Table 1.
List of all Sclerotinia sclerotiorum genes studied so far using mutant analysis. The genes are ordered in regards to their chromosomal positions (Figure 3).
| Gene Code (New) | Gene Code (Old) | Strains | Mutant Name | Gene Full Name | Mutant Type | Mutant Phenotypes | Secretion Signal | Tested Hosts | Other Functions | Reference | |||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Hyphal Growth | Sclerotial Formation | Oxalate Production | Virulence | Appressoria Formation | Induce Host HR/Resistance | ||||||||||
| sscle_01g003850 | SS1G_02068 | Ep-1PNA367, Heilongjiang, China | ssvp1 | Small secreted virulence-related protein | RNAi | - | NA | NA | + | NA | + | Yes | Canola (Zhongyou 821) | HR induction | [30] |
| sscle_01g004990 | SS1G_01919 | 1980, Nebraska, USA | svf1 | Survival factor 1 homologue | RNAi | - | NA | - | + | + | NA | No | Arabidopsis and canola (Zhongyou 821) | Cell wall integrity, ROS production | [31] |
| sscle_01g006030 | SS1G_01788 | 1980, Nebraska, USA | cna1 | Catalytic subunit calcineurin-encoding gene | RNAi | + | + | - | + | - | NA | No | Arabiposis and tomato (Bonny Best) | Hyphal elongation | [32] |
| sscle_01g007450 | SS1G_01614 | Ep-1PNA367, Heilongjiang, China | sop1 | Microbial opsin homolog gene | RNAi | + | + | NA | + | NA | NA | No | Arabidopsis | Stress responses | [33] |
| sscle_01g011030 | SS1G_01151 | 1980, Nebraska, USA | sfh1 | GATA-box and SNF5 domains containing transcription factor | RNAi | + | + | NA | + | + | NA | No | Soybean, common bean and tomato | ROS accumulation | [34] |
| sscle_02g013550 | SS1G_12694 | HA61, Jiangsu, China | shk1 | Histidine kinases | Deletion | + | + | NA | - | NA | NA | No | Rapeseed, strawberry, tomato and cucumber | Oxidative stresses, glycerol accumulation | [35] |
| sscle_02g020240 | SS1G_04003 | 1980, Nebraska, USA | mat1-1-5 | Mating-type gene | Deletion | - | - | NA | - | NA | NA | No | Tomato | Apothecial development | [29] |
| sscle_02g020250 | SS1G_04004 | 1980, Nebraska, USA | mat1-1-1 | Mating-type gene | Deletion | - | - | NA | - | NA | NA | No | Tomato | Apothecial development | [29] |
| sscle_02g020260 | SS1G_04005 | 1980, Nebraska, USA | mat1-2-4 | Mating-type gene | Deletion | - | - | NA | - | NA | NA | No | Tomato | Apothecial development, ascospore production | [29] |
| sscle_02g020270 | SS1G_04006 | 1980, Nebraska, USA | mat1-2-1 | Mating-type gene | Deletion | + | - | NA | - | NA | NA | No | Tomato | Apothecial development | [29] |
| sscle_03g025030 | SS1G_00699 | WMA1, Washington, USA | sod1 | Cu/Zn superoxide dismutase | T-DNA | - | - | - | + | NA | NA | No | Pea (Guido) | Detoxification of host ROS | [36] |
| sscle_03g025030 | SS1G_00699 | 1980, Nebraska, USA | sod1 | Cu/Zn superoxide dismutase | Deletion | + | + | + | + | NA | NA | No | Tomato (Garden Peach) and tobacco | Oxidative stress tolerance, repression of host ROS | [37] |
| sscle_03g028510 | SS1G_00263 | Isolated from an infected canola stem, Alberta, Canada | v263 | Hypothetical secreted protein | Deletion | - | NA | NA | + | NA | NA | Yes | Canola | [38] | |
| sscle_03g031470 | SS1G_13314 | Isolated from diseased rapeseed, Alberta, Canada | scd1 | Scytalone dehydratase | Deletion | + | + | NA | - | NA | NA | No | Rapeseed (Westar) | Hyphal branching | [39] |
