle, personal communication), gcn4(73), and sec3 (Fields and Schekman, personal communica-tion). mnnl and SUP71 had already been mapped in thisregion. mnnl failed to recombine with the centromere in 41tetrads, whereas SUP71 is 2.5 cM from the centromere onthe right arm (130). glc3 is 1.6 cM from the centromere, sec3is 3.4 cM from the centromere and on the right arm, and gcn4is 8.4 cM from ura3, which would place it near thecentromere (ura3 is 8.0 cM from the centromere). None ofthese five genes has been tested for linkage against eachother, so their order relative to each other or the centromereis unknown.

gln3 is 9.8 cM proximal to his] (127) and the gene NHS1,which is involved in H2S production, is 1.4 cM proximal tohom3; this places it very close to BORI (180). SUP85 mapsdistal to his] (68) and fails to recombine with SUP19,20,which also have been mapped in this region. arg84 fails torecombine with arg6; it is believed to be a mutation in thepromoter region of the argS arg6 gene complex (79).

In previous editions of the map, metS was placed betweentrp2 and radSI (132). We have now shown that this gene ismet6 and that metS maps on the right arm of chromosome X(Schild and Mortimer, in press). eth2 (sam2) had been shownto be linked to met6, so it has been assigned to chromosomeV as well. It is provisionally placed distal to met6 but it couldalso be proximal. A temperature-sensitive genetic petite,pet-ts1402, maps 50 cM distal to ilvi (116). spt2 is less thana map unit from rad4, located much further out on the rightarm of this chromosome (205). Recent data place rad24 (r1s)distal to rad3 (Game, personal communication).

Chromosome VIChromosome VI is one of the shortest in genetic map

length. It is also the second smallest chromosome as deter-mined by OFAGE (Carle and Olson, in press). Its map length

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MICROBIOL. Rfv.

TABLE 7. New tetrad analysis data for chromosome VII'Segregation Ascus type Map distance

Interval (no.) (no.) Reference

spoTi2-adeSspoT2-lys5rad54-lys5rad54-metl3cup2-lys5cup2-aro2cup2-metl3SUP38-ade5SUP38-lys5SUP38-cyh2mf4x2-lysSmfa2-aro2mfa2-metl3mfa2-cyh2mfos2-trp5hemlO-leulhem10-olelhemlO-trp5SUP,54-trp5SUP54-leulSUPS4-cen7atel-leulatel-trp5smr2-cen7smr2-cyh3tsmOI19-antltsmOl19-leulanti-leulTyl-leulrmel-trpSrmel-leulrmel-ade6supll2-trp5s;,upll2-ade6alg7-trp5alg7-cly8cdc62-leulcdc62-ade6ade6-cly8ade6-SUF4r-1y8-SUF4SUF4-ser2SUF4-ade3SUF15-ser2SUFIS-ade3ser2-ade3hipl-ade3SUP76-ade3SUF76-SUP77SUP77-ade3

FD SD PD NPD T x' (cM)

27 0 7 10.412 2 26 51.343 0 15 13.042 0 39 24.482 0 12 6.477 1 34 18.134 1 54 33.95 2 13 NL10 0 13 29.520 0 2 4.747 6 92 46.749 5 87 43.486 1 52 21.0

206 1 26 7.06 1 17 50.29 0 8 23.7

3.314.0

44 0 6 6.129 0 22 21.6

24 17 24.046 0 2 2.131 0 6 8.2

101 22 9.424 0 0 078 0 6 3.681 0 3 1.879 0 5 3.088 0 21 9.925 0 35 29.644 0 16 13.439 0 19 16.918 2 38 44.873 0 15 8.624 1 18 29.18 0 12 30.15 0 32 43.3

27 1 12 24.628 3 58 44.320 8 61 75.548 2 39 29.757 21 161 73.869 17 172 60.0160 0 18 5.1142 0 30 8.9159 0 11 3.3

10.820 0 17 23.022 0 25 27.25 7 25 NLd

SE

3.615.52.93.01.73.74.2

6.43.46.96.33.01.9

16.26.3

2.43.55.41.53.11.9

1.41.11.32.13.52.93.59.32.09.65.62.9

12.17.4

21.16.4

12.68.11.11.51.0

4.24.1

M. Tsuboi, personal communicationM. Tsuboi, personal communicationE. L. Dowling, Ph.D. thesisE. L. Dowling, Ph.D thesisJ. Welch and S. Fogel, personal communicationJ. Welch and S. Fogel, personal cornmunicationJ. Welch and S. Fogel, personal communicationB. Ono, personal communicationB. Ono, personal communicationB. Ono, personal communicationS. Caplan and J. Kurjan, personal communicationS. Caplan and J. Kurjan, personal communicationS. Caplan and J. Kurjan, personal communicationS. Caplan and J. Kurjan, personal communicationS. Caplan and J. Kurjan, personal communication193D. Urban-Grimal and R. Labbe-BoisbD. Urban-Grimal and R. Labbe-Boisb101101101160160505010; F. Boutelet and F. Hilger, personal communication10; F. Boutelet and F. Hilger, personal communication2992156156156B. Ono, personal communicationB. Ono, personal communication66P. J. flanic-Joyce, in pressP. J. Hanic-Joyce, in press2323233838555555J. Tanaka and G. R. Finkc686868

a See footnote a, Table 1.b Abstr. Int. Conf. Yeast Genet. Mol. Biol, 11th, p.87, 1982.c Abstr. Mol. Biol. Yeast Meet., Cold Spring Harbor, N.Y., 1983.d NL, Not linked.

and physical size are 140 cM and 290 kb, respectively. Ourfirst review positioned 13 genes along this chromosome; 8genes have been added since. The structural genes for actinand P-tubulin, which fail to recombine, are now the distal-most markers on the left arm and are located about 50 cMfrom cdc4 (49; J. Thomas, S. Falco, and D. Botstein,personal communication). These genes have been shown tobe separated by approximately 1.5 kb ofDNA. In the regionbetween these two genes is a sequence with close homologyto the human ras haslbas oncogene (57). tsmOO70 maps 20.4

± 4.4 cM distal to cdc4 (10; F. Boutelet and F. Hilger,personal communication), which places it close to SUF9which in turn is 16.5 ± 5.7 cM distal to the same gene (136).sec4 is about 1 map unit proximal to cdc4 (Fields andSchekman, personal communication), whereas a Tyl ele-ment is 3 cM proximal to the same gene (92). SUF20 mapsbetween SUPJl and the centromere on the right arm,whereas SUF18 is 1.1 cM proximal to cly3 further out on thischromosome arm (55). SUP57, a UAG suppressor, fails torecombine with metlO; this places it very close to SUP6



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TABLE 8. New tetrad analysis data for chromosome VIII"Segregation Ascus type Map distance

Interval (no.) (no.) ReferenceFD SD PD NPD T x' (cM) SE

dur3-spoll 186 1 62 13.7 1.9 T. Cooper and M. Mojumdar, personal communicationdur3-cen8 144 105 24.5 2.2 T. Cooper and M. Mojumdar, personal communicationdur3-arg4 129 3 117 27.4 2.7 T. Cooper and M. Mojumdar, personal communicationdur3-dur4 0.4 T. Cooper and M. Mojumdar, personal communicationspoll-cen8 98 151 39.8 3.3 T. Cooper and M. Mojumdar, personal communicationspoll-arg4 90 4 155 36.3 2.9 T. Cooper and M. Mojumdar, personal communicationardl-arg4 52 0 1 1.0 1.1 M. Whiteway and J. Szostak, personal communicationardl-cen8 49 25 18.9 3.5 T. Cooper and M. Mojumdar, personal communicationtsm0186-arg4 39 0 5 5.7 2.5 10; F. Boutelet and F. Hilger, personal communicationtsmO186-thrl 39 0 5 5.7 2.5 10; F. Boutelet and F. Hilger, personal communicationtsmO15l-arg4 37 0 7 8.0 2.8 10; F. Boutelet and F. Hilger, personal communicationtsmOI51-thrl 42 0 2 2.3 1.7 10; F. Boutelet and F. Hilger, personal communicationtsmOI51-tsmO186 40 0 4 4.6 2.3 10; F. Boutelet and F. Hilger, personal communicationput2-arg4 31 1 21 26.3 7.7 11put2-thrl 40 0 11 10.8 2.9 11put2-CUPI 41 1 10 17.4 10.4 11supllH-arg4 31 2 52 38.4 6.1 B. Ono, personal communicationsup]ll-thrl 23 0 24 26.1 4.1 B. Ono, personal communicationspol2-pet3 75 0 0 0 89


(approximately 2 cM from metlO), although these two sup- is remarkable in that all other mapped fermentation genespressors recombine (101). (reviewed in reference 132) and many of the glycolytic genes

(109, 114, 115; Z. Lobo, Ph.D. thesis, University ofChromosome VII Bombay, Bombay, India, 1976) are also on or near the ends

Chromosome VII one of the largest yeast chromosomes; of chromosomes.only IV and XII are larger and XV is near the same size On the left arm, adeS has been changed to adeS,7 (157) and(Carle and Olson, in press). Until recently, the genetic map spoT2 has been shown to map proximal to this gene (191).length of this chromosome was uncertain because a group of radS4 (E. L. Dowling, Ph.D. thesis), cup2 (Welch and Fogel,genes had been placed on the right arm only by mitotic personal communication), and cdc43 (A. Adams and J.crossing-over procedures (157). These studies indicated that Pringle, personal communication) map distal to lysS but theMALI/SUCI were the most proximal genes of this group. It order of these genes relative to lysS and ski) is unknown.has recently been shown that SUF4, which was the most sup38 maps distal to cyh2 (B. Ono, personal communica-"distal" mapped gene relative to MALI/SUCI, is in fact tion), whereas the structural gene for the ot mating typeproximal to MALJ/SUCI and linked to cly8 (23), which is factor, mfa2, maps 7.0 cM proximal to this gene (S. Caplanabout 45 cM from the centromere (130). Thus, this group of and J. Kurjan, personal communication). The gene hemlO isgenes from MALl/SUCI to SUF4 has been reversed, placing 3.3 cM from olel (Urban-Grimal, personal communication),MALl/SUCI as the most distal genes on the right arm. There but its order relative to rad6, which maps in the same region,is evidence that SUCI is the distal-most marker and is is uncertain. SUP54 maps 6.1 cM from trpS (101) and isadjacent to the telomere on the right arm of chromosome VII probably distal to this gene, whereas ate) is 2 cM from leul,(J. Celenza and M. Carlson, personal communication). This also probably distal (160). tsmOIl9 is also near leul (Boutelet

TABLE 9. New tetrad analysis data for chromosome IX'Segregation Ascus type Map distance

Interval (no.) (no.) ReferenceFD SD PD NPD T x' (cM) SE

rev7-SUC2 2 10 30 NLb C. W. Lawrence, personal communicationrev7-SUC2 11 0 14 28.1 5.1 C. W. Lawrence, personal communication

Total 13 10 44 NL

rev7-hisS 84 0 27 12.2 2.1 C. W. Lawrence, personal communicationrev7-lysJl 24 3 41 46.5 11.0 C. W. Lawrence, personal communicationlysil-hisS 16 1 27 38.2 8.4 C. W. Lawrence, personal communicationtsmO139-lys1l 28 1 26 29.7 7.1 10; F. Boutlet and F. Hilger, personal communicationtsmO139-his6 14 1 17 37.6 12.7 10; F. Boutlet and F. Hilger, personal communicationdal81-his6 69 10 126 49.2 6.9 192dal8l-dall 181 2 22 9.2 3.6 192dal81-dal4 172 1 32 9.4 2.1 192

a See footnote a, Table 1.b NL, Not linked.

