After the whole-genome duplication event leading the evolution of S. cerevisiae and related species, each of the genes that were essential would have been duplicated. The fates of each of these genes can be identified based on the knowledge of existing ohnologs (genes related by the whole-genome duplication), synthetic lethality data, and essentiality data.
The analysis to trace the fates of these duplicated essential genes in the quire programming language follows.
By far the most common event for any duplicated gene was that the duplicate copy was lost.
We can find these genes with quire.
First, let's read a gene registry for all of the S. cerevisiae genes and a list of the essential genes. (Here we are using the list of genes reported to be essential for the reference S288c strain; a small number of genes are different in different strain backgrounds.)
scgenes = registry( @ "scgene_registry.txt" )
essential = genelist( @ "essential.txt", generegistry=scgenes )
Second, we read in a network in which genes related by the whole-genome duplication are reported as "ohnolog" interactions. When the network of paralogs is read in, we specify that the symmetry is "all". This means that the A:B interaction is also stored as the B:A interaction. The actual file also has other non-whole genome duplication relationships that are labeled as "paralog" interactions, so we need to filter out only the ohnolog interactions.
paralogs_net = network( @ "paralogs.txt", generegistry=scgenes, symmetry="all", format="sif" )
ohnologs_net = filter( paralogs_net, type="ohnolog" )
Because we read in the network with a symmetry of "all", we can extract all of the genes by getting the "bait" or the "prey" genes.
ohnologs = element( ohnologs_net, type="baits" )
We can then identify the essential genes that do not have duplications in S. cerevisiae identifying those essential genes that are not present in the list of ohnologs.
essential_with_deletion = essential + ohnologs -> filter( partition="10" )
"Essential genes = " + sizeof( essential )
"Essential genes not in ohnolog pairs = " + sizeof( essential_with_deletion )
And the output is:
Essential genes = 1060
Essential genes not in ohnolog pairs = 1018
Thus only 42 of the 1060 essential genes have not lost their duplicates (ohnologs) arising from the whole-genome duplication event. Here we won't list the 1018 gene not in ohnolog pairs.
One interesting possibility is that divergence of the one of the copies allows it to adopt a different function, since the other still supports the essential function.
We can identify these types of pairs using quire.
First, let's read a gene registry for all of the S. cerevisiae genes and a list of the essential genes. (Here we are using the list of genes reported to be essential for the reference S288c strain; a small number of genes are different in different strain backgrounds.)
scgenes = registry( @ "scgene_registry.txt" )
essential = genelist( @ "essential.txt", generegistry=scgenes )
Second, we read in a network in which genes related by the whole-genome duplication are reported as "ohnolog" interactions. The actual file also has other non-whole genome duplication relationships that are labeled as "paralog" interactions, so we need to filter out only the ohnolog interactions.
When the network of paralogs is read in, we specify that the symmetry is "all". This means that the A:B interaction is also stored as the B:A interaction. If we had specified the symmetry as "none", we would only store the interaction as an A:B interaction. This is important in the essential gene paralog filtering that will happen later.
paralogs_net = network( @ "paralogs.txt", generegistry=scgenes, symmetry="all", format="sif" )
ohnologs_net = filter( paralogs_net, type="ohnolog" )
Third, we label all of the interactions based on which of the genes involved in the interaction are in list of essential genes and filter out those with only one essential gene. In this case, the interactions are given a gene partition label of "0:0" if neither gene is essential, "1:0" or "0:1" if one gene is essential, and "1:1" if both genes are essential.
Because we read the network in using the symmetry set to "all", filtering out those interactions with gene partition labels of "1:0" or "0:1" will give us each interaction twice. If the network had been read in with the symmetry set to "none", then we would have needed both. However, the advantage of using symmetry "all" with filtering for gene partition labels of "1:0" means that we always know that the essential gene is the first one (the "bait") and the non-essential gene is the second one (the "prey").
ohnologs_net::label( genepartition=essential )
only_one_essential = filter( ohnologs_net, genepartition="1:0" )
Fourth, and finally, we can now investigate all of the ohnolog pairs with one essential and one non-essential gene by examining the interactions in the resulting network.
