Table 1.
Application of CRISPR/Cas9 technology in tomato improvement for plant architecture, flower, fruit and biotic/abiotic stress resistance-related traits.
| Trait | Genotype | Target gene | Gene ID | CRISPR (DNA repair) |
Transformation | Genome editing effects | References |
|---|---|---|---|---|---|---|---|
| Plant architecture, flower and fruit traits | |||||||
| Plant architecture, lycopene, fruit weight, size, number | S. pimpinellifolium |
SP, O, MULT, FAS FW2.2, CNR, CycB |
Solyc06g074350, Solyc02g085500, Solyc02g077390, Solyc11g071380, Solyc02g077920 Solyc04g040190 |
Cas9 (knockout) |
Agrobacterium | Determinate growth habit (SP), increased fruit size (FAS), shape (O), number (MULT), fruit weight (FW2.2), and very high lycopene (CycB) content | Zsögön et al., 2018 |
| Inflorescence, fruit size, weight, yield | M82, S. pimpinellifolium |
WUS/LC, CLV3/FAS SP, S |
Solyc02g083950,Solyc11g071380, Solyc06g074350, Solyc02g077390 |
Cas9 (knockout) |
Agrobacterium | Meristem size regulator (LC-FAS), increased compound inflorescence, locule number, fruit weight, size, and yield | Rodríguez-Leal et al., 2017 |
| Plant growth habit, fruit | S. pimpinellifolium |
SP, SP5G, CLV3, WUS, GGP1 |
Solyc06g074350, Solyc05g053850, Solyc11g071380, Solyc02g083950, Solyc02g091510 |
Cas9 (knockout) |
Agrobacterium | Determinate growth habit, compact plants, synchronous fruit ripening and enlarged fruit size | Li et al., 2018d |
| Plant growth habit, flower | M82 | SP5G | Solyc05g053850 | Cas9 (knockout) |
Agrobacterium | Rapid flowering, compact determinate growth habit and early yield | Soyk et al., 2017 |
| Plant growth habit | Red Setter |
SP, SP5G |
Solyc06g074350, Solyc05g053850 |
Cas9 (Knockin/RNPs) |
Protoplast (PEG) | Protoplast-mediated transformation protocols | Liu et al., 2022 |
| Plant regeneration | S. peruvianum |
SGS3, RDR6, PR-1, ProSys, Mlo1 |
Solyc04g025300, Solyc04g014870,Solyc08g075820, Solyc09g006005, Solyc05g051750, Solyc04g049090 |
Cas9 (Knockin/RNPs) |
Protoplast (PEG) | No chromosomal changes or unintended genome editing sites, and heritable changes | Lin et al., 2022a |
| Plant regeneration | Micro-Tom | AGO7 | Solyc01g010970 | Cas9 (CBEs/ABEs) |
Protoplast (PEG) |
Successful application of cytosine base editors (CBEs) and adenine base editors (ABEs) in rice, tomato and poplar | Sretenovic et al., 2021 |
| Plant phenotype | Micro-Tom |
PDS, PIF4 |
Solyc03g123760, Solyc07g043580 |
Cas9 (knockout) |
Agrobacterium | High frequencies of homozygous and biallelic albino mutants | Pan et al., 2016 |
| Plant phenotype | Ailsa Craig and Micro-Tom |
PDS | NM_001247166 | Cas9 (knockout) |
Agrobacterium | Albino mutants | Li et al., 2018c |
| Plant regeneration | Red Setter, Ailsa Craig, M82, Moneymaker |
SP, SP5G |
Solyc06g074350, Solyc05g053850 |
Cas9 (RNPs) |
Protoplast (PEG-Ca2+) | Improved protoplast isolation and shoot regeneration | Liu et al., 2022 |
| Plant regeneration | Solanum spp. |
SHR, SCR |
Solyc02g092370, Solyc10g074680 | Cas9 (knockout) |
A.