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TABLE 10. New tetrad analysis data for chromosome XaSegregation (no.) Ascus type (no.) Map distance


50 081 0120 014 124 049 029 87 2

27 06 0

35 013 0

52 362 0

35 8

43 0

41 0

7 0

10 0

18 0

68 0

589 0266 0186 0504 49 0

32

63

2

0

11 9.163 21.929 9.931 40.837 30.848 25.082 61.644 54.438 29.60 09 10.37 17.6

2.818 11.6

9.8

1

3

1.2

3.5

14 33.5

7 20.8

1

9

2209

25013

2.8

5.9

1.80

2.318.129.7

48 37.5

5 3.7

2.5 32.1 421.7 427.7 423.4 722.7 72

11.9 729.3 723.3 72

683.1 685.5 681.6 1212.6 1213.3 10; F. Boutelet and

F. Hilger,personalcommunication

1.3 10; F. Boutelet andF. Hilger,personalcommunication

2.1 10; F. Boutelet andF. Hilger,personalcommunication

5.3 C. W. Lawrence,personalcommunication




0.4 5656

0.8 551.2 55, 565.4 Schild and

Mortimer, inpress

6.5 Schild andMortimer, inpress

1.6 Schild andMortimer, inpress


and Hilger, personal communication). There are now ninegenes that map very close to leul; most ofthese genes conferresistance to drugs (BOR2, borrelidin; cyh3 cycloheximide;till, thiaisoleucine; anti, antibiotic; AMY], antimycin; olil,oligomycin; AXE), axenomycin). It has been proposed thatthese genes are alleles or are part of a genetic complexcoding for ribosomal proteins (159). smr2, a mutation thatconfers resistance to the herbicide sulfometuron methyl,fails to recombine with cyh3 (50).A transposable element, Tyl, is on the right arm about 10

cM from leul (92), and rmel, which is possibly an allele ofdsml (156), is 13.4 cM from leul. alg7 shows linkage to trpS

and cly8 and appears to map in the region between ade6 andthe centromere (6). supl]l2 maps 8.6 cM distal to ade6 (Ono,personal communication) and cdc62 is near cly8 but itsposition relative to ade6 is unknown (Hanic-Joyce, in press).The genes SUF4 (56), SUP77 (68), SUP76 (68), and hip) (J.Tanaka and G. R. Fink, Abstr. Mol. Biol. Yeast Meet., p.284, 1983) had all been mapped distal to ade3, but, asdiscussed above, they are now proximal. hip) also is linkedto ade3 but it is not known if it is proximal or distal to thisgene (Tanaka and Fink, Abstr. Mol. Biol. Yeast Meet.,1983). As mentioned earlier, SUF4, formerly the most distalof this group, maps only 29 cM from the proximal marker

eaml-inolinol-cdc6inol-ura2inol-arg3arg3-SUP7arg3-ura2arg3-cdc6ura2-SUP7ura2-cdc6SUP74-SUP7SUP74-SUP73SUP73-SUP51cyrl-cenlOcyrl-ilv3tsmO185-cenlO

tsmO185-met3

tsmO185-ilv3

rev5-met3

rev5-ilv3

revS-cycl

rev5-cdc8

SUF23-cyclSUF23-rad7SUF23-SUP4SUF23-cdcllmetS-cdcll

hom6-cdcll

metS-hom6

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TABLE 11. New tetrad analysis data for chromosome XI'



SUP33-fasl 101 0 13 5,7

SUP33-trp3

lapl-mak9lapl-fas)lapl-lap4lapl-urallapl-trp3lap4-mqk9lap4-fas)SUP75-trp3

SUP75-ural

SUP75-metl4

SUP58-cenllSUP58-metl4SUP58-metlgcn3-met14dal8O-cdcl6dal8O-metl4dal8O-metlspoT23-metl4spoT23-metlspo)4-cdcl6

spol4-metl4

met2O-metlsirl-metl

sirl-metl4

sirl-maklS

sirrl-MAL4

SUP-lA-metlSUJP-)A-MAL4

19 33

122 0 106 23.4

25 1 13 26.3132 0 24 7.925 0 9 13.329 5 103 50.123 4 77 50.46 1 10 53.38 0 9 26.65 1 14 53.5

10 1 25 44.0

11 6 21 NLb

42.820 3 30 51.310 7 32 144.895 0 10 4.8104 1 66 21.1127 0 44 13.052 4 115 41.459 0 35 18.619 0 39 34.19 0 15 31.4

28 0 20 20.9

175 0 2 0.631 1 35 31.0

7 2 59 53.1

30 1 19 26.0

52 0

18 274 0

0 0

56 45.60 0

1.5 B. Ono, persopalcommunication

1.7 B. Ono, personalcommunication

12.0 1901.6 1903.9 1906.0 1907.2 190

30.5 1906.3 190

21.0 D. Hawthorne, personalcommunication

10.0 D. Hawthorme, personalcommunication

D. Hawthorne, personalcommuqication

7.5 10117.0 101

145.9 1011.5 1562.6 1921.8 1924.2 1922.5 1913.4 1915.1 M. Townsend, B. diDomenico,

S. Klapholz, and R.Esposito, personalcommunication

3.6 M. Townsend, B. diDomenico,S. Klapholz, and R.Esposito, persqnalcomnilunication

0.4 1195.7 J. M. Ivy, personal

communication6.8 J. M. Ivy, personal

communication8.3 J. M. Ivy, personal

communicationJ. M. Ivy, personalcommunication

6.5 1717

I See footnote a, Table 1.b NL, Not linked.

cly8. SUFI5 (55) and fol2 (J. Game, J. Little, and B.Rockmill, personal communication) also have been locatedin the SUF4-SUCJ/MALI interval.

Chromosome VIIIChromosomes VIII and V, which are about the same size

(580 kb), are in the group of medium-sized chromosomes asdetermined by OFAGE (Carle and Olson, in press). Themeiotic length of the region between mak7 (distal-most onthe left arm) and makl9 (distal-most on the right arm) hasbeen determined by collective tetrad analyses tQ be about180 cM. mak2O had been assigned to chromosome VIII bytrisomic analyses and to the left arm because of a lack ofmitotic linkage to right arm markers (198). Although trisomicanalysis has been used successfully to map many genes, it isnot always reliable. In view of this, the assignment of mak2O

to this chromosome most be considered provisional. Thegenes dur3 and dur4, which fail to recombine, map 24.5 cMfrom the centromere and 13.7 cM distl to spoil on the leftarm (T. Cooper and M. Mojumdar, personal communica-tion). On the right arm, ardl is 1 cM proximal to arg4 (M.Whiteway and J. Szostak, persontl communication), andspol3 has now been shown to be proximal, rather thandistal, to arg4 (S. Klapholz and R. E. Esposito, personalcommunication). Two temperature-sensitive lethals,tsmQl86 and tsmO151 map in the agg4-thrl interval (F.Boutelet and F. Hilger, personal communication). put2 is 2.9cM distal to thr4 (11) and suplHl appears to map close toCUP] (Ono, personal co'mmunication) but its position, prox-imal or distal, to this gene is unknown. spol2 fails torecombine with pet3 and, in addition, the order of SUF8 andpet3 on this chromosopne has been reyersed (89). Finally,

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TABLE 12. New tetrad analysis data for chromosome XII"



pprl-cenl2 55 8 6.6 2.2 104pprl-aspS 54 0 9 7.2 2.3 104SUP37-asp5 8 2 18 62.4 28.3 B. Ono, personal

communicationSUP37-gaI2 12 1 23 41.4 10.3 B. Ono, personal

communicationpep3-gal2 71 9 192 46.6 4.1 206pep3-RDNI 218 1 53 10.9 1.7 206gal2-RDNI 61 17 194 60.0 7.0 206ilv5-RDNI 19 5 22 118.5 181.8 148iIvS-ura4 48 10 82 60.8 13.9 148ilvS-car2 12 6 34 98.5 52.7 148ura4-car2 23 0 21 24.5 4.3 148nam2-ura4 18 0 13 21.1 4.5 46SUP86-ura4 24 0 0 0 68cup3-ura4 86 2 37 20.4 4.8 J. Welch and S. Fogel,

personal communicationcup3-car2 14 0 6 15.1 5.3 J. Welch and S. Fogel,

personal communicationmar2-ura4 29 0 48 31.5 3.0 A. Klar, personal

communlicationmar2(ste8)-ura4 26 2 29 38.3 10.7 R. K. Chan, personal

communicationTotal 55 2 77 33.5 3.9

sst2-ura4 14 1 12 36.4 17.6 R. K. Chan, personalcommunication

sst2-mar2(ste8) 23 0 4 7.5 3.6 R. K. Chan, personalcommunication

See footnote a, Table 1.