In the following for loop, the value i is assigned to each interaction in the network. In quire, the value is provided as a list data type containing the named elements bait, prey, partition, genepartition, and obs. The obs named element is a list of lists containing each observation for that interaction. These observation sublists contain the named elements pmid, type, id, obspartition and symmgen. Here, only the bait and prey named elements of the interaction list are important.
n = 1
for i in only_one_essential
"Interaction " + n
"\tEssential: " + i["bait"] + " " + tolocus( i["bait"], scgenes )
"\tNon-essential: " + i["prey"] + " " + tolocus( i["prey"], scgenes )
""
n = n+1
rof
The results are:
Interaction 1
Essential: YDR303C RSC3
Non-essential: YHR056C RSC30
Interaction 2
Essential: YOR326W MYO2
Non-essential: YAL029C MYO4
Interaction 3
Essential: YMR113W FOL3
Non-essential: YKL132C RMA1
Interaction 4
Essential: YOR110W TFC7
Non-essential: YNL108C YNL108C
Interaction 5
Essential: YNR026C SEC12
Non-essential: YCR067C SED4
Interaction 6
Essential: YKL180W RPL17A
Non-essential: YJL177W RPL17B
Interaction 7
Essential: YPL228W CET1
Non-essential: YMR180C CTL1
Interaction 8
Essential: YLR457C NBP1
Non-essential: YPR174C CSA1
Interaction 9
Essential: YOR182C RPS30B
Non-essential: YLR287C-A RPS30A
Interaction 10
Essential: YLR293C GSP1
Non-essential: YOR185C GSP2
Interaction 11
Essential: YJR045C SSC1
Non-essential: YEL030W ECM10
Interaction 12
Essential: YMR047C NUP116
Non-essential: YKL068W NUP100
Interaction 13
Essential: YLR249W YEF3
Non-essential: YNL014W HEF3
Interaction 14
Essential: YCR052W RSC6
Non-essential: YNR023W SNF12
Interaction 15
Essential: YMR079W SEC14
Non-essential: YKL091C YKL091C
Interaction 16
Essential: YJL026W RNR2
Non-essential: YGR180C RNR4
Interaction 17
Essential: YLR223C IFH1
Non-essential: YDR223W CRF1
Interaction 18
Essential: YOL066C RIB2
Non-essential: YDL036C PUS9
Interaction 19
Essential: YDR341C YDR341C
Non-essential: YHR091C MSR1
Interaction 20
Essential: YCL043C PDI1
Non-essential: YDR518W EUG1
Interaction 21
Essential: YKL035W UGP1
Non-essential: YHL012W YHL012W
Interaction 22
Essential: YML085C TUB1
Non-essential: YML124C TUB3
Interaction 23
Essential: YNL162W RPL42A
Non-essential: YHR141C RPL42B
Interaction 24
Essential: YLR310C CDC25
Non-essential: YLL016W SDC25
Interaction 25
Essential: YLR310C CDC25
Non-essential: YLL017W YLL017W
Interaction 26
Essential: YBL030C PET9
Non-essential: YBR085W AAC3
Interaction 27
Essential: YKL104C GFA1
Non-essential: YMR084W YMR084W
Interaction 28
Essential: YKL104C GFA1
Non-essential: YMR085W YMR085W
Interaction 29
Essential: YPR162C ORC4
Non-essential: YLR453C RIF2
Interaction 30
Essential: YGL225W VRG4
Non-essential: YER039C HVG1
Interaction 31
Essential: YML125C PGA3
Non-essential: YML087C AIM33
Interaction 32
Essential: YOL120C RPL18A
Non-essential: YNL301C RPL18B
Interaction 33
Essential: YAL038W CDC19
Non-essential: YOR347C PYK2
Interaction 34
Essential: YNR016C ACC1
Non-essential: YMR207C HFA1
Interaction 35
Essential: YEL034W HYP2
Non-essential: YJR047C ANB1
Interaction 36
Essential: YLR029C RPL15A
Non-essential: YMR121C RPL15B
Interaction 37
Essential: YOR204W DED1
Non-essential: YPL119C DBP1
Interaction 38
Essential: YBR049C REB1
Non-essential: YDR026C NSI1
Interaction 39
Essential: YOR167C RPS28A
Non-essential: YLR264W RPS28B
Interaction 40
Essential: YGL075C MPS2
Non-essential: YPL200W CSM4
Interaction 41
Essential: YKL141W SDH3
Non-essential: YMR118C SHH3
Interaction 42
Essential: YLR277C YSH1
Non-essential: YOR179C SYC1
Interaction 43
Essential: YKL203C TOR2
Non-essential: YJR066W TOR1
Interaction 44
Essential: YML065W ORC1
Non-essential: YLR442C SIR3
Several different kinds of diversification can be observed in this list.
