Standard name Systematic name Name description Description
PDS5 YMR076C Cohesion maintenance factor Cohesion maintenance factor; involved in sister chromatid condensation and cohesion; colocalizes with cohesin on chromosomes; performs its cohesin maintenance function in pre-anaphase cells by protecting the integrity of the cohesion complex; regulates homolog pairing and facilitates synaptonemal complex formation, axis formation, interhomolog recombination and synapsis during meiosis; relocalizes to the cytosol in response to hypoxia; scaffolding protein contains 26 HEAT repeats
Visualize the interactome of PDS5
Links to external databases:
SGD   |   KEGG   |  

Cell cycle phase and timing of transcription peak

Cell cycle phase Time (min)
G1/S 55

CYCLoPs localization & abundance

WT1 WT2 WT3
Nucleolus 0.203 0.190 0.210
Nucleus 0.794 0.796 0.790
Mito NaN NaN 0.356

GFP localization & abundance

GFP localization GFP abundance
nucleus 7720

GO term count

Metabolism Cell cycle Cell division Signal transduction DNA replication
PDS5 2 7 0 0 0

List of interactors

Use the search utility to find the interactor you are looking for. By clicking the column headers the table will be sorted on that column. Use shift+click to sort on multiple columns. Default sorting is on number of experiments, number of unique methods and the number of publications, and alphabetical on standard name, in that order.
Standard name Systematic name Name description Type #Experiments #Methods #Publications Evidence
MCD1 YDL003W Essential alpha-kleisin subunit of the cohesin complex physical 13 6 9 Affinity Capture-Western, FRET, Affinity Capture-Western, Affinity Capture-MS, Affinity Capture-Western, Affinity Capture-Western, Reconstituted Complex, Co-crystal Structure, Reconstituted Complex, Affinity Capture-Western, Reconstituted Complex, Co-purification, Affinity Capture-MS
ECO1 YFR027W Acetyltransferase required for sister chromatid cohesion genetic 8 3 5 Synthetic Lethality, Dosage Rescue, Dosage Rescue, Synthetic Rescue, Synthetic Rescue, Synthetic Rescue, Synthetic Rescue, Synthetic Rescue
RAD61 YDR014W Subunit of a complex that inhibits sister chromatid cohesion physical 5 2 3 Affinity Capture-Western, Affinity Capture-Western, Affinity Capture-Western, Two-hybrid, Affinity Capture-Western
FKH1 YIL131C Forkhead family transcription factor regulation 4 2 4 chromatin immunoprecipitation-chip evidence, chromatin immunoprecipitation- exonuclease evidence, chromatin immunoprecipitation-chip evidence, chromatin immunoprecipitation- exonuclease evidence
ELG1 YOR144C Subunit of an alternative replication factor C complex genetic 4 3 3 Synthetic Rescue, Synthetic Rescue, Phenotypic Suppression, Negative Genetic
RAD61 YDR014W Subunit of a complex that inhibits sister chromatid cohesion genetic 3 3 3 Phenotypic Suppression, Negative Genetic, Phenotypic Enhancement
IRR1 YIL026C Subunit of the cohesin complex physical 3 2 3 Affinity Capture-MS, Affinity Capture-Western, Affinity Capture-Western
SMC3 YJL074C Subunit of the multiprotein cohesin complex physical 3 2 3 Affinity Capture-Western, Affinity Capture-MS, Affinity Capture-Western
SMC1 YFL008W Subunit of the multiprotein cohesin complex physical 3 3 2 Affinity Capture-Western, FRET, Affinity Capture-MS
POL30 YBR088C Proliferating cell nuclear antigen (PCNA) genetic 3 3 2 Dosage Rescue, Phenotypic Suppression, Negative Genetic
GIM3 YNL153C Subunit of the heterohexameric cochaperone prefoldin complex genetic 3 1 2 Negative Genetic, Negative Genetic, Negative Genetic
MMS21 YEL019C Highly conserved SUMO E3 ligase subunit of SMC5-SMC6 complex genetic 3 3 1 Synthetic Growth Defect, Phenotypic Enhancement, Phenotypic Suppression
ECO1 YFR027W Acetyltransferase required for sister chromatid cohesion physical 2 2 2 Biochemical Activity, Affinity Capture-Western
HOS1 YPR068C Class I histone deacetylase (HDAC) family member genetic 2 2 2 Phenotypic Suppression, Positive Genetic
TOP2 YNL088W Topoisomerase II genetic 2 2 2 Dosage Rescue, Synthetic Growth Defect
SGS1 YMR190C RecQ family nucleolar DNA helicase genetic 2 2 2 Synthetic Lethality, Positive Genetic
SLX5 YDL013W Subunit of the Slx5-Slx8 SUMO-targeted Ub ligase (STUbL) complex genetic 2 2 2 Synthetic Rescue, Negative Genetic
CHD1 YER164W Chromatin remodeler that regulates various aspects of transcription genetic 2 1 2 Negative Genetic, Negative Genetic
CHL4 YDR254W Outer kinetochore protein required for chromosome stability genetic 2 1 2 Negative Genetic, Negative Genetic
KAP95 YLR347C Karyopherin beta genetic 2 1 2 Negative Genetic, Negative Genetic
DHH1 YDL160C Cytoplasmic DEAD-box helicase and mRNA decapping activator physical 2 1 2 Affinity Capture-RNA, Affinity Capture-RNA
MAD2 YJL030W Component of the spindle-assembly checkpoint complex genetic 2 1 2 Negative Genetic, Negative Genetic
ARP6 YLR085C Actin-related protein that binds nucleosomes genetic 2 1 2 Negative Genetic, Negative Genetic
PAN1 YIR006C Part of actin cytoskeleton-regulatory complex Pan1p-Sla1p-End3p genetic 2 1 2 Positive Genetic, Positive Genetic