Celenza and Carlson (personal communication) have deter-mined by the spoll mapping method that SUC7 is onchromosome VIII but have not localized the gene on thischromosome.

Chromosome IXThere have been relatively few changes on chromosome

IX since our last review. Chromosome IX is the fourthsmallest chromosome (460 kb) as determined by OFAGE(Carle and Olson, in press) and it is still rather sparselypopulated with genes. rev7 has been mapped against SUC2,hisS, and lysll, genes already located on the left arm, andappears to be located close to SUP22 (C. Lawrence, per-sonal communication). Since rev7 has not been mappedagainst this suppressor, the order of these two genes relativeto outside markers is unknown. The temperature-sensitivelethal tsmOl39 maps between lysli and his6 (Boutelet andHilger, personal communication) and is probably locatedproximal to SUP17. Finally, two genes have been added tothe dal cotnplex on the right arm; dal8J, a regulatory site,maps proximal to the complex, whereas dal3 maps distal(192).

Chromosome XIn order of increasing physical size, chromosome X ranks

seventh (Carle and Olson, in press). Its genetic map lengthfrom cdc6 on the left arm to hom6 on the right arm isapproximately 200 cM. The killer maintenance gene makl7has been mapped as the distal-most gene on the left arm bymitotic crossing-over procedures (198). It fails to showmeiotic linkage to ura2 but has not yet been tested for

meiotic linkage to cdc6 which is more distal. The genes eami(3) and inol (42) map between cdc6 and ura2 but their order,relative to these two flanking mnarkers, is unknown. arg3maps in the ura2-SUP7 interval about midway betweenthese two genes (72). SUP74, a UGA suppressor, fails torecombine with SUP7, an ochre suppressor (68). It seemsunlikely that these suppressors are alleles; SUP7 is a tyro-sine-inserting ochre suppressor and it is improbable that agene coding for tRNATYr could mutate to recognize a UGAcodon. SUP73, another UGA suppressor, maps 10.3 cMproximal to SUP74 (68). cyrl defines the adenylate cyclasegene (121) and tsmOJ85, which maps in the same region andis allelic with cdc35, is deficient in adenylate cyclase activity(Boutelet and Hilger, personal communication). revS (Lawr-ence, personal communication) and SUF23 (55, 56) havebeen added to the cluster of genes that map very close tocycl, the structural gene of iso-1-cytochrome c. revS appearsto be the most proximal of this group and SUF23 failed torecombine in 266 tetrads with rad7, which is also in thisgroup (55, 56). Two new genes have been added to the distalregion of the right arm of chromosome X. metS and hom6,which are only 3.6 cM apart, map 37.5 cM distal to cdcll(Schild and Mortimer, in press). The prefetrred order of thesetwo genes places met5 proximal, but this is still uncertain.

Chromosome XIChromosome XI is unusual in that a large group of linked

genes, extending from mak9 to cly7, has been located on thischromosome only by trisomic analyses, and this association,at least for ural which is part of this group, has been knownfor at least 25 years. Remarkably, no meiotic linkage be-

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TABLE 13. New tetrad analysis data for chromosome XIII"

Segregation (no.) Ascus type (no.) Map distanceInterval Reference


45 18.2 3.023 26.8 3.813 21.1 4.50 0

6 13.8 4.930 42.5 8.236 32.7 5.8

Schild and Mortimer, in pressSchild and Mortimer, in pressSchild and Mortimer, in pressSchild and Mortimer, in press68Schild and Mortimer, in press

SUF22-SUF7SUF22-cenl3SUF7-cenl3tsmOIll-cenl3

74 10379 9826 1

tsmOII-tsmO800

tsmOIJJ-arg8O

tsmO800-cenl3

tsmO800-arg8O

secS9-radS2

secS9-cenl3

radS2-cenl3

spoTI-cenl3

165 0

43 0

18 0

9

9

22

90 25

79 0

58 0

34

71

100 8

61 44

224 0

spoTI-lys7

spoTI-cdcS

14 1

13 0

mcml-lys7

adh3-lys7

SMRI-ilv2SMRI-lys7pet-ts2858-ilv2pet-ts2858-rnalilv2-rnal

supl13-cenl3

supl13-lys7

ilv2-lys7ilv2-lys7

Total

ilv2-arg8OSUP78-SUP8spoT7-rnal

31 039 135 114 412 5

11 27

21 0

82 514 1

96 6

9 0

5 0245 0

spo7T7-mak27

prcl-rnal

118 0

36 8

prcl-SUP8

ade4-rnalade4-SUP8cdc61-rnalcdc6)-ade4

See footnote a, Table 1.

79 0

19 116 018 115 0

22

6

0

44183339

2.6 0.9 5537.4 3.6 5534.8 3.3 551.9 1.9 10; F. Boutelet and F. Hilger,

personal communication8.7 2.7 10; F. Boutelet and F. Hilger,




personal communication27.7 2.3 C. Fields and R. Schekman,



personal communication0 M. Tsuboi, personal

communication38.9 10.3 M. Tsuboi, personal


communication2 G. Maine and B. Tye,

personal communication16 M. Ciriacy, personal

communication0

30.023.266.975.2

Unlinked

15

11941

160

1086

20

113

30

49112126

20.9

37.142.3

38.2

504.6 507.8 116

23.9 11625.7 D. Hawthorne, personal

communicationB. Ono, personalcommunication

4.2 B. Ono, personalcommunication

4.1 1486.1 D. Hawthorne, personal

communication3.4

26.5 5.9 14831.0 7.1 681.2 0.5 M. Tsuboi, personal


communication54.8 7.5 E. Jones, M. Aynardi, and M.

Kolodny, personalcommunication

14.0 2.3 E. Jones, M. Aynardi, and M.Kolodny, personalcommunication

40.1 5.1 Schild and Mortimer, in press20.5 4.8 Schild and Mortimer, in press34.8 9.8 Hanic-Joyce, in press31.8 3.8 Hanic-Joyce, in press

GAL80-radS2GAL80-SUP79rad52-SUP5GA,L80-SUP5SUP79-radS2SUP79-rad52

Total

95 120 018 011 016 012 128 1

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TABLE 14. New tetrad analysis data for chromosome XIVaSegregation (no.) Ascus type (no.) Map distance


43.43.8

14.642.2

10.2

43.212.514.0

12.7

7.6 560.9 562.2 564.3 C. Fields and R.


1.6 C. Fields and R.Schekman, personalcommunication

7.2 931.9 935.5 J. Game, personal

communication

5.9 1.2 930 J. Game, personal

communicatioh5.5 1.1

22.40

0

3.7 93J. Game, personalcommunication

J. Game, personalcommunication

41.7 5.0 3512.1 2.2 3546.9 3.9 3591.0 21.0 3566.0 15.4 350 Chang et al., in press12.9 3.4 Williamson et al., in

press4.7 2.3 Williamson et al., in

press8.2 2.9 Williamson et al., in

press16.5 3.3 K. Voelker, S.

DiNardo, and R.Sternglanz, personalcommunication

14.3

44.2

2.5

5.9 K. Voelker, S.DiNardo, and R.Sternglanz, personalcommunication

2.0 2.2 883.6 1.3 L. Robinson and K.

Tatchell, personalcommunication

3.2 1.1

63.7

76.550.895.970.3

64.0

68.5

14.8

13.79.9

95.413.3

14.6

6.8

L. Robinson and K.Tatchell, personalcommunication

9019988K. Voelker, S.DiNardo, and R.Sternglanz, personalcommunication

L. Robinson and K.Tatchell, personalcommunication


SUF6-pha2SUF6-met2SUF6-pet2sec2-pha2

88 9185 0147 158 5

981552126

37

3533S

38

230

23

sec2-met2

radSO-pha2radSO-met2rad5O-met2

Total

rad5O-pet2rad5O-pet2

Total

radS0-petxfoll-pet2

foll-rad5O

sufl4-pet2sufl4-petxsufl4-met4met4-pet2met4-petxleu4-met4pmsl-met4

pmsl-top2

top2-met4

top2-met4

total

top2-pet8

148 0

12 199 013 0

112 0

173 013 0

186 0

26 013 0

29 0

77 78i 0

115 1543 2227 821 032 0

39 0

36 0

90 1

126 1

38 4

26 090 0

116 0

25 7

48 2024 45 2

22 10

24 7

123 43

RASsc2-met4RASsc2rthet4

210

0

12425

261137700

11

4

7

35

42

84

17

8

65

151571194

66

379

Total

RASs2-pet8

met4-pet8met4-pet8met4-pet8met4-pet8

met4-pet8

Total

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TABLE 14-Continued

Segregation (no.) Ascus type (no.) Map distance


spol-pet8 107 0 9 3.9 1.3 90; B. diDomenico andR. Esposito, personalcommunication

spol-rna2 32 0 7 9.0 3.2 90; B. diDomenico andR. Esposito, personalcommunication

pet8-rna2 35 0 5 6.3 2.7 90; B. diDomenico andR. Esposito, personalcommunication

TylaTYlb 88 0 7 3.7 1.4 92Tylapet8 16 0 8 16.8 5.0 92Tylb-met4 20 3 40 49.5 12.0 92Tylb-pet8 28 0 16 18.2 3.7 92Tylb-lys9 33 2 54 37.8 5.9 92pet494-rna2 62 3 123 37.8 3.3 138pet494-lys9 273 0 14 2.4 0.6 138SUP28-pet8 25 1 66 39.3 3.9 B. Ono, personal

communicationSUP28-lys9 86 0 12 6.2 1.7 B. Ono, personal

communicationholl-lys9 4.7 P. Farabaugh, personal

communication


tween any of these genes and other chromosome XI markershas as yet been detected. Very recently, B. Glassner andR. K. Mortimer (personal communication) have shown mi-totic linkage between cdc16 and ural. Added to this group oforphaned genes since our last review (132) are SUP33 nearfasi (Ono, personal communication), lap) and lap4 (190),located between fasi and mnn4, and SUP75, near cly7 (D.Hawthorne, personal communication).SUP58 shows meiotic linkage to metl4, located near the

centromere of this chromosome, and is the proximal-mostmarker on the left arm (101). On the right arm, gcn3 is 4.8 cMdistal to metl4 (73) and dal80 maps 13.0 cM distal to metl4(192). Near dal80 are spol4 (M. Townsend, B. di Domenico,S. Klapholz, and R. E. Esposito, personal communication)and spoT23 (191), which map in the same region; these twosporulation genes may be allelic. The gene met20 maps about0.5 cM from met) (119), and sirl was mapped by theintegrated plasmid method and shown to be very close toMAL4 (J. M. Ivy, personal communication). A cloned DNAfragment that behaves as an amber suppressor, SUP-IA,when present in high copy number, also fails to recombinewith MAL4 (17).