| sscle_03g031480 | SS1G_13315 | Isolated from diseased rapeseed, Alberta, Canada | thr1 | Trihydroxynaphthalene reductase | Deletion | + | + | NA | - | NA | NA | No | Rapeseed (Westar) | Hyphal branching | [39] |
| sscle_03g031520 | SS1G_13322 | 1980, Nebraska, U.S.A.; UF1, Florida; WMA, Washington, USA | pks13 | Polyketide synthase | Deletion by CRISPR | - | - | NA | - | + | NA | No | Soybean, canola, tomato (Better Boy), faba bean (Windsor) and pea (Sugar Daddy) | Pigmentation of compound appressoria | [40] |
| sscle_03g031670 | SS1G_13339 | 274, Nebraska, USA | pth2 | Peroxysomal carnitine acetyl transferase | Deletion | + | + | + | + | + | NA | No | Soybean | Oxalic acid accumulation | [41] |
| sscle_04g034810 | SS1G_02462 | Isolated from an infected canola stem, Alberta, Canada | axp | Arabinofuranosidase/β-xylosidase | Deletion | NA | NA | NA | + | NA | NA | Yes | Canola | [42] | |
| sscle_04g034960 | SS1G_02486 | Sunf-M, Inner Mongolia, China | caf1 | Secreted protein with a putative Ca2+-binding EF-hand motif | RNAi, T-DNA | + | + | - | + | + | + | Yes | Arabidopsis, rapeseed, pak choi cabbage, hot pepper, tomato, cucumber and soybean | [43] | |
| sscle_04g037170 | SS1G_02784 | 1980, Nebraska, USA | scat1 | Type A catalase | Deletion | + | + | NA | + | NA | NA | No | Tomato (Garden Peach) | Modulation of ROS | [44] |
| sscle_04g038020 | SS1G_02904 | Ep-1PNA367, Heilongjiang, China | cvnh | Secreted protein | RNAi | + | + | NA | + | NA | NA | Yes | Arabidopsis | [45] | |
| sscle_05g046240 | SS1G_05917 | SUN-F-M, Inner Mongolia, China | sl2 | Cell wall protein | RNAi | - | + | NA | - | NA | NA | Yes | Rapeseed and Arabidopsis | Cellular integrity | [46] |
| sscle_05g046390 | SS1G_05899 | 1980, Nebraska, USA | trr1 | Thioredoxin reductase | RNAi | NA | + | NA | + | NA | NA | No | Arabidopsis and tobacco | Oxidative stress tolerance | [47] |
| sscle_05g046790 | SS1G_05839 | 1980, Nebraska, USA | bi1 | Bax inhibitor-1 protein | RNAi | + | - | - | + | NA | NA | No | Arabidopsis and tomato | Stress responses, hyphal tip branching | [48] |
| sscle_05g046830 | SS1G_05834 | JRUF1, Florida, USA | foxe2 | Forkhead-box transcription factor family gene | Deletion, T-DNA | - | - | NA | - | - | NA | No | Tomato | Apothecial development | [49] |
| sscle_05g048220 | SS1G_05661 | 1980, Nebraska, USA | nox1 | NADPH oxidase | RNAi | - | + | + | + | NA | NA | No | Tomato (Rutger) | ROS regulation | [50] |
| sscle_06g049430 | SS1G_07404 | 1980, Nebraska, USA | rhs1 | Rearrangement hot spot repeat-containing protein | RNAi | + | + | - | + | + | NA | Yes | Arabidopsis and canola (Zhongshuang 9) | [51] | |
| sscle_06g049780 | SS1G_07360 | 1980, Nebraska, USA | fkh1 | Atypical forkhead (FKH)-box-containing protein | RNAi | + | + | NA | + | NA | NA | No | Tomato | Cellular integrity | [52] |
| sscle_06g049830 | SS1G_07355 | 1980, Nebraska, USA | pac1 | pH-Responsive transcription factor | Deletion | - | + | + | + | NA | NA | No | Arabidopsis and tomato (Bonnie Best) | [53] | |
| sscle_06g049890 | SS1G_07345 | NGA4, Anhui, China | pemg1 | Elicitor-homologous protein | RNAi | + | NA | - | + | + | NA | No | Oilseed rape and soybean | Negative regulator of growth and virulence | [54] |