CIN2 YPL241C GTPase-activating protein (GAP) for Cin4p genetic 2 1 2 Negative Genetic, Negative Genetic
SMC1 YFL008W Subunit of the multiprotein cohesin complex genetic 2 1 2 Negative Genetic, Negative Genetic
SPC105 YGL093W Subunit of a kinetochore-microtubule binding complex genetic 2 1 2 Negative Genetic, Negative Genetic
RIO1 YOR119C Serine kinase involved in cell cycle regulation and rDNA integrity genetic 2 1 2 Negative Genetic, Negative Genetic
SRS2 YJL092W DNA helicase and DNA-dependent ATPase genetic 2 1 2 Negative Genetic, Negative Genetic
MOB2 YFL034C-B Activator of Cbk1p kinase genetic 2 2 1 Positive Genetic, Negative Genetic
NFI1 YOR156C SUMO E3 ligase genetic 2 2 1 Synthetic Rescue, Phenotypic Suppression
SMT3 YDR510W Ubiquitin-like protein of the SUMO family genetic 2 2 1 Synthetic Rescue, Phenotypic Suppression
SLX8 YER116C Subunit of Slx5-Slx8 SUMO-targeted ubiquitin ligase (STUbL) complex genetic 2 2 1 Synthetic Rescue, Phenotypic Suppression
CTF19 YPL018W Outer kinetochore protein, needed for accurate chromosome segregation physical 2 1 1 Two-hybrid, Two-hybrid
CTF19 YPL018W Outer kinetochore protein, needed for accurate chromosome segregation genetic 2 1 1 Negative Genetic, Negative Genetic
CTF3 YLR381W Outer kinetochore protein that forms a complex with Mcm16p and Mcm22p genetic 2 1 1 Negative Genetic, Negative Genetic
MCM22 YJR135C Outer kinetochore protein and component of the Ctf3 subcomplex genetic 2 1 1 Negative Genetic, Negative Genetic
IML3 YBR107C Outer kinetochore protein and component of the Ctf19 complex genetic 2 1 1 Negative Genetic, Negative Genetic
KIN3 YAR018C Nonessential serine/threonine protein kinase physical 2 1 1 Two-hybrid, Two-hybrid
GIN4 YDR507C Protein kinase involved in bud growth and assembly of the septin ring physical 2 1 1 Two-hybrid, Two-hybrid
CTF4 YPR135W Chromatin-associated protein genetic 2 1 1 Negative Genetic, Negative Genetic
CLB2 YPR119W B-type cyclin involved in cell cycle progression genetic 2 1 1 Negative Genetic, Negative Genetic
MCM16 YPR046W Component of the Ctf19 complex and the COMA subcomplex genetic 2 1 1 Negative Genetic, Negative Genetic
BUB1 YGR188C Protein kinase involved in the cell cycle checkpoint into anaphase genetic 2 1 1 Negative Genetic, Negative Genetic
CTF8 YHR191C Subunit of a complex with Ctf18p genetic 2 1 1 Negative Genetic, Negative Genetic
CIN1 YOR349W Tubulin folding factor D involved in beta-tubulin (Tub2p) folding genetic 2 1 1 Negative Genetic, Negative Genetic
SSA1 YAL005C ATPase involved in protein folding and NLS-directed nuclear transport physical 2 1 1 Affinity Capture-MS, Affinity Capture-MS
SSB1 YDL229W Cytoplasmic ATPase that is a ribosome-associated molecular chaperone physical 2 1 1 Affinity Capture-MS, Affinity Capture-MS
STP22 YCL008C Component of the ESCRT-I complex physical 2 1 1 Two-hybrid, Two-hybrid
URM1 YIL008W Ubiquitin-like protein involved in thiolation of cytoplasmic tRNAs genetic 2 1 1 Positive Genetic, Positive Genetic
RPN4 YDL020C Transcription factor that stimulates expression of proteasome genes genetic 2 1 1 Negative Genetic, Negative Genetic
SGF73 YGL066W Subunit of DUBm module of SAGA and SLIK genetic 2 1 1 Negative Genetic, Negative Genetic
SMI1 YGR229C Protein involved in the regulation of cell wall synthesis genetic 2 1 1 Negative Genetic, Negative Genetic
HCM1 YCR065W Forkhead transcription factor genetic 2 1 1 Negative Genetic, Negative Genetic
DCC1 YCL016C Subunit of a complex with Ctf8p and Ctf18p genetic 2 1 1 Negative Genetic, Negative Genetic
ICE2 YIL090W Integral ER membrane protein with type-III transmembrane domains genetic 2 1 1 Positive Genetic, Positive Genetic
FKH2 YNL068C Forkhead family transcription factor regulation 1 1 1 chromatin immunoprecipitation-chip evidence
REB1 YBR049C RNA polymerase I enhancer binding protein regulation 1 1 1 chromatin immunoprecipitation-chip evidence
CKA2 YOR061W Alpha' catalytic subunit of casein kinase 2 (CK2) genetic 1 1 1 Positive Genetic
CKB2 YOR039W Beta' regulatory subunit of casein kinase 2 (CK2) genetic 1 1 1 Positive Genetic
NIP7 YPL211W Nucleolar protein required for 60S ribosome subunit biogenesis genetic 1 1 1 Synthetic Lethality
NOG1 YPL093W Putative GTPase genetic 1 1 1 Negative Genetic
NOP7 YGR103W Component of several different pre-ribosomal particles physical 1 1 1 Affinity Capture-MS
ADH1 YOL086C Alcohol dehydrogenase genetic 1 1 1 Negative Genetic
KAP123 YER110C Karyopherin beta genetic 1 1 1 Negative Genetic
BRE1 YDL074C E3 ubiquitin ligase genetic 1 1 1 Positive Genetic
MCM21 YDR318W Component of the kinetochore sub-complex COMA genetic 1 1 1 Negative Genetic
LSM1 YJL124C Lsm (Like Sm) protein genetic 1 1 1 Negative Genetic
CDC48 YDL126C AAA ATPase with protein-unfoldase activity genetic 1 1 1 Negative Genetic
LRP1 YHR081W Nuclear exosome-associated nucleic acid binding protein genetic 1 1 1 Positive Genetic
MED7 YOL135C Subunit of the RNA polymerase II mediator complex genetic 1 1 1 Negative Genetic