(1) Some ohnolog pairs contain genes that have adopted different functions and one of the pair has lost the "essential" function. For example, in the TOR2/TOR1 pair, both genes encode protein kinases that play a role in nutrient sensing, but only TOR2 maintains an essential role in actin polarization. In the ORC4/RIF2 pair, ORC4 is an essential gene that encodes a subunit of the DNA replication origin recognition complex (Orc1-6), whereas RIF2 encodes a protein that binds Rap1 and helps control telomere length and telomeric silencing. Remarkably, Rif2 does not appear to interact with any of the Orc1-6 subunits. Similarly, ORC1 encodes a member of the origin-recognition complex, whereas SIR3 is involved in transcriptional silencing and is recruited to DNA through interaction with Rap1. In the RSC6/SNF12 pair, RSC6 encodes a subunit of the RSC chromatin remodeling complex, and SNF12 encodes a subunit of the SWI/SNF chromatin remodeling complex.
(2) Some ohnolog pairs contain genes in which one gene in the pair is split: GFA1/YMR084W-YMR085W * and CDC25/SDC25-YLL017W. In strains related to S288c, YMR084W-YMR085W exists as a single full-length gene and as do SDC25-YLL017W*. This sort of mutagenic loss of the copy is unsurprising in a situation where there are two redundant genes.
(3) Some ohnolog pairs appear to have alternative regulation: RNR2/RNR4, TUB1/TUB3, and CDC19/PYK2. In some cases, such as RPL15A/RPL15B, one of the genes is very poorly expressed, which is conceptually similar to the mutagenic copy loss of genes in category 2.
(4) Some ohnolog pairs appear to have changes in subcellular localization: ACC1/HFA1, FOL3/RMA1, and YDR341C/MSR1.
In cases where an essential gene plays an essential role in two different processes, it's possible to envision that gene duplication could allow each copy to specialize for each process. In this case, each of the duplicates would be expected to be essential.
We can identify these types of pairs using quire.
First, let's read a gene registry for all of the S. cerevisiae genes and a list of the essential genes. (Here we are using the list of genes reported to be essential for the reference S288c strain; a small number of genes are different in different strain backgrounds.)
scgenes = registry( @"scgene_registry.txt" )
essential = genelist( @"essential.txt", generegistry=scgenes )
Second, we read in a network in which genes related by the whole-genome duplication are reported as "ohnolog" interactions. The actual file also has other non-whole genome duplication relationships that are labeled as "paralog" interactions, so we need to filter out only the ohnolog interactions.
When the network of paralogs is read in, we specify that the symmetry is "none", unlike the cases described above. This means that the A:B interaction are only stored as the B:A interaction. If we had specified the symmetry as "all", we would store the interaction both A:B and B:A interactions. This would duplicate all of the interactions we are going to label below.
paralogs_net = network( @ QUIRE_DIR+ "scerevisiae/paralogs.txt", generegistry=scgenes, symmetry="none", format="sif" )
ohnologs_net = filter( paralogs_net, type="ohnolog" )
Third, we label all of the interactions based on which of the genes involved in the interaction are in list of essential genes and filter out those with only one essential gene. In this case, the interactions are given a gene partition label of "0:0" if neither gene is essential, "1:0" or "0:1" if one gene is essential, and "1:1" if both genes are essential.