Chromosome XIIChromosome XII is very likely the largest of the yeast

chromosomes. Its meiotic length from makl2 on the left armto car2 on the right arm is approximately 300 cM. Inaddition, this chromosome carries about 120 copies of therDNA genes of yeasts (150); each copy has 9.1 kb of DNA.As yet unexplained is the observation that meiotic re-combination in the rDNA region is only about 1% of thatexpected on the basis of the amount of DNA (150). If wewere to assume normal recombination frequencies in thisregion (ca. 0.3 cM/kb), chromosome XII would have arecombination length of approximately 630 cM. This wouldmake it genetically larger than chromosome IV, which has arecombination length of about 500 cM. It is interesting thatduring OFAGE analysis chromosome XII frequently failed

to leave the sample well, and when it did, it left in a fashionsuggesting to the authors that its movement was restrictedby factors (e.g., nucleolus) other than those governingnormal DNA migration (Carle and Olson, in press). Ouranalysis indicates that chromosome XII probably is thelargest yeast chromosome and suggests that perhaps only thephysical size of this chromosome determines its migrationbehavior.Nine genes have been added to this chromosome since our

last review. These include pprl, which maps between thecentromere and aspS on the right arm (104), and SUP37(Ono, personal communication) and pep3 (E. Jones, per-sonal communication; 206), which map proximal to therDNA cluster. In addition, ilvS (148) maps distal to therDNA genes and shows meiotic linkage to this cluster ofgenes. This linkage analysis was accomplished by integratingthe LEU2 gene into the rDNA sequences and then using thismarker to score these repeated genes. The meiotic distancefrom rDNA to ilvS is over 100 cM. Distal, but linked, to ilv5is a group of genes whose order on the chromosome is stilluncertain. The genes nam2 (46), SUP86 (68), cup3 (Welchand Fogel, personal communication), and mar2 (A. Klar,personal communication) have been added to the three genesalready in this group. The mutations mar2, ste8, and sir3have been shown to be allelic (Klar, personal communica-tion; R. Chan, personal communication). All three affectexpression of the silent copies of the mating type locus.Finally, sstl is 7 cM distal to mar2 (24; Chan, personalcommunication).

Chromosome XIIISeveral changes have been made to the map of chromo-

some XIII since our last major review (132). The genes eth2and met6, which had tentatively been placed distal to lys7(117), have been found to be located on chromosome V neartrp2 (Schild and Mortimer, in press). The "metS" markerthat was mapped in this location was, in fact, met6; metS hasnow been shown to map on chromosome X (Schild and

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TABLE 15. New tetrad analysis data for chromosome XVaSegregation (no.) Ascus type (no.) Map distance

IntervalSD

ReferenceFD SD PD NPD T x' (cM) SE

23.6 2.3 5675.7 13.0 5679.3 8.4 55, 5645 M. Ciriacy, personal

communication27 M. Ciriacy, personal

communication15 J. Pringle, personal

communication71.9 8.6 552.9 0.6 559.9 2.2 G. Fabian and E. Jones,

personal communication9.2 2.2 G. Fabian and E. Jones,





personal communication25.1 4.4 M. Tsuboi, personal



communication0.602.8

11.229.90.7

25.235.59.70.3

33.939.014.17.2

12.8

0.6 99

2.0 92.7 95.3 90.6 1867.0 1862.5 561.1 560.2 562.4 569.0 1822.1 1823.5 881.8

105.4 111.4 888.8 2.2 M. Tsuboi, personal



communication30.8 2.5 5024.5 5.4 5011.8 6.3 16618.0 4.6 16629.9 5.9 D. Barnes and J. Thorner,

personal communication20.5 4.8 D. Barnes and J. Thorner,

personal communication42.8 20.3 D. Barnes and J. Thorner,

personal communication69.8 19.3 Schild and Mortimer, in press28.6 3.9 Schild and Mortimer, in press52.9 24.3 D. Barnes and J. Thorner,

personal communication37.1 7.3


SUFI-arg8SUFI-argiarg8-argladhl-arg8

adhi-argi

glc4-arglSUF17-arg8SUFJ7-arglpepl2-tup4

pepl2-cenlS

pepl2-makl

71 15

113 143 19125 59

86 34375 069 0

61 0

46 0

17 0

9 1

tup4-makl

90150438

2852317

25

40

17

23

tup4-cenlS

makl-cenl5

56 30

76 10

spoTII-petl7

spoTlI-argl

80 7

80 1

69 1

150 192

spoTII-ceniS

tup7-cenlStup7-pho8Otup7-makltup7-tup4tup7-tmpltopl-cenlStopl-petl7sufll-cenlSsufll-petl7sufll-cdc21sufll-ade2RAS"]petl7RASsc1_ade2RASs]ade2

Total

RAScsJ-his3spoTIS-ade2

spoTIS-petl7

spoTIS-his3

84 035 058 013 0

7 0

278 0342 0138 526 287 024 0

111 0

021618

7

662

20135344

38

8 361 0

31 0

12 6

smr3-ade2smr3-his3ste4-his3ste4-met7stel3-his3

stel3-met7

42 067 210 018 011 0

1513

29

53

66393

1015

11

12

462418

42

stel3-cdc3l

cdc31-ade2cdc3l-his3cdc3l-his3

16 0

10 1

Total

12 518 013 2

31 2

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TABLE 15-ContinuedSegregation (no.) Ascus type (no.) Map distance


cdc31-met7 44 0 8 7.7 2.6 Schild and Mortimer, in presscdc3l-met7 17 0 16 25.1 5.1 D. Barnes and J. Thorner,

personal communicationTotal 61 0 24 14.2 2.5

sufl3-his3 43 8 147 51.3 5.4 35sufl3-tra3 128 1 77 20.2 2.2 35sufl3-prtl 48 11 155 55.2 6.4 35his3-gcdl 86 1 113 29.8 2.3 35cdc64-his3 8 10 45 NL Hanic-Joyce, in presscdc64-met7 9 0 20 34.6 4.4 Hanic-Joyce, in presscdc64-prtl 46 1 67 32.1 3.4 Hanic-Joyce, in presscdc66-cdc64 42 0 12 11.2 2.9 P. Hanic-Joyce and D. R.

Carruthers, personalcommunication

cdc66-prtl 11 0 30 36.6 3.5 P. Hanic-Joyce and D. R.Carruthers, personalcommunication

cdc63-prtl 372 0 0 0 fianic-Joyce, in pressphrl-prtl 43 0 16 13.6 2.9 161phr2-phrl 27 0 16 18.7 3.7 112


Mortimer, in press). Fragment V, containing arg8l, SUP5, arm (10; Boutelet and Hilger, personal communication;and GAL80, has been located on the left arm of chromosome Fields and Schekman, persohal commnunication). meml isXIII, using the radS2 chromosome loss mapping method only 2 cM from lys7but the order of these two gends relative(Schild and Mortimer, in press). Independently this group of to arg8O is unknown (G. Maine and B. Tye, personalgenes had been placed on chromosome XIII by showing that communication). supll3 maps 20.9 cM distal to lys7 (Ono,both GAL80 and SUPS hybridize to the OFAGE-separated personal communication), whereas iIv2 is 38.2 cM distal toband corresponding to this chromosome (Carle and Olson, in this marker (148; Hawthorne, personal communication).press). SUP79 (68) and SUF22 (55) also tnap in this region, pet-ts2858 maps 23.2 cM distal to ilv2. This petite mutationdistal to rad52. spoT2 maps very close to the centromere also shows loose linkage to rnal which serves to join the(191), whereas tsmOll, secS9, and tsmO800, respectively, SUP8-rnal group meiotically to the rest of the chromosomenmap 1.9, 3.8, and 11.5 cM from the centromere on the right and to orient this group, with rnal most proximal (116).

TAB3LE 16. New tetrad analysis data for chromosome XVI'Segregation Ascus type (no.) Map distance

Interval (no.) ReferenceFD SD PD NPD T x' (CM) SE

spoT16-gal4 100 0 5 2.4 1.1 M. Tsuboi, personal communicationpep4-gal4 50 3 118 40.1 3.6 F. Park and E. Jones, personal comnmunicationcdc60-gal4 37 6 98 50.0 6.9 Hanic-Joyce, in presscdc60-pep4 36 0 3 3.9 2.3 Hanic-Joyce, in presstsrmOll5-radl 83 0 27 12.5 2.2 199tsmO115-cenl6 91 81 28.1 2.9 199tsmO115-mak6 10 0 13 28.4 5.3 199tsmO115-aro7 27 8 66 67.5 17.0 199nibl-radl 37 0 0 0 74nibl-cenl6 12 5 16.1 6.8 74SUF21-cenl6 159 3 0.9 0.5 55SUF21-aro7 46 1 96 35.7 2.8 55glnl-cenl6 45 59 36.0 4.5 A. Mitchell, personal communicationglnl-mak6 28 1 24 29.0 7.4 A. Mitchell, personal communicationglnl-mak3 85 0 36 15.1 2.2 A. Mitchell, personal communicationglnl-aro7 155 0 72 16.0 1.6 A. Mitchell, personal communicationtsmO120-cenl6 27 28 31.0 5.4 10; F. Boutelet and F. Hilger, personal communicationtsmO120-mak3 47 0 8 7.3 2.4 10; F. Boutelet and F. Hilger, personal communicationtsmO120-aro7 45 0 10 9.1 2.7 10; F. Boutelet and F. Hilger, personal communicationspoT20-aro7 22 0 1 2.3 2.5 191spoT20-cenl6 61 109 43.5 4.4 191Tyl-aro7 i9 4 52 54.7 11.6 92


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TABLE 17. New tetrad analysis data for chromosome XVII"Segrega- Ascus type Map

Interval tion (no.) (no.) distance ReferenceFD SD PD NPD T x'(CM) SE

KRBI-cenl7 268 13 2.3 0.6 202KRBI-cenl7 20 1 2.4 2.4 199

Total 288 14 2.3 0.6


SUP78 (68) and spoT7 (191) map close to rnal. ade4 (Schildand Mortimer, in press) and prcl (E. Jones, M. Aynardi, andM. Kolodny, personal communication) have been shown tomap distal to SUP8, with ade4 probably the most distal.adh3 maps 16 cM from lys7 (M. Ciriacy, personal commu-nication) and adh2 is 8 kb (ca. 3 cM) removed from ade4(27). cdc6l has been mapped 31.8 cM from ade4 (Hanic-Joyce, in press). Finally, Celenza and Carlson (personalcommunication) have placed SUC4 on chromosome XIII buthave not mapped it to a specific location on this chromo-some.