DST1 YGL043W General transcription elongation factor TFIIS genetic 1 1 1 Negative Genetic
NAP1 YKR048C Histone chaperone and assembly chaperone for Rps6a/b genetic 1 1 1 Negative Genetic
HCR1 YLR192C eIF3j component of translation initiation factor 3 (eIF3) genetic 1 1 1 Negative Genetic
ACT1 YFL039C Actin, structural protein in cytoskeleton genetic 1 1 1 Positive Genetic
CCR4 YAL021C Component of the CCR4-NOT transcriptional complex physical 1 1 1 Affinity Capture-RNA
ACC1 YNR016C Acetyl-CoA carboxylase, biotin containing enzyme genetic 1 1 1 Negative Genetic
APC1 YNL172W Largest subunit of the Anaphase-Promoting Complex/Cyclosome genetic 1 1 1 Negative Genetic
ARC15 YIL062C Subunit of the ARP2/3 complex genetic 1 1 1 Positive Genetic
ASF1 YJL115W Nucleosome assembly factor genetic 1 1 1 Negative Genetic
BEM2 YER155C Rho GTPase activating protein (RhoGAP) genetic 1 1 1 Positive Genetic
CDC28 YBR160W Cyclin-dependent kinase (CDK) catalytic subunit genetic 1 1 1 Negative Genetic
CDC3 YLR314C Component of the septin ring that is required for cytokinesis physical 1 1 1 Two-hybrid
BUD31 YCR063W Component of the SF3b subcomplex of the U2 snRNP genetic 1 1 1 Positive Genetic
DCP1 YOL149W Subunit of the Dcp1p-Dcp2p decapping enzyme complex genetic 1 1 1 Negative Genetic
DOA1 YKL213C WD-repeat protein involved in ubiquitin-mediated protein degradation genetic 1 1 1 Positive Genetic
CDC16 YKL022C Subunit of the anaphase-promoting complex/cyclosome (APC/C) genetic 1 1 1 Negative Genetic
CDC23 YHR166C Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C) genetic 1 1 1 Negative Genetic
EAF3 YPR023C Component of the Rpd3S histone deacetylase complex genetic 1 1 1 Negative Genetic
FKS1 YLR342W Catalytic subunit of 1,3-beta-D-glucan synthase genetic 1 1 1 Negative Genetic
CCT4 YDL143W Subunit of the cytosolic chaperonin Cct ring complex genetic 1 1 1 Negative Genetic
HHF1 YBR009C Histone H4 genetic 1 1 1 Negative Genetic
ISW1 YBR245C ATPase subunit of imitation-switch (ISWI) class chromatin remodelers physical 1 1 1 Affinity Capture-RNA
HSL1 YKL101W Nim1p-related protein kinase genetic 1 1 1 Negative Genetic
END3 YNL084C EH domain-containing protein involved in endocytosis physical 1 1 1 Two-hybrid
MCD1 YDL003W Essential alpha-kleisin subunit of the cohesin complex genetic 1 1 1 Negative Genetic
MCM5 YLR274W Component of the Mcm2-7 hexameric helicase complex genetic 1 1 1 Negative Genetic
MED1 YPR070W Subunit of the RNA polymerase II mediator complex genetic 1 1 1 Negative Genetic
HSC82 YMR186W Cytoplasmic chaperone of the Hsp90 family genetic 1 1 1 Synthetic Growth Defect
HRR25 YPL204W Conserved casein kinase, monopolin subunit genetic 1 1 1 Negative Genetic
MAM33 YIL070C Acidic protein of the mitochondrial matrix genetic 1 1 1 Positive Genetic
ITC1 YGL133W Subunit of ATP-dependent Isw2p-Itc1p chromatin remodeling complex genetic 1 1 1 Negative Genetic
NUP60 YAR002W FG-nucleoporin component of central core of the nuclear pore complex genetic 1 1 1 Positive Genetic
ORC2 YBR060C Subunit of the origin recognition complex (ORC) genetic 1 1 1 Negative Genetic
PAN2 YGL094C Catalytic subunit of the Pan2p-Pan3p poly(A)-ribonuclease complex genetic 1 1 1 Negative Genetic
HSP82 YPL240C Hsp90 chaperone genetic 1 1 1 Synthetic Growth Defect
DAM1 YGR113W Essential subunit of the Dam1 complex (aka DASH complex) genetic 1 1 1 Negative Genetic
MET18 YIL128W Component of cytosolic iron-sulfur protein assembly (CIA) machinery genetic 1 1 1 Positive Genetic
DEP1 YAL013W Component of the Rpd3L histone deacetylase complex genetic 1 1 1 Negative Genetic
CTI6 YPL181W Component of the Rpd3L histone deacetylase complex genetic 1 1 1 Negative Genetic
NUM1 YDR150W Protein required for nuclear migration physical 1 1 1 Two-hybrid
CSE1 YGL238W Nuclear envelope protein that acts as a recycling factor genetic 1 1 1 Negative Genetic
NEM1 YHR004C Probable catalytic subunit of Nem1p-Spo7p phosphatase holoenzyme genetic 1 1 1 Positive Genetic
HEM13 YDR044W Coproporphyrinogen III oxidase genetic 1 1 1 Negative Genetic
BCY1 YIL033C Regulatory subunit of the cyclic AMP-dependent protein kinase (PKA) genetic 1 1 1 Positive Genetic
BRO1 YPL084W Cytoplasmic class E vacuolar protein sorting (VPS) factor genetic 1 1 1 Positive Genetic
HYP2 YEL034W Translation elongation factor eIF-5A genetic 1 1 1 Negative Genetic
CDC5 YMR001C Polo-like kinase required for mitotic exit physical 1 1 1 Affinity Capture-MS
ARO4 YBR249C 3-deoxy-D-arabino-heptulosonate-7-phosphate (DAHP) synthase physical 1 1 1 Affinity Capture-MS
BUD13 YGL174W Subunit of the RES complex genetic 1 1 1 Negative Genetic
GET2 YER083C Subunit of the GET complex genetic 1 1 1 Positive Genetic
CDC9 YDL164C DNA ligase I found in nucleus and mitochondria genetic 1 1 1 Negative Genetic
CYC8 YBR112C General transcriptional co-repressor genetic 1 1 1 Positive Genetic
DDC1 YPL194W DNA damage checkpoint protein genetic 1 1 1 Negative Genetic
PBR1 YNL181W Putative oxidoreductase genetic 1 1 1 Negative Genetic