ohnologs_net::label( genepartition=essential )
two_essential = filter( ohnologs_net, genepartition="1:1" )
sizeof( two_essential )
And the output is zero, as there are no ohnolog pairs where both genes are essential. To confirm this, we can count the number of ohnolog pairs with no essential genes, add it to the 44 ohnolog pairs identified above with one essential gene, and ensure that it matches the total number of ohnolog pairs.
no_essential = filter( ohnologs_net, genepartition="0:0" )
"Count = " + sizeof( no_essential ) + " + 44 + " + sizeof( two_essential ) + " = " + ( sizeof(no_essential) + 44 )
"Total = " + sizeof( ohnologs_net )
And the output is:
Count = 506 + 44 + 0 = 550
Total = 550
A final interesting possibility is that S. cerevisiae has retained both copies of the gene in the ohnolog pair. In this case, the gene that was essential in the pre-WGD species is now no long essential; however, simultaneous deletion of both copies should be lethal. Hence the genes in the ohnolog pair should show a synthetic lethal interaction.
We can identify these types of pairs using quire.
First, let's read a gene registry for all of the S. cerevisiae genes.
scgenes = registry( @"scgene_registry.txt" )
Second, we read in a network in which genes related by the whole-genome duplication are reported as "ohnolog" interactions. The actual file also has other non-whole genome duplication relationships that are labeled as "paralog" interactions, so we need to filter out only the ohnolog interactions.
When the network of paralogs is read in, we specify that the symmetry is "none". This means that the A:B interaction is not also stored as the B:A interaction. Because the paralog file was created non-redundantly, this means that after combining with the synthetic lethal network, we don't see both the A:B and B:A version of each interaction.
paralogs_net = network( @ "paralogs.txt", generegistry=scgenes, symmetry="none", format="sif" )
ohnologs_net = filter( paralogs_net, type="ohnolog" )
Third, we need to read in the synthetic lethal interaction data for genes in S. cerevisiae. We will do this reading in a (modified) version of the BioGRiD interaction database, and extract out those interactions corresponding to "Synthetic Lethality". We will read in the data with a symmetry type of "all" to ensure that we don't miss overlaps with the ohnolog network.
biogridfile = "BIOGRID-ORGANISM-Saccharomyces_cerevisiae_S288c-3.5.168-modified.tab2.txt"
biogrid_net = network( @ biogridfile, format="biogrid_tab2", generegistry=scgenes, symmetry="all", sourcename="biogrid", organismbait="559292", organismprey="559292" )
sl_net = filter( biogrid_net, type="Synthetic Lethality" )
Fourth, we will extract out the gene pairs that are both synthetic lethal and ohnologs by adding together the two networks and extracting those interactions in common. Interactions in common have a partition of "11", which means that they were present in the ohnolog network (first "1") and in the synthetic lethality network (second "1").