| sscle_06g051560 | SS1G_07136 | UF1, Florida, USA | ste12 | Downstream transcription factor of MAPK pathway | RNAi | + | + | NA | + | + | NA | No | Bush bean and tomato | [13] | |
| sscle_07g055970 | SS1G_03527 | SUN-F-M, Inner Mongolia, China | hex1 | Woronin body major protein | RNAi | - | + | NA | NA | NA | NA | No | Cellular integrity | [46] | |
| sscle_07g058030 | SS1G_03252 | UF1, Florida, USA | ams2 | Cell-cycle-regulated GATA transcription factor | RNAi | + | - | NA | + | + | NA | No | Common bean | Chromosome segregation, number and distribution of sclerotia | [55] |
| sscle_07g058620 | SS1G_03171 | 1980, Nebraska, USA | pka1 | Protein kinase A | Deletion | - | - | - | NA | NA | NA | No | PKA activity | [56] | |
| sscle_07g059700 | MK992913 | 1980, Nebraska, USA | qdo | Quercetin dioxygenase gene | Deletion | - | NA | NA | + | NA | NA | Yes | Arabidopsis | Flavonol degradation | [57] |
| sscle_08g064670 | SS1G_05163 | 1980, Nebraska, USA | cry1 | Cryptochrome family CRY-DASH ortholog | Deletion | - | - | NA | - | NA | NA | No | Arabidopsis and tomato | Sclerotial mass, response to UV | [58] |
| sscle_08g065550 | SS1G_05284 | NGA4, Anhui, China | nacα | Nascent polypeptide-associated complex α-subunit | RNAi | - | - | - | + | NA | NA | No | Oilseed rape and tobacco | [59] | |
| sscle_08g066770 | SS1G_05445 | 1980, Nebraska, USA | smk3 | Slt2 ortholog | Deletion | + | + | NA | + | + | NA | No | Canola (Westar) | Cuticle penetration, cell wall integrity | [60] |
| sscle_08g067830 | SS1G_05588 | 1980, Nebraska, U.S.A. | mads | MADS-box proteins | RNAi | + | - | NA | + | NA | NA | No | Tomato | [61] | |
| sscle_08g068500 | SS1G_14133 | Ep-1PNA367, Heilongjiang, China | itl | Integrin-like protein | RNAi | + | + | NA | + | NA | + | Yes | Arabidopsis and canola | Hyphal branching, suppression of host defense | [62] |
| sscle_08g068530 | SS1G_14127 | 1980, Nebraska, USA | ggt1 | γ-Glutamyl transpeptidase | Deletion | - | + | NA | - | + | NA | Yes | Tomato | [63] | |
| sscle_09g069850 | SS1G_10796 | 1980, Nebraska, USA | odc2 | Oxalate decarboxylases | Deletion | - | - | + | + | + | NA | Yes | Common bean (Bush Blue Lake 47), soybean (Harosoy), tomato (Bonnie Best) and celery | [64] | |
| sscle_10g075560 | SS1G_08218 | WMA1, Washington, USA | oah | Oxaloacetate acetylhydrolase | T-DNA | - | - | + | + | NA | NA | No | Faba bean (Broad Windsor), pea (Guido), green bean (Great North Tara) and soybean (Skylla) | [65] | |
| sscle_10g075560 | SS1G_08218 | 1980, Nebraska, USA | oah1 | Oxaloacetate acetylhydrolase | Deletion | - | + | + | + | + | NA | No | Tomato (Bonnie Best), common bean (Bush Blue Lake 47), soybean (Harosoy), canola, Arabidopsis and sunflower | [66] | |
| sscle_10g075560 | SS1G_08218 | 1980, Nebraska, U.S.A.; UF1, Florida; WMA, Washington, USA | oah1 | Oxaloacetate acetylhydrolase | Deletion by CRISPR | - | + | + | + | + | NA | No | Soybean, canola, tomato (Better Boy), faba bean (Windsor), and pea (Sugar Daddy) | [40] | |
| sscle_10g077630 | SS1G_08489 | 1980, Nebraska, USA | pph1 | Type 2A Ser/Thr phosphatase catalytic subunit PP2Ac | RNAi | + | + | NA | NA | NA | NA | No | [67] | ||
| sscle_10g079320 | SS1G_14065 | 1980, Nebraska, USA | ssp1 | Development-specific protein | Deletion | - | - | NA | NA | NA | NA | No | Resistance to glycoside-containing antibiotics | [68] | |