FUM1 YPL262W Fumarase genetic 1 1 1 Positive Genetic
CDC20 YGL116W Activator of anaphase-promoting complex/cyclosome (APC/C) genetic 1 1 1 Negative Genetic
ELM1 YKL048C Serine/threonine protein kinase genetic 1 1 1 Negative Genetic
HSL7 YBR133C Protein arginine N-methyltransferase genetic 1 1 1 Negative Genetic
CDC37 YDR168W Essential Hsp90p co-chaperone genetic 1 1 1 Negative Genetic
AIM46 YHR199C Protein of unknown function genetic 1 1 1 Negative Genetic
MCX1 YBR227C Non-proteolytic ATPase of the AAA family genetic 1 1 1 Negative Genetic
KEL2 YGR238C Protein that negatively regulates mitotic exit physical 1 1 1 Two-hybrid
CAC2 YML102W Subunit of chromatin assembly factor I (CAF-1), with Rlf2p and Msi1p genetic 1 1 1 Negative Genetic
ISU1 YPL135W Conserved protein of the mitochondrial matrix genetic 1 1 1 Negative Genetic
NUS1 YDL193W Forms dehydrodolichyl diphosphate synthase complex with RER2 or SRT1 genetic 1 1 1 Negative Genetic
HAL5 YJL165C Snf1p-related nutrient-responsive protein kinase physical 1 1 1 Affinity Capture-MS
PAC2 YER007W Microtubule effector required for tubulin heterodimer formation genetic 1 1 1 Negative Genetic
DEF1 YKL054C RNAPII degradation factor genetic 1 1 1 Negative Genetic
BUB3 YOR026W Kinetochore checkpoint WD40 repeat protein genetic 1 1 1 Negative Genetic
IST3 YIR005W Component of the U2 snRNP genetic 1 1 1 Positive Genetic
CDC7 YDL017W S/T kinase and catalytic subunit of DDK (Dbf4-dependent kinase) genetic 1 1 1 Negative Genetic
MMS22 YLR320W Subunit of E3 ubiquitin ligase complex involved in replication repair genetic 1 1 1 Negative Genetic
DAD1 YDR016C Essential subunit of the Dam1 complex (aka DASH complex) genetic 1 1 1 Negative Genetic
DID4 YKL002W Class E Vps protein of the ESCRT-III complex genetic 1 1 1 Positive Genetic
ATG1 YGL180W Protein serine/threonine kinase genetic 1 1 1 Negative Genetic
CSM1 YCR086W Nucleolar protein that mediates homolog segregation during meiosis I genetic 1 1 1 Negative Genetic
MCM10 YIL150C Essential chromatin-associated protein genetic 1 1 1 Positive Genetic
BIM1 YER016W Microtubule plus end-tracking protein genetic 1 1 1 Negative Genetic
GNA1 YFL017C Glucosamine-6-phosphate acetyltransferase genetic 1 1 1 Negative Genetic
IPK1 YDR315C Inositol 1,3,4,5,6-pentakisphosphate 2-kinase genetic 1 1 1 Negative Genetic
HSP10 YOR020C Mitochondrial matrix co-chaperonin genetic 1 1 1 Negative Genetic
NUP57 YGR119C FG-nucleoporin component of central core of the nuclear pore complex genetic 1 1 1 Negative Genetic
JSN1 YJR091C Member of the Puf family of RNA-binding proteins genetic 1 1 1 Negative Genetic
ASK1 YKL052C Essential subunit of the Dam1 complex (aka DASH complex) genetic 1 1 1 Negative Genetic
JLP1 YLL057C Fe(II)-dependent sulfonate/alpha-ketoglutarate dioxygenase genetic 1 1 1 Positive Genetic
MDY2 YOL111C Protein involved in inserting tail-anchored proteins into ER membranes genetic 1 1 1 Negative Genetic
HUA1 YGR268C Cytoplasmic protein containing a zinc finger domain genetic 1 1 1 Positive Genetic
APC5 YOR249C Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C) genetic 1 1 1 Negative Genetic
APC9 YLR102C Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C) genetic 1 1 1 Negative Genetic
CDC27 YBL084C Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C) genetic 1 1 1 Negative Genetic
IPL1 YPL209C Aurora kinase of chromosomal passenger complex genetic 1 1 1 Negative Genetic
HTL1 YCR020W-B Component of the RSC chromatin remodeling complex genetic 1 1 1 Positive Genetic
CHL1 YPL008W Probable DNA helicase genetic 1 1 1 Negative Genetic
DAL81 YIR023W Positive regulator of genes in multiple nitrogen degradation pathways genetic 1 1 1 Positive Genetic
CBF1 YJR060W Basic helix-loop-helix (bHLH) protein genetic 1 1 1 Negative Genetic
MRS1 YIR021W Splicing protein genetic 1 1 1 Positive Genetic
BFR1 YOR198C Component of mRNP complexes associated with polyribosomes physical 1 1 1 Two-hybrid
BRN1 YBL097W Subunit of the condensin complex genetic 1 1 1 Negative Genetic
GIM4 YEL003W Subunit of the heterohexameric cochaperone prefoldin complex genetic 1 1 1 Negative Genetic
MAD1 YGL086W Coiled-coil protein involved in spindle-assembly checkpoint genetic 1 1 1 Negative Genetic
MCM3 YEL032W Protein involved in DNA replication genetic 1 1 1 Negative Genetic
MPS1 YDL028C Dual-specificity kinase genetic 1 1 1 Negative Genetic
CBF2 YGR140W Essential kinetochore protein genetic 1 1 1 Negative Genetic
ORC3 YLL004W Subunit of the origin recognition complex (ORC) genetic 1 1 1 Negative Genetic
KTI11 YBL071W-A Zn-ribbon protein that co-purifies with Dph1 and Dph2 genetic 1 1 1 Negative Genetic
HSF1 YGL073W Trimeric heat shock transcription factor genetic 1 1 1 Negative Genetic
MPT5 YGL178W mRNA-binding protein of the PUF family physical 1 1 1 Affinity Capture-RNA
FCP1 YMR277W Carboxy-terminal domain (CTD) phosphatase genetic 1 1 1 Positive Genetic
ESC2 YDR363W Sumo-like domain protein genetic 1 1 1 Positive Genetic
GAL1 YBR020W Galactokinase genetic 1 1 1 Negative Genetic