sl_ohnologs = ohnologs_net + sl_net -> filter( partition="11" )
sizeof( sl_ohnologs )
This analysis gives us 110 gene pairs (we would have 220 gene pairs with both A:B and B:A version of each interaction if we had read the ohnolog network with a symmetry of "all"). We can output these with the following for loop:
n = 1
for i in sl_ohnologs
"Interaction " + n
"\tGene A: " + i["bait"] + " " + tolocus( i["bait"], scgenes )
"\tGene B: " + i["prey"] + " " + tolocus( i["prey"], scgenes )
""
n = n+1
rof
This gives us the result:
~~~
Interaction 1
Gene A: YBR048W RPS11B
Gene B: YDR025W RPS11A
Interaction 2
Gene A: YLR264W RPS28B
Gene B: YOR167C RPS28A
Interaction 3
Gene A: YGR118W RPS23A
Gene B: YPR132W RPS23B
Interaction 4
Gene A: YNL301C RPL18B
Gene B: YOL120C RPL18A
Interaction 5
Gene A: YLR448W RPL6B
Gene B: YML073C RPL6A
Interaction 6
Gene A: YMR246W FAA4
Gene B: YOR317W FAA1
Interaction 7
Gene A: YDL083C RPS16B
Gene B: YMR143W RPS16A
Interaction 8
Gene A: YML124C TUB3
Gene B: YML085C TUB1
Interaction 9
Gene A: YHR021C RPS27B
Gene B: YKL156W RPS27A
Interaction 10
Gene A: YBR031W RPL4A
Gene B: YDR012W RPL4B
Interaction 11
Gene A: YBR169C SSE2
Gene B: YPL106C SSE1
Interaction 12
Gene A: YGL071W AFT1
Gene B: YPL202C AFT2
Interaction 13
Gene A: YDR099W BMH2
Gene B: YER177W BMH1
Interaction 14
Gene A: YIL105C SLM1
Gene B: YNL047C SLM2
Interaction 15
Gene A: YGR085C RPL11B
Gene B: YPR102C RPL11A
Interaction 16
Gene A: YIL149C MLP2
Gene B: YKR095W MLP1
Interaction 17
Gene A: YBR161W CSH1
Gene B: YPL057C SUR1
Interaction 18
Gene A: YHL001W RPL14B
Gene B: YKL006W RPL14A
Interaction 19
Gene A: YMR186W HSC82
Gene B: YPL240C HSP82
Interaction 20
Gene A: YER056C-A RPL34A
Gene B: YIL052C RPL34B
Interaction 21
Gene A: YGL076C RPL7A
Gene B: YPL198W RPL7B
Interaction 22
Gene A: YIL133C RPL16A
Gene B: YNL069C RPL16B
Interaction 23
Gene A: YOR054C VHS3
Gene B: YKR072C SIS2
Interaction 24
Gene A: YMR230W RPS10B
Gene B: YOR293W RPS10A
Interaction 25
Gene A: YDR471W RPL27B
Gene B: YHR010W RPL27A
Interaction 26
Gene A: YLR287C-A RPS30A
Gene B: YOR182C RPS30B
Interaction 27
Gene A: YMR199W CLN1
Gene B: YPL256C CLN2
Interaction 28
Gene A: YGR109C CLB6
Gene B: YPR120C CLB5
Interaction 29
Gene A: YDL229W SSB1
Gene B: YNL209W SSB2
Interaction 30
Gene A: YGR214W RPS0A
Gene B: YLR048W RPS0B
Interaction 31
Gene A: YOR234C RPL33B
Gene B: YPL143W RPL33A
Interaction 32
Gene A: YDR144C MKC7
Gene B: YLR120C YPS1
Interaction 33
Gene A: YBR181C RPS6B
Gene B: YPL090C RPS6A
Interaction 34
Gene A: YDR436W PPZ2
Gene B: YML016C PPZ1
Interaction 35
Gene A: YMR121C RPL15B
Gene B: YLR029C RPL15A
Interaction 36
Gene A: YOR231W MKK1
Gene B: YPL140C MKK2
Interaction 37
Gene A: YAL028W FRT2
Gene B: YOR324C FRT1
Interaction 38
Gene A: YER131W RPS26B
Gene B: YGL189C RPS26A
Interaction 39
Gene A: YCR031C RPS14A