Chromosome XIVIn our earlier review (132), chromosome XIV was identi-

fied by the centromere-linked marker pet8, and rna2 and lys9were located distal to pet8 pn the right arm. Also, near thecentromere were $UFIO and spol. Another group of linkedgenes that inclpded mak26, ski4, hex2, petx, and ski3 hadbeen located on this chromosome only by trisomic analysis.The markers met4 and met2 had been eliminated from allidentified chromosomes, including XIV, by aneuploid anal-yses and hence assigned to chromosome XVII (130). Severalgenes were subseqqently shown to be linked to markers onboth of these "chromospmes." It has since been found thatthe original aneuploio analysis that established the indepen-dence of these chromosomes was faulty (90); the strain thatwas disomic for met4 in fact contained two copies of the leftarm of chromosome XIV and one copy of the right arm. Thispartial disome Was crossed to lys9, a right arm marker, and2:2 segregation for Lys+:Lys- was obtained, whereasdisomic ratios for met4 were observed. This indicated incor-rectly that met4 was not on the same chromosome as lys9.The spoll mapping method was subsequently used to showthat all of these markers were on the same chromosome (90).This is another example of the pitfalls of aneuploid analysisas a mapping method. It is our view that assignment of agene to a chromosome by this method must be consideredprovisional until confirmed by tetrad analysis or by mitoticcrossing-over procedures.To the combined chromosome XIV have been added

SUF6 distal to met2 (56), sec2 distal to pet2 (Fields andSchekman, personal communication), and foll and radSOproximal to pet2 (93; Game, personal communication). Inaddition, suf74 is 42 cM proximal to pet2 (35). The genes

pmsl (M. S. Williamson, J. C. Game, and S. Fogel, Genet-ics, in press), RASSC2 (88), top2 (K. Voelkel, S. DiNardo,and R. Sternglanz, manuscript in preparation), and Ieu4

(L. L. Chang, P. R. Gatzek, and G. B. Kohlhaw, Gene, inpress) all map near met4. These genes are of great interest:pmsl leads to a considerable increase in the frequency ofpostmeiotic segregation, RASSC2 is a close analog of a humanoncogene, and top2 codes for or controls the synthesis oftopoisomerase 2. Ieu4 is one of two genes coding for a-

isopropyl malate synthase (Chang et al., in press). Two Tyl

TABLE 18. Glossary of gene symbolsSymbol Definition

act.... Actinade ... . Adenine requiringadh.... Defective in alcohol utilizationadr ... . Alcohol nonutilizeralg.... Asparagine-linked glycosylation deficientAMY .... Antimycin resistanceanp ... . Sensitive to ANP and osmotic sensitiveant ... . Antibiotic resistanceard ... . Defective in cell cycle arrest at startarg ... . Arginine requiringaro ... . Aromatic amino acid requiringasp ... . Aspartic acid requiringate.... Arginyl-tRNA-protein transferase deficientAXE.... Axenomycin resistancebar ... . a cells lack barrier effect on a factorBOR.... Borrelidin resistancecan ... . Canavanine resistancecar ... . Arginine catabolism defectivecdc ... . Cell division cycle blocked at 36°Ccen ... . Centromerechl.... Chromosome losscho.... Choline requiringcly.... Cell lysis at 36°Ccpa ... . Arginine requiring in presence of excess uracilcry ..... . Cryptopleurine resistanceCUP/cup .... Copper resistancecyc ... . Cytochrome c deficiencycyh.... Cycloheximide resistancecyr ... . Adenylate cyclase deficientcys ... . Cysteine requiringdal.... Allantoin degradation deficientdbl ... . Alcian blue dye binding deficientdsm.... Premeiotic DNA synthesis deficientdur ... . Urea degradation deficienteam.... Phenotypic revertants of chol mutantserg ... . Defective in ergosterol biosynthesis; many also

nystatin resistanteth ... . Ethionine resistancefas.... Fatty acid synthetase deficientfdp .....,.. Unable to grow on glucose, fructose, sucrose, or

mannoseflk.... Resistance to catabolite repressionFLO.... Flocculationfol.... Folinic acid requiringfro.... Frothinggal.... Galactose nonutilizergcd ... . Depressed for general control of amino acid syn-

thesisgcn ... . Non-derepressible general control of amino acid

synthesisglc.... Glycogen storageglk.... Glucokinase deficient (unable to use glucose)gln.... Unable to derepress glutamine synthetasehem ... . Heme synthesis deficienthip ... . Histidine-specific permeasehis.... Histidine requiringHML .... Mating type cassetteHMR .... Mating type cassetteHO.... Homothallic switchinghol ... . Histidinol uptake proficienthom ... . Homoserine requiringhxk.... Hexokinase deficientils .. .. Isoleucyl-transfer RNA synthetase deficient; no

growth at 36°Cilv.... Isoleucine-plus-valine requiringino ... . Inositol deficientkar ... . Nuclear fusion defectivekex ... . Unable to express killer phenotype


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TABLE 18-ContinuedSymbol Definition

KRB... Suppression of some mak mutationslap... Leucine aminopeptidase deficientlet... Lethalleu... Leucine requiringIts .. . Low-temperature sensitiveIys... Lysine requiringmak .. . Maintenance of killer deficientMAL ... Maltose fermentation positivemar... Partial expression of mating type cassettesMAT .,.... Mating type locusmcm... Minichromosome maintenance deficientmes... Methionyl-transfer RNA synthetase defective; no

growth at 36°Cmet.. Methionine requiringmfa... a-mating factorMGL .. a-Methylglucoside fermentermin.. Inhibited by methioninemnn.. Mannan synthesis defectivemut.. Elevated spontaneous mutation ratenam. . Nuclear suppressor of mitochondrial mutationsNHS.. Hydrogen sulfide production inhibitornib. . Nibbled colony phenotype due to 2,um DNAnul. . Nonmaterole.. Oleic acid requiringoli.. Oligomycin resistanceosm.. Sensitive to low osmotic pressurepdx.. Pyridoxine requiringpep.. Proteinase deficientpet. . Petite; unable to grow on nonfermentable carbon

sourcespgi.. Phosphoglucose isomerase deficientpgk.. 3-Phosphoglycerate kinase deficientpha.. Phenylalanine requiringpho.. Phosphatase deficientphr. . Photoreactivation repair deficientPHS .. Hydrogen sulfide production deficientpms.. Increased postmeiotic segregationppr. . Defective in pyrimidine biosynthetic pathway reg-

ulationprb. . Proteinase B deficientprc. . Proteinase C deficientprt.. Protein synthesis defective at 36°Cpur. . Purine excretionput. . Proline nonutilizerpyk.. Pyruvate kinase deficientrad. . Radiation (ultraviolet or ionizing) sensitiveRASSc.. Homologous to RAS proto-oncogeneRDN.. Ribosomal RNA structural genesrev.. Nonrevertiblerme.. Meiosis independent of mating type heterozygos-

ityrna. . Unable to grow at 36°C; block in RNA synthesisROC.. Roccal resistancesam.. S-Adenosylmethionine deficientsec. . Secretion deficientser.. Serine requiringsir.. Defective in regulation of silent mating type infor-

mationski.. SuperkillerSMR/smr.... Sulfometuron methyl resistancesnf.... Deficient in derepression of many glucose-

repressible genessot.... Suppression of deoxythymidine monophosphatc

uptakespd ... . Sporulation not repressed on rich mediaspe ... . Spermidine resistancespo ... . Sporulation deficientspt.... Suppressor of Ty-mediated expression of his4ssn ... . Suppressor of snfl

Continued

TABLE 18-ContinuedSymbol Definition

sst.... Supersensitive to a factorste.... SterileSUC.... Sucrose fermenterSUF/suf.... Suppression of frameshift mutationsuh ... . Suppression of his2-1SUP/sup .... Suppression of nonsense mutationSUS ... . Suppression of serlswi ... . Homothallic switching deficienttcm.... Trichodermin resistanceTEF ... . Translational elongation factorthi.... Thiamine requiringthr.... Threonine requiringtil .. .. Thiaisoleucine resistancetmp.... Thymidine monophosphate requiringtop ... . Topoisomerase deficienttra.... Triazylalanine resistanttmn.... Proline transfer RNA genetrp. .... Tryptophan requiringtsm.... Lethal, temperature sensitivetub ... . Tubulin; MBC resistancetup ... . Deoxythymidine monophosphate uptake positivetyr.... Tyrosine requiringumr.... Non-ultraviolet revertibleura ... . Uracil requiring

sequences map between met4 and the centromere (92). Thegene spol has been moved from the right to the left arm butis still very close to the centromere (di Domenico andEsposito, personal communication). Three genes have beenmapped near lys9. These are SUP28 (Ono, personal commu-nication), pet494 (138), and holl (P. Farabaugh, personalcommunication). The order of these three genes relative toeach other, lys9, or proximal markers is unknown.