GPR1 YDL035C Plasma membrane G protein coupled receptor (GPCR) genetic 1 1 1 Negative Genetic
NNF2 YGR089W Protein that exhibits physical and genetic interactions with Rpb8p physical 1 1 1 Two-hybrid
IVY1 YDR229W Phospholipid-binding protein that interacts with both Ypt7p and Vps33p physical 1 1 1 Two-hybrid
PBP1 YGR178C Component of glucose deprivation induced stress granules physical 1 1 1 Affinity Capture-MS
PBP1 YGR178C Component of glucose deprivation induced stress granules genetic 1 1 1 Negative Genetic
MCA1 YOR197W Ca2+-dependent cysteine protease genetic 1 1 1 Negative Genetic
CHS5 YLR330W Component of the exomer complex physical 1 1 1 Affinity Capture-MS
PAM16 YJL104W Subunit of the import motor (PAM complex) genetic 1 1 1 Positive Genetic
BET4 YJL031C Alpha subunit of Type II geranylgeranyltransferase genetic 1 1 1 Negative Genetic
AIM44 YPL158C Regulator of Cdc42p and Rho1p genetic 1 1 1 Negative Genetic
PRE1 YER012W Beta 4 subunit of the 20S proteasome physical 1 1 1 Affinity Capture-MS
XRN1 YGL173C Evolutionarily-conserved 5'-3' exonuclease physical 1 1 1 Affinity Capture-MS
XRN1 YGL173C Evolutionarily-conserved 5'-3' exonuclease genetic 1 1 1 Positive Genetic
SMC3 YJL074C Subunit of the multiprotein cohesin complex genetic 1 1 1 Negative Genetic
REC8 YPR007C Meiosis-specific component of the sister chromatid cohesion complex physical 1 1 1 Affinity Capture-MS
REC8 YPR007C Meiosis-specific component of the sister chromatid cohesion complex genetic 1 1 1 Phenotypic Enhancement
TSC11 YER093C Subunit of TORC2 (Tor2p-Lst8p-Avo1-Avo2-Tsc11p-Bit61p) physical 1 1 1 Two-hybrid
YGR130C YGR130C Component of the eisosome with unknown function physical 1 1 1 Two-hybrid
YAP1802 YGR241C Protein of the AP180 family, involved in clathrin cage assembly physical 1 1 1 Two-hybrid
SMY1 YKL079W Kinesin-like myosin passenger-protein physical 1 1 1 Two-hybrid
YAP5 YIR018W Basic leucine zipper (bZIP) iron-sensing transcription factor physical 1 1 1 Two-hybrid
SNF7 YLR025W One of four subunits of the ESCRT-III complex physical 1 1 1 Two-hybrid
VPS64 YDR200C Protein required for cytoplasm to vacuole targeting of proteins physical 1 1 1 Two-hybrid
SPO21 YOL091W Component of the meiotic outer plaque of the spindle pole body physical 1 1 1 Two-hybrid
SCC2 YDR180W Subunit of cohesin loading factor (Scc2p-Scc4p) physical 1 1 1 Affinity Capture-MS
SCC2 YDR180W Subunit of cohesin loading factor (Scc2p-Scc4p) genetic 1 1 1 Negative Genetic
PUF4 YGL014W Member of the PUF protein family physical 1 1 1 Affinity Capture-RNA
RET1 YOR207C Second-largest subunit of RNA polymerase III physical 1 1 1 Affinity Capture-MS
SLT2 YHR030C Serine/threonine MAP kinase physical 1 1 1 PCA
SIS1 YNL007C Type II HSP40 co-chaperone that interacts with the HSP70 protein Ssa1p physical 1 1 1 Affinity Capture-MS
ULP2 YIL031W Peptidase that deconjugates Smt3/SUMO-1 peptides from proteins genetic 1 1 1 Dosage Rescue
SEC15 YGL233W Essential 113 kDa subunit of the exocyst complex genetic 1 1 1 Synthetic Lethality
TAF1 YGR274C TFIID subunit, involved in RNA pol II transcription initiation genetic 1 1 1 Synthetic Lethality
TEL2 YGR099W Subunit of the ASTRA complex, involved in chromatin remodeling genetic 1 1 1 Positive Genetic
RET2 YFR051C Delta subunit of the coatomer complex (COPI) genetic 1 1 1 Positive Genetic
RPB3 YIL021W RNA polymerase II third largest subunit B44 genetic 1 1 1 Positive Genetic
SSL2 YIL143C Component of RNA polymerase transcription factor TFIIH holoenzyme genetic 1 1 1 Positive Genetic
SEC11 YIR022W 18kDa catalytic subunit of the Signal Peptidase Complex (SPC) genetic 1 1 1 Positive Genetic
RPB11 YOL005C RNA polymerase II subunit B12.5 genetic 1 1 1 Positive Genetic
SEC23 YPR181C GTPase-activating protein, stimulates the GTPase activity of Sar1p genetic 1 1 1 Positive Genetic
VID24 YBR105C GID Complex regulatory subunit genetic 1 1 1 Positive Genetic
YSA1 YBR111C Nudix hydrolase family member with ADP-ribose pyrophosphatase activity genetic 1 1 1 Positive Genetic
SNF5 YBR289W Subunit of the SWI/SNF chromatin remodeling complex genetic 1 1 1 Positive Genetic
RHB1 YCR027C Putative Rheb-related GTPase genetic 1 1 1 Positive Genetic
SED4 YCR067C Integral ER membrane protein that stimulates Sar1p GTPase activity genetic 1 1 1 Positive Genetic
RPL2A YFR031C-A Ribosomal 60S subunit protein L2A genetic 1 1 1 Positive Genetic
SLX9 YGR081C Protein required for pre-rRNA processing genetic 1 1 1 Positive Genetic
RNR3 YIL066C Minor isoform of large subunit of ribonucleotide-diphosphate reductase genetic 1 1 1 Positive Genetic
SMT1 YJL147C Translational repressor of the mitochondrial ATP6/8 mRNA genetic 1 1 1 Positive Genetic
UFD4 YKL010C Ubiquitin-protein ligase (E3) genetic 1 1 1 Positive Genetic
SPE1 YKL184W Ornithine decarboxylase genetic 1 1 1 Positive Genetic
TRM2 YKR056W tRNA methyltransferase genetic 1 1 1 Positive Genetic
RPS0B YLR048W Protein component of the small (40S) ribosomal subunit genetic 1 1 1 Positive Genetic
PSY3 YLR376C Component of Shu complex (aka PCSS complex) genetic 1 1 1 Positive Genetic