Gene B: YJL191W RPS14B
Interaction 40
Gene A: YDR450W RPS18A
Gene B: YML026C RPS18B
Interaction 41
Gene A: YKL068W NUP100
Gene B: YMR047C NUP116
Interaction 42
Gene A: YLR441C RPS1A
Gene B: YML063W RPS1B
Interaction 43
Gene A: YAL030W SNC1
Gene B: YOR327C SNC2
Interaction 44
Gene A: YGR070W ROM1
Gene B: YLR371W ROM2
Interaction 45
Gene A: YDR502C SAM2
Gene B: YLR180W SAM1
Interaction 46
Gene A: YJL129C TRK1
Gene B: YKR050W TRK2
Interaction 47
Gene A: YBR118W TEF2
Gene B: YPR080W TEF1
Interaction 48
Gene A: YDR490C PKH1
Gene B: YOL100W PKH2
Interaction 49
Gene A: YDR500C RPL37B
Gene B: YLR185W RPL37A
Interaction 50
Gene A: YBR191W RPL21A
Gene B: YPL079W RPL21B
Interaction 51
Gene A: YGL135W RPL1B
Gene B: YPL220W RPL1A
Interaction 52
Gene A: YDR385W EFT2
Gene B: YOR133W EFT1
Interaction 53
Gene A: YER019C-A SBH2
Gene B: YER087C-B SBH1
Interaction 54
Gene A: YMR192W GYL1
Gene B: YPL249C GYP5
Interaction 55
Gene A: YDR312W SSF2
Gene B: YHR066W SSF1
Interaction 56
Gene A: YIL148W RPL40A
Gene B: YKR094C RPL40B
Interaction 57
Gene A: YJL177W RPL17B
Gene B: YKL180W RPL17A
Interaction 58
Gene A: YDL191W RPL35A
Gene B: YDL136W RPL35B
Interaction 59
Gene A: YBL027W RPL19B
Gene B: YBR084C-A RPL19A
Interaction 60
Gene A: YNL096C RPS7B
Gene B: YOR096W RPS7A
Interaction 61
Gene A: YDL061C RPS29B
Gene B: YLR388W RPS29A
Interaction 62
Gene A: YGL049C TIF4632
Gene B: YGR162W TIF4631
Interaction 63
Gene A: YNL098C RAS2
Gene B: YOR101W RAS1
Interaction 64
Gene A: YMR194W RPL36A
Gene B: YPL249C-A RPL36B
Interaction 65
Gene A: YMR109W MYO5
Gene B: YKL129C MYO3
Interaction 66
Gene A: YDL082W RPL13A
Gene B: YMR142C RPL13B
Interaction 67
Gene A: YHR135C YCK1
Gene B: YNL154C YCK2
Interaction 68
Gene A: YGR192C TDH3
Gene B: YJR009C TDH2
Interaction 69
Gene A: YDL175C AIR2
Gene B: YIL079C AIR1
Interaction 70
Gene A: YJL136C RPS21B
Gene B: YKR057W RPS21A
Interaction 71
Gene A: YIR033W MGA2
Gene B: YKL020C SPT23
Interaction 72
Gene A: YDR447C RPS17B
Gene B: YML024W RPS17A
Interaction 73
Gene A: YDL192W ARF1
Gene B: YDL137W ARF2
Interaction 74
Gene A: YMR104C YPK2
Gene B: YKL126W YPK1
Interaction 75
Gene A: YDL161W ENT1
Gene B: YLR206W ENT2
Interaction 76
Gene A: YGR124W ASN2
Gene B: YPR145W ASN1
Interaction 77
Gene A: YJR094W-A RPL43B
Gene B: YPR043W RPL43A
Interaction 78
Gene A: YBR189W RPS9B
Gene B: YPL081W RPS9A
Interaction 79
Gene A: YER027C GAL83
Gene B: YGL208W SIP2
Interaction 80
Gene A: YER031C YPT31
Gene B: YGL210W YPT32
Interaction 81
Gene A: YGR108W CLB1
Gene B: YPR119W CLB2
Interaction 82
Gene A: YBL079W NUP170
Gene B: YER105C NUP157
Interaction 83
Gene A: YGR032W GSC2
Gene B: YLR342W FKS1
Interaction 84
Gene A: YDL188C PPH22
Gene B: YDL134C PPH21
Interaction 85
Gene A: YEL022W GEA2
Gene B: YJR031C GEA1
Interaction 86
Gene A: YDR303C RSC3
Gene B: YHR056C RSC30
Interaction 87
Gene A: YOR226C ISU2
Gene B: YPL135W ISU1
Interaction 88
Gene A: YBL085W BOI1
Gene B: YER114C BOI2
Interaction 89
Gene A: YBL068W PRS4