Chromosome XVChromosome XV is one of the larger chromosomes of

yeasts, with a physical size comparable to that of chromo-some VII and with only chromosomes IV and XII larger(Carle and Olson, in press). Its meiotic length is about 375cM and 39 genes have been mapped along it. Since our lastreview, the frameshift suppressors SUFI and SUFJ7 havebeen added to the left arm (55, 56). In addition, adhl(Ciriacy, personal communication) and glc4 (J. Pringle,personal communication) have been assigned to the arg8-argi interval on this chromosome arm. spoTIl is near thecentromere on the left arm (191). The gene makJ, which hadbeen mapped on the right arm near the centromere, has beenshown to be an allele of topi (186), the gene that controls thesynthesis of topoisomerase 1 of yeasts. Interestingly, eithertopoisomerase 1 or topoisomerase 2 is sufficient for' most ofthe functions of the yeast cell cycle; topoisomerase 2 alone isnecessary for separation of replicated chromosomes in mi-tosis (41). mak8 has been shown to be allelic to temi (204),which had previously been mapped proximal to petl7 (62).Very near to tsm8740, a cell cycle mutant, is RASSCl ahuman oncogene analog (82, 182). It appears, however, thatthese genes are not alleles. A few map units distal to tsm8740is spoT15 (191). Distal to his3 are ste4 (166) and stel3 (D.Barnes and J. Thorner, personal communication), and cdc3lmaps further out on the chromosome distal to met7 (Schildand Mortimer, in press). The recessive frameshift suppres-sor sufl3 is near cpal (35) and distal to these genes is cdc64(Hanic-Joyce, in press). Both sufl3 and cdc64 show linkageto prtl. This latter gene had earlier been located on chromo-

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TABLE 19. List of mapped genes

Gene Map Reference(s)position

4R See gcn211R See gcn35R See gcn46L 491R 129, 15815R 1297R 82, 129, 18513R Schild and Mortimer, in press7L 129, 140, 2017R 69, 165; D. J. Plotkin, Ph.D. thesis, Uni-

versity of Chicago, Chicago, Ill., 19784R 2, 130; G. E. Jones, Ph.D. thesis, Uni-

versity of California, Berkeley, 197015R 37, 1297R 8215L M. Ciriacy, personal communication13R 27; M. Ciriacy, personal communication13R M. Ciriacy, personal communication4R J. Wood and C. L. Denis, personal com-

munication2R 337R 67L 1102L 110, 199SL 1237L 298R M. Whiteway and J. Szostak, personal

communication15L 30, 7110L 728R 69, 129, 1984 200SR 53, 107, 129, 141; D. P. Morrison, Ph.D.

thesis, University of Alberta, Edmon-ton, 1978

15L 71SR 12913R 7213L 72; Schild and Mortimer, in press4R 72SR 794R 129, 130, 2037L 129, 140, 165, 20116R 70, 103, 145, 2004R 83; Jones, Ph.D. thesis12R 43, 78, 129, 1507L 1607L S. Sora, personal communication9L G. Sprague and I. Herskowitz, personal

communicationSR 1417L 141SL 130, 164, 20012R 724L 66, 91, 130, 2036L 38, 13013L 38, 13010L F. Hilger, personal communication; Ka-

wasaki, Ph.D. thesis4L 130, 1451OR 97, 1304L 66, 1303R 34, 1301OR 97, 1308R 38, 2006R 38, 1301R 13011L 66, 2031L Kawasaki, Ph.D. thesis

TABLE 19-Continued


cdc21cdc24cdc26cdc28cdc29cdc31cdc35cdc36cdc37cdc39cdc4O

cdc43

cdc6Ocdc61cdc62cdc63cdc64cdc66

chilcholcly2cly3cly7cly8cpalcrylCUPJcup2

cup3

cupS

cup14

cyclcyc2cyc3cyc7cyc8cyc9cyhlcyh2cyh3cyh4cyhlOcyrlcysI

cys3dalldal2dal3

dal4dal80dal8idblidsml

durldur2dur3

dur4

15R1L6R2R9L15R1OR4L4R3R4R

7L

16L13R7R15R15R15R

16LSR2L6R11L7R15R3R8R7L

12R

SL

4R

1OR15R1LSL2R3R2L7L7L15R2R1OR1L

1L9R9R9R

9R11R9RIlL7R

2R2R8L

8L

Continued

58, 20085Kawasaki, Ph.D. thesis32, 66, 20066Schild and Mortimer, in pressSee tsmO185166166166Y. Kassir, M. Kupiec, A. Shalom, andG. Simchen, personal communication

A. Adams and J. Pringle, personal com-munication

Hanic-Joyce, in pressHanic-Joyce, in pressHanic-Joyce, in pressHanic-Joyce, in pressHanic-Joyce, in pressP. Hanic-Joyce and D. R. Carruthers,

personal communication1084, 107130, 199130130130; Plotkin, Ph.D. thesis7261, 125, 17069, 129J. W. Welch and S. Fogel, personalcommunication

J. W. Welch and S. Fogel, personalcommunication



97, 130158158164158158129129, 140, 165; Plotkin, Ph.D. thesis129129167121158, 203; S. Halos, Ph.D. thesis, Univer-

sity of California, Berkeley, 197614731, 9831, 98T. G. Cooper and H. S. Yoo, personalcommunication

31, 981921925, 54D. Fast, Ph.D. thesis, University of Chi-

cago, Chicago, Ill., 19783232T. G. Cooper and M. Mojumdar, per-

sonal communicationT. G. Cooper and M. Mojumdar, per-

sonal communication


aasiaas2aas3actiadelade2ade3ade4ade5,7ade6

ade8

ade9ADEJSadhiadh2adh3adrl

alglalg7AMY]AMY2anpiantiardl

arglarg3arg4arg5,6

arg8arg9arg8Oarg8iarg82arg84arolaro2aro7aspiasp5ateiAXEIbarl

BORiBOR2canicar2cdc2cdc4cdcScdc6

cdc7cdc8cdc9cdciOcdclIcdci2cdci4cdci5cdci6cdcl9

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TABLE 19-Continued


253122117, 118; Schild and Mortimer, in press16, 3619417575, 158, 177See FLOIJ. Game and J. Little, personal commu-

nicationJ. Game and J. Little, and B. Rockmill,

personal communication184, 185184, 1858, 43, 107, 12943, 129, 15043, 80J. Haber, personal communication448, 43, 107, 1298, 43, 107, 129Schild and Mortimer, in press; Carle and

Olson, in press12073; D. Yep, Ph.D. thesis, Cornell Uni-

versity, Ithaca, N.Y., 1973737373J. Pringle, personal communicationJ. Pringle, personal communicationP. Oeller and J. Pringle, personal com-

municationJ. Pringle, personal communication115A. P. Mitchell, personal communication127193Tanaka and Fink, Abstr. Mol. Biol.

Yeast Meet., 198353, 107, 129, 141; E. Savage, Ph.D. the-

sis, University of Alberta, Edmonton,1979

40, 70, 129, 13037, 12934, 69, 185, 196107, 129, 14469, 130, 144129; Plotkin, Ph.D. thesisSee his36565Kawasaki, Ph.D. thesisP. Farabaugh, personal communication129, 13052, 129, 141Schild and Mortimer, in press109; Lobo, Ph.D. thesis11512476, 107, 12914870, 97, 129, 1301484247, 90201188, 201

Continued

TABLE 19-Continued

Gene Map Reference(s)position Rfrnes

KRBIlap]lap3lap4letlletlMlet3let5let6leulleu2leu4ltsllts3lts4ItslOlyslIys2lys4

lysSlys7lys9lysllmaklmak3mak4makSmak6mak7mak8mak9maklOmaklImakl2makl3makl4maklSmakl,6makl,7makl8makl,9mak20mak2lmak22mak24mak26mak27MALIMAL2MAL3MAL4marlmar2(ste8)MATmcml

meslmetlmet2met3met4metSmet6met7met8metlO

metl3

17RllL14LllL1R13R10L10L6L7L3L14L7L7L4R4R9R2R4R

7L13R14R9L15R16R2R2R16R8L15RllLSLllL12L9R3R11R1L.10L8R8R8L4R12L7L14L13R7R3R2R11R4L12R3R13R

7R11R14L1OR14L1ORSR15R2R6R

7L

19919090, 19019013013913913913969, 109, 129, 16534, 69, 107, 129Chang et al., in press16716716716731, 69, 98, 107, 14432, 69, 129, 200; Plotkin, Ph.D. thesisR. Contopoulou, personal communica-

tion129, 140, 165, 20136, 66, 129, 130129, 130, 201107, 129, 144200145, 200200200145, 200197, 198, 200200203164, 20020320319820320320319819819819820319819819820323, 82, 129, 18512, 69, 109114; Kawasaki, Ph.D. thesis70, 129, 130, 20391A. Klar, personal communication61, 69, 125, 129, 170G. Maine and B. Tye, personal commu-

nication13269, 129, 13090, 13097, 129, 14590, 130Schild and Mortimer, in press117, 118; Schild and Mortimer, in pressLowenstein, Ph.D. thesis32, 70, 12940, 107, 109, 129; Kawasaki, Ph.D.

thesis129, 140, 165, 168, 201


DUR80eamlerg6eth2(sam2)fasifdplflkiFLO]FLO4foil

fo/2

frolfro2gallgal2gal3gal4galSgal7gall,Ogal80

GAL83gcdl(tra3)a

gcn2(aasl)gcn3(aas2)gcn4(aas3)gllgIc3gIc4

glc6glklglnlgIn3hem,10hipi

his]

his2his3his4A,B,ChisShis6his7his8HMLHMRHOhollhom2hom3hom6hxklhxk2ilsiilvlilv2iIv3ilvSinolkarlkexlkex2

2R10L4R5RllL2R3R1R1R14L

7R

7R7R2R12R4R16LF62R2R13L

5R15R

4R11RSR2R5L15L

2R3L16RSR7L7R

SR

6R15R3L9L9L2R15R3L3R4L14R4RSR1OR6R7L2L5R13R1OR12R10L14L7L14L

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Reference(s)

66, 77, 129, 203119C. Shari and J. Kuran, personal com-

munication114; Kawasaki, Ph.D. thesis1641

60604618074130155, 16515916816869206; E. Jones, personal communicationF. Park and E. Jones, personal commu-

nicationE. Jones and M. Kolodny, personalcommunication

G. Fabian and E. Jones, personal com-munication

E. Jones, personal communication69, 12990, 129, 13038, 130, 198, 20038, 129, 130, 20143, 129, 14232, 70, 129130, 20370, 130, 158, 200130, 2001381161162011149590, 129, 13018764, 171, 183, 189187869187187161112179Williamson et al., in press104207E. Jones, M. Aynardi, and M. Kolodny,

personal communication13090, 130130211113, 158, 174See rad24108, 153, 154132172; Morrison, Ph.D. thesis