TMA23 YMR269W Nucleolar protein implicated in ribosome biogenesis genetic 1 1 1 Positive Genetic
YMR315W YMR315W Protein with NADP(H) oxidoreductase activity genetic 1 1 1 Positive Genetic
RPS10A YOR293W Protein component of the small (40S) ribosomal subunit genetic 1 1 1 Positive Genetic
PHO11 YAR071W One of three repressible acid phosphatases genetic 1 1 1 Positive Genetic
SSD1 YDR293C Translational repressor genetic 1 1 1 Positive Genetic
SWA2 YDR320C Auxilin-like protein involved in vesicular transport genetic 1 1 1 Positive Genetic
VPS29 YHR012W Subunit of the membrane-associated retromer complex genetic 1 1 1 Positive Genetic
SDS3 YIL084C Component of the Rpd3L histone deacetylase complex genetic 1 1 1 Positive Genetic
RPL40A YIL148W Ubiquitin-ribosomal 60S subunit protein L40A fusion protein genetic 1 1 1 Positive Genetic
STU1 YBL034C Microtubule plus-end-tracking non-motor protein genetic 1 1 1 Negative Genetic
RER2 YBR002C Forms the dehydrodolichyl diphosphate synthase (DDS) complex with NUS1 genetic 1 1 1 Negative Genetic
TSC3 YBR058C-A Protein that stimulates the activity of serine palmitoyltransferase genetic 1 1 1 Negative Genetic
SLI15 YBR156C Subunit of the conserved chromosomal passenger complex (CPC) genetic 1 1 1 Negative Genetic
TSC13 YDL015C Enoyl reductase genetic 1 1 1 Negative Genetic
RPN6 YDL097C Essential, non-ATPase regulatory subunit of the 26S proteasome lid genetic 1 1 1 Negative Genetic
QRI1 YDL103C UDP-N-acetylglucosamine pyrophosphorylase genetic 1 1 1 Negative Genetic
SCM3 YDL139C Nonhistone component of centromeric chromatin genetic 1 1 1 Negative Genetic
RPN5 YDL147W Subunit of the CSN and 26S proteasome lid complexes genetic 1 1 1 Negative Genetic
PSF1 YDR013W Subunit of the GINS complex (Sld5p, Psf1p, Psf2p, Psf3p) genetic 1 1 1 Negative Genetic
RPS13 YDR064W Protein component of the small (40S) ribosomal subunit genetic 1 1 1 Negative Genetic
TAF12 YDR145W Subunit (61/68 kDa) of TFIID and SAGA complexes genetic 1 1 1 Negative Genetic
TAF10 YDR167W Subunit (145 kDa) of TFIID and SAGA complexes genetic 1 1 1 Negative Genetic
UBC1 YDR177W Ubiquitin-conjugating enzyme genetic 1 1 1 Negative Genetic
SPC19 YDR201W Essential subunit of the Dam1 complex (aka DASH complex) genetic 1 1 1 Negative Genetic
SRP101 YDR292C Signal recognition particle (SRP) receptor alpha subunit genetic 1 1 1 Negative Genetic
RSC3 YDR303C Component of the RSC chromatin remodeling complex genetic 1 1 1 Negative Genetic
YCG1 YDR325W Subunit of the condensin complex genetic 1 1 1 Negative Genetic
YDR341C YDR341C Arginyl-tRNA synthetase genetic 1 1 1 Negative Genetic
SEC20 YDR498C Membrane glycoprotein v-SNARE genetic 1 1 1 Negative Genetic
RBA50 YDR527W Protein involved in transcription genetic 1 1 1 Negative Genetic
PRP22 YER013W DEAH-box RNA-dependent ATPase/ATP-dependent RNA helicase genetic 1 1 1 Negative Genetic
UTP7 YER082C Nucleolar protein genetic 1 1 1 Negative Genetic
SCC4 YER147C Subunit of cohesin loading factor (Scc2p-Scc4p) genetic 1 1 1 Negative Genetic
TUB2 YFL037W Beta-tubulin genetic 1 1 1 Negative Genetic
RPN11 YFR004W Metalloprotease subunit of 19S regulatory particle genetic 1 1 1 Negative Genetic
TAF6 YGL112C Subunit (60 kDa) of TFIID and SAGA complexes genetic 1 1 1 Negative Genetic
RPS2 YGL123W Protein component of the small (40S) subunit genetic 1 1 1 Negative Genetic
SEC27 YGL137W Essential beta'-coat protein of the COPI coatomer genetic 1 1 1 Negative Genetic
UFD1 YGR048W Substrate-recruiting cofactor of the Cdc48p-Npl4p-Ufd1p segregase genetic 1 1 1 Negative Genetic
PHS1 YJL097W Essential 3-hydroxyacyl-CoA dehydratase of the ER membrane genetic 1 1 1 Negative Genetic
SPC42 YKL042W Central plaque component of spindle pole body (SPB) genetic 1 1 1 Negative Genetic
YJU2 YKL095W Essential protein required for pre-mRNA splicing genetic 1 1 1 Negative Genetic
SRP102 YKL154W Signal recognition particle (SRP) receptor beta subunit genetic 1 1 1 Negative Genetic
TFA2 YKR062W TFIIE small subunit genetic 1 1 1 Negative Genetic
SFI1 YLL003W Centrin (Cdc31p)-binding protein required for SPB duplication genetic 1 1 1 Negative Genetic
SDO1 YLR022C Guanine nucleotide exchange factor (GEF) for Ria1p genetic 1 1 1 Negative Genetic
STU2 YLR045C Microtubule polymerase of the XMAP215/Dis1 family genetic 1 1 1 Negative Genetic
TUB4 YLR212C Gamma-tubulin genetic 1 1 1 Negative Genetic
SRP1 YNL189W Karyopherin alpha homolog genetic 1 1 1 Negative Genetic
PRP2 YNR011C RNA-dependent DExD/H-box ATPase genetic 1 1 1 Negative Genetic
RFC4 YOL094C Subunit of heteropentameric Replication factor C (RF-C) genetic 1 1 1 Negative Genetic
TOA1 YOR194C TFIIA large subunit genetic 1 1 1 Negative Genetic
RPT4 YOR259C ATPase of the 19S regulatory particle of the 26S proteasome genetic 1 1 1 Negative Genetic
RPN8 YOR261C Essential non-ATPase regulatory subunit of the 26S proteasome genetic 1 1 1 Negative Genetic
TAF3 YPL011C TFIID subunit (47 kDa) genetic 1 1 1 Negative Genetic
TIM50 YPL063W Essential component of the TIM23 complex genetic 1 1 1 Negative Genetic