Gene B: YER099C PRS2
Interaction 90
Gene A: YNL302C RPS19B
Gene B: YOL121C RPS19A
Interaction 91
Gene A: YBL087C RPL23A
Gene B: YER117W RPL23B
Interaction 92
Gene A: YGR143W SKN1
Gene B: YPR159W KRE6
Interaction 93
Gene A: YDR309C GIC2
Gene B: YHR061C GIC1
Interaction 94
Gene A: YIL131C FKH1
Gene B: YNL068C FKH2
Interaction 95
Gene A: YBR082C UBC4
Gene B: YDR059C UBC5
Interaction 96
Gene A: YLR450W HMG2
Gene B: YML075C HMG1
Interaction 97
Gene A: YMR242C RPL20A
Gene B: YOR312C RPL20B
Interaction 98
Gene A: YHL033C RPL8A
Gene B: YLL045C RPL8B
Interaction 99
Gene A: YNL299W TRF5
Gene B: YOL115W PAP2
Interaction 100
Gene A: YHR203C RPS4B
Gene B: YJR145C RPS4A
Interaction 101
Gene A: YHR141C RPL42B
Gene B: YNL162W RPL42A
Interaction 102
Gene A: YHL003C LAG1
Gene B: YKL008C LAC1
Interaction 103
Gene A: YER074W RPS24A
Gene B: YIL069C RPS24B
Interaction 104
Gene A: YEL034W HYP2
Gene B: YJR047C ANB1
Interaction 105
Gene A: YDR409W SIZ1
Gene B: YOR156C NFI1
Interaction 106
Gene A: YBL039C URA7
Gene B: YJR103W URA8
Interaction 107
Gene A: YGR092W DBF2
Gene B: YPR111W DBF20
Interaction 108
Gene A: YBL072C RPS8A
Gene B: YER102W RPS8B
Interaction 109
Gene A: YFR031C-A RPL2A
Gene B: YIL018W RPL2B
Interaction 110
Gene A: YER062C GPP2
Gene B: YIL053W GPP1
~~~
There is a very clear signal of ohnologs that duplicate genes encoding protein components of both the small and large ribosomal subunits (RPS0A/B, RPS1A/B, RPS4A/B, RPS6A/B, RPS7A/B, RPS8A/B, RPS9A/B, RPS10A/B, RPS11A/B, RPS14A/B, RPS16A/B, RPS17A/B, RPS18A/B, RPS19A/B, RPS21A/B, RPS23A/B, RPS24A/B, RPS26A/B, RPS27A/B, RPS28A/B, RSP29A/B, RPS30A/B, RPL1A/B, RPL2A/B, RPL4A/B, RPL6A/B, RPL7A/B, RPL8A/B, RPL11A/B, RPL13A/B, RPL14A/B, RPL15A/B, RPL17A/B, RPL18A/B, RPL19A/B, RPL20A/B, RPL21A/B, RPL23A/B RPL27A/B, RPL33A/B, RPL34A/B, RPL35A/B, RPL36A/B, RPL37A/B, RPL40A/B, RPL43A/B). Remarkably, RPL15A/B appears as both a synthetic lethal pair and as a pair in which one of the genes is essential, which likely also represent differences between S. cerevisiae strains.
Additionally, the ohnolog pair RSC3/RSC30 appears as a pair in which only one of the two genes is essential (RSC3) or both of the genes are redundant. This is due to differences in essential genes among different S. cerevisiae strains. In the S288c background, RSC3 is essential, whereas it is not in the W303 background.
Others: ANB1/HYP2, AFT1/2, AIR1/2, ARF1/2, ASN1/2 BMH1/2, BOI1/2, CLB1/2, CLB5/6, CLN1/2, CSH1/SUR1, DBF2/20, EFT1/2, ENT1/2, FAA1/4, FHK1/2, FKS1/GSC2, FRT1/2, GAL83/SIP2 GEA1/2, GIC1/2, GPP1/2, GYL1/GYP5, HSC82/HSP82, HMG1/2, ISU1/2, KRE6/SKN1, LAC1/LAG1 MGA2/SPT23, MKC7/YPS1, MKK1/2, MLP1/2, MYO3/5, NFI1/SIZ1 NUP100/NUP116, NUP170/157, PAP2/TRF5, PKH1/2, PPH21/22 PPZ1/2, PRS2/4, RAS1/2, ROM1/2, RSC3/30, SAM1/2, SBH1/2, SIS2/VHS3, SLM1/2, SNC1/2, SSB1/2, SSE1/2, SSF1/2, TIF4631/4632 TEF1/2, TDH2/3, TRK1/2, TUB1/3, UBC4/5, URA7/8, YCK1/2, YPK1/2, YPT31/32