Continued

TABLE 19-Continued

Gene Map Reference(s)

rad4 5R 172; Morrison, Ph.D. thesisradS 12R 99rad6 7L 59rad7 1OR 97rad9 4R Dowling, Ph.D. thesisradi8 3R 130, 158rad23 5L 123rad24 SR F. Eckardt and J. Game, personal com-

municationradSO 14L 93; J. Game, personal communicationradSl SR Morrison, Ph.D. thesisradS2a 13L 154rad54 7L J. Game, personal communication;

Dowling, Ph.D. thesisradSS 4R 130rad56 16R 59rad57 4R 59, 145RASSCI 15R 88, 182RASSC2 14L 88; L. Robinson and K. Tatchell, per-

sonal communicationRDNI 12R 150rev2 12R See radSrev5 1OR C. W. Lawrence, personal communica-

tionrev7 9L C. W. Lawrence, personal communica-

tionrmel 7R 156rnal 13R 130, 198, 203rna2 14R 38, 130, 201rna3 4R 130rnaS 2R 130; Kawasaki, Ph.D. thesisrna6 2R See tsm7269rnall 4L 130, 187, 203ROCI 12R 129sam2 (eth2) SR 117, 118; Schild and Mortimer, in presssecl 4R C. Fields and R. Schekman, personal

communicationsec2 14L C. Fields and R. Schekman, personal

communicationsec3 SR C. Fields and R. Schekman, personal

communicationsec4 6L C. Fields and R. Schekman, personal

communicationsecS 4R C. Fields and R. Schekman, personal

communicationsec7 4R C. Fields and R. Schekman, personal

communicationseci8 2R C. Fields and R. Schekman, personal

communicationsec55 3R C. Fields and R. Schekman, personal

communicationsecS9 13R C. Fields and R. Schekman, personal

communicationserl 15R 129; J. R. Johnston, Ph.D. thesis, Uni-

versity of California, Berkeley, 1962;Lowenstein, Ph.D. thesis

ser2sirlsir2sir3sir4skilski3

ski4SMRIsmr2smr3snflsotl

7R 8211R J. M. Ivy, personal communication4L See mar]12R See mar2 and ste84R J. M. Ivy, personal communication7L 18814L 18814L 18813R 507L 5015R 504R 2216L 153


TABLE 19-Continued

Gene

meti4met2Omfca2

MGL2minimnni

mnn2

mnn4mutimut2nam2NHSJnibinul3olelolilosmlosm2pdx2pep3pep4

pep7

pepi2

pepi6petlpet2pet3pet8pet9petl Ipeti4peti 7peti8pet494pet-tsl402pet-ts2858petxpgilpgklpha2pho2pho3,5pho4pho8pho8OPHO82pho85phrlphr2PHSJpmslpprlprblprcl

prtlprt2prt3purSput2pykl

radlrad2rad3

Mapposition

11R11R7L

2R5LSC2R11LF1lF1l12RSR16L4R7L7L1OR16RF612R16L

4R

15L

12R8R14L8R14R2L2R4R15R3R14R5R13R14L2R3R14L4L2R6R4R15R6R16L15R15R4R14L12R5L13R

15R14LSL4R8R1L5R16L7RSR

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TABLE 19-Continued

Gene position Reference(s)

I. Dawes, personal communication30, 7190; B. DiDomenico and R. E. Esposito,

personal communicationR. E. Esposito and C. Waddell, personalcommunication

S. Klapholz and R. E. Esposito, person-al communication

89S. Klapholz and R. E. Esposito, person-

al communicationM. Townsend, B. DiDomenico, S. Kia-

pholz, and R. E. Esposito, personalcommunication

M. Tsuboi, personal communicationM. Tsuboi, personal communication191M. Tsuboi, personal communication191M. Tsuboi, personal communicationM. Tsuboi, personal communicationM. Tusboi, personal communication1911912052052121R. K. Chan, personal communicationR. K. Chan, personal communication166166166R. K. Chan, personal communicationSee sir4D. Barnes and J. Thorner, personalcommunication

23, 82, 129, 157, 185107, 129, 144114; Kawasaki, Ph.D. thesis20; Kawasaki, Ph.D. thesis56345623, 56375638383838563535355555555555555555555555, 5613017

Continued

TABLE 19-Continued

Gene position Reference(s)

SUP2SUP3SUP4

SUPSSUP6SUP7SUP8SUPIISUP15,16SUP17SUP19,20SUP22SUP25SUP26SUP27SUP28SUP29SUP30SUP33sup35sup36SUP37SUP38sup45SUP46sup47SUPSOSUPS1SUP52SUP53

SUP54

SUP56SUPS7SUP58SUP61SUP71SUP72SUP73SUP74SUP75SUP76SUP77SUP78SUP79SUP80SUP85SUP86SUP87SUP88sup)l)sup,112sup] 13SUSIswil

temlTEF2thilthrithr4till

tmpltop2

4R15L1OR

13L6R10L13R6R16R9LSR9L11R12R4R14R10c10c11L4R4R12R7L2R2R2RF610c10c3L

7L

1R6R11L3R5R2R10L10L11L7R7R13R13L4R5R12R2R4R8R7R13R5L16L

15R2RF68R3R7L

15R14L

13070, 7197, 130; R. Gilmore, Ph.D. thesis, Uni-

versity of California, Berkeley, 196645; Schild and Mortimer, in press40, 70, 13070; Gilmore, Ph.D. thesis130, 19840, 70, 13070, 103, 145144130, 14514470, 130145145B. Ono, personal communication145130B. Ono, personal communication129143B. Ono, personal communicationB. Ono, personal communication70, 171143, 146143129, 130130102152; C. Reed and S. Liebman, personalcommunication

101, 152; C. Reed and S. Liebman, per-sonal communication

10110110112, 130130686868D. Hawthorne, personal communication686868686868686868B. Ono, personal communicationB. Ono, personal communicationB. Ono, personal communication12963; J. Haber and L. Rowe, personalcommunication

62, 20416245, 129, 13069, 12995, 97, 100, 129, 181, 196J. I, Stiles and F. Sherman, personal

communication13, 58, 200K. Voelkel, S. DiNardo, and R. Stern-

glanz, personal communication


spdlspe2spol

spo7

15L15L14L

1L

spoil

spol2spol3

spol4

spoT)spoT2spoT4spo27spoT8spoTl)spoT15spoTl6spol20spoT23spt2spt3ssn2ssn6(cyc8)sstlsst2ste4steSste7ste8(mar2)ste9stel3

SUCISUC2SUC3SUC5SUFISUF2SUF3SUF4SUF5SUF6SUF7SUF8SUF9SUFIOsuflIsufl2sufl3sufl4SUFISSUF16SUF17SUF18SUF19SUF20SUF21SUF22SUF23SUF24SUF25suh2SUP-IA

8L

8R8R

11R

13C7L4L13R2R15L15R16L16R11RSR4R4R2R9L12R15R4R4L12R4R15R

7R9L2R4L15L3R4R7R15R14L13L8R6L14L15R4R15R14L7R3R15L6RSL6R16R13L1OR4R4L12R11R

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TABLE 19-Continued

Gene Map Reference(s)position Rfrnes

15R 73; D. Yep, Ph.D. thesis, Cornell Uni-versity, Ithaca, N.Y., 1975 (see gedi)

1R C. Cummins and M. Culbertson,personal communication

4R 43, 59, 67, 129, 130, 145, 203SR 53, 107, 129, 14111L 5, 16, 36, 129, 2034R 83, 130; Jones, Ph.D. thesis7L 69, 129, 140, 141, 155, 165, 168; Plotkin,

Ph.D. thesis3R G. Sprague and J. Herskowitz, personal

communication3R G. Fink, personal communication; J.

McCusker and J. Haber, personalcommunication

10101010200101064, 114, 130101010

10F. Hilger, personal communication13010721072

SR6L4R13R16L7L16R2R9L8R1OR

8R4L7L13R16R4R2R

15R 726L J. Thomas, S. C. Falco, and D. Bot-

stein, personal communication3R 100, 19615L 20015R 92R 32, 64, 129, 158; Plotkin, Ph.D. thesis3R 10011L 5, 16, 34, 129, 20310L 130; Kawasaki, Ph.D. thesisSL 1, 4, 129, 130, 16412R 130, 150

a The reference to the mapping of radS2 and tra3 (gcdl) was incorrect in ourprevious review (132). radS2 was mapped by M. Resnick (154) and tra3 was

mapped by D. Yep (Ph.D. thesis, 1975).

(130). The cell division cycle mutation cdc63 has also beenmapped in this region and has been shown to be allelic withprtl (Hanic-Joyce, in press). At 13.6 cM from prtl is phrl(161), although the order of these two genes on the chromo-some is still unknown. It has also been reported that phr2 islinked to phrl at a distance of 18 cM (112).

Chromosome XVIgal4 is the most distal marker on the left arm of chromo-

some XVI; close to this gene is the sporulation-defectivegene spoT16 (191). About 40 cM proximal to gal4 is pep4 (E.Jones, personal communication) and 3.9 cM from pep4 is thecell cycle mutant cdc6O (Hanic-Joyce, in press). This groupof genes had been located on the left arm of chromosomeXVI because gaI4 shows mitotic linkage to radl (J. Mc-Cusker and J. Haber, personal communication); however,

TABLE 20. Genetic and physical sizes of yeast chromosomesRecombinant DNA x 10-16 DNA DNABand Chromosome length (cM) ga (kb)b (kb)C

1 I 98 14 198 2602 VI 137 16 225 2903 III 137 22 311 3704 IX 198 35 495 4605a VIII 183 45 635 5805b V 233 50 709 5806 XI 242 54 763 7007 X 200 59 8358 XIV 283 61 8649 II 244 67 94810 XIII 229 70 99110b XVI 176 74 1,047lla XV 361 81 1,146llb VII 391 91 1,28512 IV 485 111 1,571(13) XIId 294 155 2,194a DNA content/meiotic bivalent (4c), assuming 250 x 10-16 g per haploid

genome (94).b DNA content/chromosome, assuming 14,000 kb per haploid genome (96).c As determined from size standards, using OFAGE (Carle and Olson, in

press).d See discussion of chromosome XII.

no meiotic linkage exists between gal4 and radl. Neitherpep4 nor cdc60, the two most proximal genes of this group,has been tested for linkage against other more proximal leftarm markers. To this proximal group of genes have beenadded tsmO15 (199) and nib] (74). The frameshift suppres-sor SUF21 is approximately 1 cM from the centromere onthe right arm (55). Distal to mak6 are glnl (A. Mitchell,personal communication) and tsmOJ20 (Boutelet and Hilger,personal communication). Finally, a Tyl sequence mapsapproximately 55 cM distal to aro7 (92), which places it nearradS6.