SUI3 YPL237W Beta subunit of the translation initiation factor eIF2 genetic 1 1 1 Negative Genetic
RPN7 YPR108W Essential non-ATPase regulatory subunit of the 26S proteasome genetic 1 1 1 Negative Genetic
VPS8 YAL002W Membrane-binding component of the CORVET complex genetic 1 1 1 Negative Genetic
SWC3 YAL011W Protein of unknown function genetic 1 1 1 Negative Genetic
SHE1 YBL031W Mitotic spindle protein genetic 1 1 1 Negative Genetic
PIN4 YBL051C Protein involved in G2/M phase progression and response to DNA damage genetic 1 1 1 Negative Genetic
SWC5 YBR231C Component of the SWR1 complex genetic 1 1 1 Negative Genetic
RRT2 YBR246W Methylesterase performing penultimate step of diphthamide biosynthesis genetic 1 1 1 Negative Genetic
RER1 YCL001W Protein involved in retention of membrane proteins genetic 1 1 1 Negative Genetic
STE50 YCL032W Adaptor protein for various signaling pathways genetic 1 1 1 Negative Genetic
YCR090C YCR090C Putative protein of unknown function genetic 1 1 1 Negative Genetic
RPP1A YDL081C Ribosomal stalk protein P1 alpha genetic 1 1 1 Negative Genetic
YGL036W YGL036W Putative protein of unknown function genetic 1 1 1 Negative Genetic
RAD54 YGL163C DNA-dependent ATPase that stimulates strand exchange genetic 1 1 1 Negative Genetic
PMR1 YGL167C High affinity Ca2+/Mn2+ P-type ATPase genetic 1 1 1 Negative Genetic
RPS25A YGR027C Protein component of the small (40S) ribosomal subunit genetic 1 1 1 Negative Genetic
TIM21 YGR033C Nonessential component of the TIM23 complex genetic 1 1 1 Negative Genetic
RPL11B YGR085C Ribosomal 60S subunit protein L11B genetic 1 1 1 Negative Genetic
PRE9 YGR135W Alpha 3 subunit of the 20S proteasome genetic 1 1 1 Negative Genetic
YTA7 YGR270W Chromatin-binding ATPase that regulates histone gene expression genetic 1 1 1 Negative Genetic
RRM3 YHR031C DNA helicase involved in rDNA replication and Ty1 transposition genetic 1 1 1 Negative Genetic
SRB2 YHR041C Subunit of the RNA polymerase II mediator complex genetic 1 1 1 Negative Genetic
UBA4 YHR111W E1-like protein that activates Urm1p before urmylation genetic 1 1 1 Negative Genetic
STB5 YHR178W Transcription factor genetic 1 1 1 Negative Genetic
YJL163C YJL163C Putative protein of unknown function genetic 1 1 1 Negative Genetic
YJL206C YJL206C Putative protein of unknown function genetic 1 1 1 Negative Genetic
TMA19 YKL056C Protein that associates with ribosomes genetic 1 1 1 Negative Genetic
RAD5 YLR032W DNA helicase/Ubiquitin ligase genetic 1 1 1 Negative Genetic
RIC1 YLR039C Protein involved in retrograde transport to the cis-Golgi network genetic 1 1 1 Negative Genetic
SIC1 YLR079W Cyclin-dependent kinase inhibitor (CKI) genetic 1 1 1 Negative Genetic
UPS1 YLR193C Phosphatidic acid transfer protein genetic 1 1 1 Negative Genetic
YLR415C YLR415C Putative protein of unknown function genetic 1 1 1 Negative Genetic
TUB3 YML124C Alpha-tubulin genetic 1 1 1 Negative Genetic
YDJ1 YNL064C Type I HSP40 co-chaperone genetic 1 1 1 Negative Genetic
SWM2 YNR004W Protein with a role in snRNA and snoRNA cap trimethylation genetic 1 1 1 Negative Genetic
RCN2 YOR220W Protein of unknown function genetic 1 1 1 Negative Genetic
POC4 YPL144W Component of a heterodimeric Poc4p-Irc25p chaperone genetic 1 1 1 Negative Genetic
TGS1 YPL157W Trimethyl guanosine synthase, conserved nucleolar methyl transferase genetic 1 1 1 Negative Genetic
YPR089W YPR089W Protein of unknown function genetic 1 1 1 Negative Genetic
URN1 YPR152C Protein of unknown function containing WW and FF domains genetic 1 1 1 Negative Genetic
SHP1 YBL058W UBX domain-containing substrate adaptor for Cdc48p genetic 1 1 1 Negative Genetic
YBR221W-A YBR221W-A Putative protein of unknown function genetic 1 1 1 Negative Genetic
RAM1 YDL090C Beta subunit of the CAAX farnesyltransferase (FTase) genetic 1 1 1 Negative Genetic
YDL218W YDL218W Putative protein of unknown function genetic 1 1 1 Negative Genetic
SWI5 YDR146C Transcription factor that recruits Mediator and Swi/Snf complexes genetic 1 1 1 Negative Genetic
SWR1 YDR334W Swi2/Snf2-related ATPase genetic 1 1 1 Negative Genetic
SPT3 YDR392W Subunit of the SAGA and SAGA-like transcriptional regulatory complexes genetic 1 1 1 Negative Genetic
RPL12B YDR418W Ribosomal 60S subunit protein L12B genetic 1 1 1 Negative Genetic
TOG1 YER184C Transcriptional activator of oleate genes genetic 1 1 1 Negative Genetic
RPL9A YGL147C Ribosomal 60S subunit protein L9A genetic 1 1 1 Negative Genetic
YGR174W-A YGR174W-A Putative protein of unknown function genetic 1 1 1 Negative Genetic
PEX21 YGR239C Peroxin required for peroxisomal matrix protein targeting genetic 1 1 1 Negative Genetic
SPT8 YLR055C Subunit of the SAGA transcriptional regulatory complex genetic 1 1 1 Negative Genetic
PER33 YLR064W Protein that localizes to the endoplasmic reticulum genetic 1 1 1 Negative Genetic
PNP1 YLR209C Purine nucleoside phosphorylase genetic 1 1 1 Negative Genetic
ZWF1 YNL241C Glucose-6-phosphate dehydrogenase (G6PD) genetic 1 1 1 Negative Genetic