| sscle_11g082700 | SS1G_07871 | 1980, Nebraska, USA | rgb1 | Type 2A Ser/Thr phosphatase B subunit | RNAi | - | + | NA | + | + | NA | No | Arabidopsis and tomato (Bonny Best) | MAPK pathway | [67] |
| sscle_11g083230 | SS1G_07798 | SUN-F-M, Inner Mongolia, China | gpd | glyceraldehyde-3-phosphate dehydrogenase | RNAi | NA | + | NA | NA | NA | NA | No | [46] | ||
| sscle_11g083680 | SS1G_07749 | 1980, Nebraska, USA | xyl1 | Endo-β-1,4-xylanase | Deletion | + | + | NA | + | NA | NA | Yes | Canola and Arabidopsis | Apothecia formation | [69] |
| sscle_11g083950 | SS1G_07715 | 1980, Nebraska, USA | sac1 | Adenylate cyclase | Deletion | + | + | - | + | NA | NA | No | Tomato (Bonnie Best) | Apothecia production, cAMP-signaling | [70] |
| sscle_12g087830 | SS1G_11172 | 1980, Nebraska, USA | nox2 | NADPH oxidase | RNAi | - | + | - | - | NA | NA | No | Tomato (Rutger) | ROS regulation | [50] |
| sscle_12g090900 | SS1G_11866 | 1980, Nebraska, USA | smk1 | ERK -type MAP kinase | RNAi | + | + | NA | NA | NA | NA | No | pH-dependent regulation | [71] | |
| sscle_14g099710 | SS1G_08814 | 1980, Nebraska, USA | odc1 | Oxalate decarboxylases | Deletion | - | NA | NA | - | NA | NA | Yes | Common bean (Bush Blue Lake 47), soybean (Harosoy) and tomato (Bonnie Best) | [64] | |
| sscle_16g107670 | SS1G_10096 | 1980, Nebraska, USA | cp1 | Cerato-platanin protein | Deletion | - | - | NA | + | NA | + | Yes | Arabidopsis and tobacco | Interaction with PR1 | [72] |
| sscle_16g107670 | SS1G_10096 | FXGD2, Anhui, China | sm1 | Cerato-platanin protein | RNAi | + | + | - | + | NA | NA | Yes | Tobacco, oilseed rape and soybean | [73] | |
| sscle_16g107930 | SS1G_10135 | 1980, Nebraska, USA | fdh1 | Formaldehyde dehydrogenase | Deletion | - | + | NA | + | + | NA | No | Common bean and tomato | Osmotic oxidative stress resistance | [74] |
| sscle_16g109570 | ANQ80447 | 1980, Nebraska, USA | nsd1 | GATA-type Ivb zinc-finger transcription factor | Deletion | - | - | NA | + | + | NA | No | Tomato (Bonnie Best), celery and tomato | Ascogonia formation, apothecium development | [75] |
The assembly genome for the new version gene code is ASM185786v1 published in 2017 and ASM14694v1 for the old version published in 2011. ‘+’ in the table represents that the corresponding phenotype of mutant is altered compared to wild type strain, while ‘-’ means that the phenotype remains unchanged. NA means not assessed. The secretion signals were found using ‘SignalP-5.0’. Unless specified, all deletions were generated with homologous recombination. The Latin names of host species used for pathogenicity test of mutants are: Apium graveolens (celery), Arabidopsis thaliana (Arabidopsis), Brassica napus (canola, oilseed rape, rapeseed), Brassica rapa subsp. chinensis (pak choi cabbage), Capsicum frutescens (hot pepper), Cucumis sativus (cucumber), Fragaria ananassa (strawberry), Glycine max (soybean), Helianthus annuus (sunflower), Lycopersicon solanum (tomato), Nicotiana benthamiana (tobacco), Phaseolus vulgaris (bush bean, common bean, green bean), Pisum sativum (pea), Vicia faba (faba bean). The ecotype of A. thaliana mentioned in the table is Columbia-0. The specific local cultivars of host species are shown in brackets.