Chromosome XVIIChromosome XVII is identified only by the centromere-

linked marker KRBI (killer replication bypass) (199, 202). Asshown in these studies, this marker arises spontaneously inmak7 segregants from crosses that carry the killer plasmid.Such segregants normally are killer minus (K-) becausemak7 strains cannot maintain the killer plasmid. KRBI is achromosomal mutation that in effect suppresses the mak7mutation. It is tightly centromere linked (<2 cM) and is notlinked to markers near the centromeres of chromosomes I toXVI. KRBI strains have normal killer plasmids and normalkiller double-stranded RNA. Also, KRBI does not appear tobe a translational-type suppressor. However, the behavior ofKRBI is not typical of mutations in nuclear genes. It arises ata high frequency in cells or segregants of cells that carry thekiller plasmid. Also, it disappears when mak7 KRBI cells aregrown at 37°C or are exposed to cycloheximide, treatmentsknown to cure cells of the killer plasmid. Finally, a crossbetween two mak7 KRBI strains, which should yield 4:0segregations for killer-nonkiller, more frequently yields 2:2segregations, and experiments indicate that one of the twoKRBI mutations has been lost.

If chromosome XVII, as defined by KRBI were "nor-mal," i.e., typical of chromosomes I to XVI, one wouldexpect to find centromere-linked genes on this chromosomeat a frequency similar to that seen for the other chromo-somes. For chromosomes I and XVI there are 144 markerswithin 25 cM of the centromere (centromere linked) for anaverage of 9 centromere-linked markers per centromere(range, 4 to 17). Assuming random association of markers

tra3a

trni

trpltrp2trp3trp4trp5

tsml

tsm5

tsmOO39tsmOO70tsmOO80tsmOlI ItsmOlJStsmOII9tsmO120tsm134tsmO139tsmOISItsmO185

(cdc35)tsmO186tsmO225tsm437tsmO800tsm4572tsm5162tsm7269

(rna6)tsm8740tub2

tupltup4tup7tyrlumr7uralura2ura3ura4

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a, 1400

1200-

1000 xv i

600_VIII/IX

400 _

/11

600 _ v

/

JI0 100 200 300 400 500 600 700

Chromosome Map Length (cM)FIG. 2. Plot of chromosome sizes versus genetic map lengths.

The points with arrows to the tight represent minimum map lengths;these six chromosomes have regions whose lengths have not beendetermined by tetrad analysis.

with centromeres aid equal probabilities of association/cen-tromere, the probability of finding a centromere with onlyone centromere marker is very small (<10-3).Two recent studies which permit identification of individ-

ual chromosomes have found evidence for only 16 chromo-somes. OFAGE reveals only 16 chromosomal bands (Carleand Olson, in press) and DAPI (4',6-diamidino-2-phenylin-dole) staining of meiotic bivalents also reveals only 16 suchstructures (94). One must conclude that the chromosomeidentified by KRBJ is anomolous. Although it segregates atthe first meiotic division, it fails to do so regularly. It appearsand disappears in a manner completely atypical of nucleargenes. Also, were chromosome XVII typical of the otherchromosomes, several other genes by now would have beenfound to be linked to KRBI. This presents a dilemma.A possible explanation, which has features similar to one

proposed by Wickner and Leibowitz (202), is that KRBI islocated on a small linear or circular piece of DNA that alsocontains sequences that behave as a centromere. This pieceofDNA could be a reverse transcript of all or a portion of thekiller double-stranded RNA. This could account for the highfrequency of appearance of KRBI in cells containing suchRNA. Its small size would account for its instability incertain circumstances just as cen plasmids are somewhatunstable (28). The OFAGE and DAPI staining experiments,discussed above, may have been carried out on killer-minusstrains which lack this small chromosome.

DISCUSSION

The genetic map of S. cerevisiae presented in this articledescribes the location of 568 genes distributed over 16metacentric chromosomes plus a single gene, KRBI, locatedon a 17th chromosome. Since our last major review (132),251 genes have been added to the genetic map. In addition,several linkages which had been established only by mitoticor aneuploid analyses have been confirmed by tetrad analy-sis. Only chromosomes VIII, IX, X, XI, XII, and XVIremain with regions not confirmed by tetrad analysis. As-suming a minimum of 100 cM for these regions, the totalminimum length of the yeast map is now 4,500 cM, which is100 cM less than our estimate made 5 years ago. This and thefact that the total number of mapped genes has increased by79% argue that the current lengths of the yeast chromosomesare close to their actual lengths.As discussed in detail above, three chromosomes have

undergone major revisions since our last review. These arechromosomes VII, XIII, and XIV. On the right arm ofchromosome VII, a large group of genes has been reversedin order relative to the centromere on the basis of meioticlinkage between SUF4 and cly8. This places SUCJIMALJ asthe most distal markers on this chromosome, where beforethis pair had been assigned as the proximal-most geneslocated distal to cly8. Chromosome XIII has had severalmajor changes. The genes eth2 and met6 have been moved tochromosome V and the group of genes on the right arm,which had been located on this chromosome by mitoticanalyses, has been shown to be meiotically linked to genesnearer the centromere. A group of genes previously identi-fied as fragment V has been shown to be located on the leftarm of this chromosome, and several other genes identifiedwith this chromosome or fragment have been assigned tospecific sites. Altogether, 17 genes have been added to thischromosome and its length has been extended to 229 cM.The most significant change on the genetic map since our lastreview is the joining of chromosomes XIV and XVII. Thesewere incorrectly identified as separate chromosomes on thebasis of aneuploid analyses, but subsequent studies showedthat they were in fact parts of a single chromosome. Thiscombined chromosome is metacentric and is one of themedium-sized chromosomes of yeasts. Another significantchange is the substitution of KRBI as the centromere markerto define chromosome XVII. This is still problematicalbecause of the anomalous genetic behaviour of this gene (seecomments on chromosome XVII).By and large, functionally related genes in yeasts are

distributed somewhat randomly over the yeast genome. Forexample, the five genes involved in tryptophan biosynthesisare located on five different chromosomes. Nevertheless,there are some notable exceptions. On chromosome III,three enzymatic activities in histidine biosynthesis are clus-tered at the his4 locus. In fact, this gene codes for a singlepolypeptide that has all three activities. The three genesgal7, gallO, and gall are tightly linked on chromosome II,yet they do not represent an operon in the bacterial sense.All three genes are independently under the control of afourth gene, gal4, located on chromosome XVI. This gene,in turn, is regulated by GAL80, located on chromosome XII.On chromosome V is located the arg5-arg6 cluster. Thesegenes appear to be under the control of the tightly linkedgene arg8O in an operon-like assembly, although it has beenargued that this group also is not like a bacterial operon. Acomplete review of genetic regulation in yeasts with appro-priate references has been presented by Jones and Fink (81).

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Two remarkable developments have added new dimen-sions to our throughts about the yeast genome; both of thesehave appeared within the last year or two. A new procedurefor gel electrophoresis allows separation of individual chro-mosomes (163; Carle and Olson, in press). In addition, aprocedure involving DAPI staining has permitted identifica-tion of individual meiotic bivalents (94). By use of the novelgel electrophoresis procedure, Carle and Olson (in press)have electrophoretically resolved 16 chromosomal bands(albeit by using four strains) and have identified each of thesewith particular chromosomes by probing Southern blots withclones of genes known to be located on particular chromo-somes. No evidence for a 17th chromosome could be found.The DAPI staining procedure clearly resolved 16 meioticbivalents and photographs of these were scanned to deter-mine their relative sizes. These sizes were calibrated byassuming the DNA content of a haploid as 250 x 10-16 g(1.56 x 101 daltons). With this value, sizes ranged from 14x 10-6 to 155 x 10-16 g per bivalent. A more accurate valuefor the DNA content of a haploid genome is 0.9 x 1010daltons (96), which leads to a chromosomal size range of 198to 2,194 kb. By assuming the same order, small to large, asdetermined by the electrophoretic and DAPI staining meth-ods, it was possible to assign a physical size to each of thechromosomes (Table 20). This chromosomal size (convertedto kilobases) is plotted against genetic map length incentimorgans (Fig. 2). A remarkable linear relation betweenphysical size and recombination length is observed, and theslope of this line is 3.6 kb/cM. Lauer et al. (96) estimated thesize of the largest yeast chromosome to be 1.5 x 109 to 2.2x 109 daltons (2,200 to 3,330 kb). These values are inreasonable agreement with the estimated size of chromo-some XII (Table 20).

ACKNOWLEDGMENTS

We thank Rebecca Contopoulou for many useful comments on themanuscript and Diana Morris for help in its preparation. Most of allwe acknowledge and thank the many investigators who providedunpublished mapping data and copies of articles containing data. Westrongly encourage researchers who are citing map positions ormapping data to reference the original source, wherever possible,rather than this review.

This work was supported by a grant from the Office of Health andEnvironmental Research of the U.S. Department of Energy undercontract DE-AC03-76SFQ0098 and by Public Health Service grantGM30990 from the National Institutes of Health.

ADDENDUM IN PROOF

The centromere-linked gene rad57 has recently been shownto map on the left arm of chromosome IV proximal to rnal linstead ofon the right arm (R. Contopoulou and R. Mortimer,unpublished data) and cdc29 has been mapped distal ratherthan proximal to his6 on chromosome IX (G. Basile and R. K.Mortimer, unpublished data). The previous edition of thegenetic map (136) mistakenly identified the ssn6 marker onchromosome II as ssnl; ssnl has not been mapped yet (M.Carlson, personal communication).

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