YNR075C-A YNR075C-A Protein of unknown function genetic 1 1 1 Negative Genetic
RTS1 YOR014W B-type regulatory subunit of protein phosphatase 2A (PP2A) genetic 1 1 1 Negative Genetic
YOR072W-B YOR072W-B Putative protein of unknown function genetic 1 1 1 Negative Genetic
YOR293C-A YOR293C-A Putative protein of unknown function genetic 1 1 1 Negative Genetic
VIK1 YPL253C Protein that forms a kinesin-14 heterodimeric motor with Kar3p genetic 1 1 1 Negative Genetic
SWD1 YAR003W Subunit of the COMPASS (Set1C) complex genetic 1 1 1 Positive Genetic
STI1 YOR027W Evolutionarily-conserved Hsp90 cochaperone genetic 1 1 1 Negative Genetic
VAM7 YGL212W Vacuolar SNARE protein genetic 1 1 1 Positive Genetic
SWC4 YGR002C Component of the Swr1p complex that incorporates Htz1p into chromatin genetic 1 1 1 Negative Genetic
RIM20 YOR275C Protein involved in proteolytic activation of Rim101p genetic 1 1 1 Positive Genetic
ILV6 YCL009C Regulatory subunit of acetolactate synthase genetic 1 1 1 Negative Genetic
ARO8 YGL202W Aromatic aminotransferase I genetic 1 1 1 Negative Genetic
OKP1 YGR179C Outer kinetochore protein required for accurate chromosome segregation genetic 1 1 1 Negative Genetic
CSF1 YLR087C Protein with structural similarity to lipid transport protein Vps13p genetic 1 1 1 Positive Genetic
MMS1 YPR164W Subunit of E3 ubiquitin ligase complex involved in replication repair genetic 1 1 1 Negative Genetic
BAP3 YDR046C Amino acid permease genetic 1 1 1 Negative Genetic
ERI1 YPL096C-A Endoplasmic reticulum membrane protein that binds and inhibits Ras2p genetic 1 1 1 Positive Genetic
DSF2 YBR007C Deletion suppressor of mpt5 mutation genetic 1 1 1 Negative Genetic
HRQ1 YDR291W 3'-5' DNA helicase of the conserved RecQ family physical 1 1 1 Affinity Capture-MS
HRQ1 YDR291W 3'-5' DNA helicase of the conserved RecQ family genetic 1 1 1 Positive Genetic
MSS4 YDR208W Phosphatidylinositol-4-phosphate 5-kinase genetic 1 1 1 Negative Genetic
KXD1 YGL079W Subunit of the BLOC-1 complex involved in endosomal maturation genetic 1 1 1 Negative Genetic
BRL1 YHR036W Essential nuclear envelope/ER integral membrane protein genetic 1 1 1 Positive Genetic
MTC1 YJL123C Protein of unknown function that may interact with ribosomes genetic 1 1 1 Negative Genetic
GEP7 YGL057C Protein of unknown function genetic 1 1 1 Negative Genetic
MSS1 YMR023C Mitochondrial protein physical 1 1 1 Affinity Capture-MS
ORC4 YPR162C Subunit of the origin recognition complex (ORC) genetic 1 1 1 Negative Genetic
MCY1 YGR012W Putative cysteine synthase genetic 1 1 1 Negative Genetic
GPI17 YDR434W Transmembrane protein genetic 1 1 1 Positive Genetic
DPH5 YLR172C Methyltransferase required for synthesis of diphthamide genetic 1 1 1 Negative Genetic
CLB4 YLR210W B-type cyclin involved in cell cycle progression genetic 1 1 1 Negative Genetic
OPY2 YPR075C Integral membrane protein that acts as a membrane anchor for Ste50p genetic 1 1 1 Negative Genetic
ALR2 YFL050C Probable Mg(2+) transporter genetic 1 1 1 Negative Genetic
APJ1 YNL077W Hsp40 chaperone with a role in SUMO-mediated protein degradation physical 1 1 1 Affinity Capture-MS
CDA1 YLR307W Chitin deacetylase genetic 1 1 1 Positive Genetic
ORM1 YGR038W Protein that mediates sphingolipid homeostasis genetic 1 1 1 Negative Genetic
GAS3 YMR215W Putative 1,3-beta-glucanosyltransferase genetic 1 1 1 Positive Genetic
MPS3 YJL019W Nuclear envelope protein physical 1 1 1 Affinity Capture-MS
NCB2 YDR397C Subunit of a heterodimeric NC2 transcription regulator complex genetic 1 1 1 Positive Genetic
KIP3 YGL216W Kinesin-related antiparallel sliding motor protein genetic 1 1 1 Negative Genetic
LDB18 YLL049W Component of the dynactin complex genetic 1 1 1 Negative Genetic
MSA1 YOR066W Activator of G1-specific transcription factors MBF and SBF genetic 1 1 1 Negative Genetic
NEO1 YIL048W Phospholipid translocase (flippase) genetic 1 1 1 Positive Genetic
DYN3 YMR299C Dynein light intermediate chain (LIC) genetic 1 1 1 Negative Genetic
CCW12 YLR110C Cell wall mannoprotein genetic 1 1 1 Negative Genetic
OSI1 YKL071W NADH-dependent aldehyde reductase genetic 1 1 1 Positive Genetic
ATG41 YPL250C Protein of unknown function genetic 1 1 1 Positive Genetic
NCS2 YNL119W Protein required for uridine thiolation of Lys(UUU) and Glu(UUC) tRNAs genetic 1 1 1 Negative Genetic
MNT2 YGL257C Mannosyltransferase genetic 1 1 1 Negative Genetic
CYT2 YKL087C Cytochrome c1 heme lyase genetic 1 1 1 Negative Genetic
EFM4 YIL064W Lysine methyltransferase genetic 1 1 1 Negative Genetic
IRC15 YPL017C Microtubule associated protein genetic 1 1 1 Negative Genetic
EMC2 YJR088C Member of conserved ER transmembrane complex genetic 1 1 1 Negative Genetic
ICT1 YLR099C Lysophosphatidic acid acyltransferase genetic 1 1 1 Negative Genetic
NFG1 YLR042C Cell wall protein of unknown function genetic 1 1 1 Negative Genetic
APQ12 YIL040W Nuclear envelope/ER integral membrane protein genetic 1 1 1 Positive Genetic
KRE5 YOR336W Protein required for beta-1,6 glucan biosynthesis genetic 1 1 1 Positive Genetic