rhizogenes |
Hairy root transformation | Ron et al., 2014 |
| Dwarf plant | Moneymaker | PRO | Solyc11g011260 | Cas9 (knockout) |
Agrobacterium | Gibberellins-responsive dominant dwarf mutant with loss of function and deletion in DELLA allele | Tomlinson et al., 2019 |
| Leaf shape | M82 | AGO7 |
Solyc01g010970, Solyc07g021170, Solyc08g041770, Solyc12g044760 |
Cas9 (knockout) |
Agrobacterium | Change in leaf shape from compound flat to needle like leaves | Brooks et al., 2014 |
| Leaf shape | Micro-Tom |
DELLA, ETR1 |
Solyc11g011260, Solyc12g011330 |
Cas9 (base editing, Target-AID) |
Agrobacterium | Reduced serrated leaflets and insensitivity to ethylene | Shimatani et al., 2017 |
| Male sterility | Micro-Tom |
Ms1035
, GSTAA |
Solyc02g079810, Solyc02g081340 |
Cas9 (knockout) |
Agrobacterium | Male-sterile line with a green hypocotyl or trichome density | Liu et al., 2021a |
| Male sterility | KS-13 | MS10 | Solyc02g079810 | Cas9 (knockout) |
Agrobacterium | Male sterile line | Jung et al., 2020 |
| Male sterility | Alisa Craig | RBOH, RBOHE | Solyc01g099620, Solyc07g042460 | Cas9 (knockout) | Agrobacterium | Male sterile line | Dai et al., 2022 |
| Male sterility | Ailsa Craig | AMS | QDO73362.1 | Cas9 (knockout) | Agrobacterium | Non-viable pollens | Bao et al., 2022 |
| Male sterility | Ailsa Craig | CMT4 | Solyc08g005400 | Cas9 (knockout) | Agrobacterium | Suppressed pollen tube growth | Guo et al., 2022 |
| Jointless fruit | M82 | J2 | Solyc12g038510, | Cas9 (knockout) | Agrobacterium | Jointless inflorescence, large sepals | Roldan et al., 2017 |
| Leaf, flower, fruit shape | M82 |
BOP1/2/3
(TMF) |
Solyc04g040220, Solyc10g079460, Solyc10g079750 |
Cas9 (knockout) | Agrobacterium | Defects in fruit shape and altered leaf | Xu et al., 2016 |
| Fruit size, locule number | S. pimpinellifolium | ENO | Solyc03g117230 | Cas9 (knockout) | Agrobacterium | Increased fruit locule number, fruit size and yield | Yuste-Lisbona et al., 2020 |
| Fruit weight, number | S. pimpinellifolium |
KLUH
(CYP78A family) |
M9 SNP | Cas9 (knock-out) | Agrobacterium | Increased fruit weight and decreased number of small fruits | Li et al., 2022b |
| Fruit size | Ailsa Craig |
SUN, BZR1.7 |
Solyc10g079240, Solyc10g076390 |
Cas9 (knockout) | Agrobacterium | Elongated fruit shape | Yu et al., 2022 |
| Fruit size | S. lycopersicum, S. pimpinellifolium |
TRM3/4/5, OFP |
Solyc07g008670, Solyc10g076180 | Cas9 (knockout) | Agrobacterium | Slightly flatter fruit | Wu et al., 2018 |
| Fruit ripening | Micro-Tom |
MIR164A
(targets: NAM2, NAM3) |
Solyc03g115850, Solyc06g069710 | Cas9 (knockout) | Agrobacterium | Accelerated fruit ripening | Lin et al., 2022b |
| Fruit ripening | Ailsa Craig | RIN | Solyc05g012020 | Cas9 (knockout) | Agrobacterium | Partial ripening and moderate red pigmentation | Ito et al., 2015; Ito et al., 2017; Ito et al., 2020 |
| Fruit ripening | Ailsa Craig |
NOR, CNR |
Solyc10g006880
Solyc02g077920 |
Cas9 (knockout) | Agrobacterium | Delayed ripening by 2-3 days | Gao et al., 2019 |
| Fruit ripening | Moneyberg, Ailsa Craig |
AP2a, NAC-NOR, FUL1, FUL2 |
Solyc03g044300, Solyc10g006880, Solyc06g069430, Solyc03g114830 |
Cas9 (knockout) | Agrobacterium | Early fruit ripening but not fully ripen | Wang et al., 2019a |
| Fruit ripening | Ailsa Craig |
PL, PG2a, TBG4 |
Solyc03g111690, Solyc10g080210, Solyc12g008840, |
Cas9 (knockout) | Agrobacterium | Influenced fruit firmness, weight and color | Wang et al., 2019a |
| Fruit ripening | Ailsa Craig | NAM1 | Solyc06g060230 | Cas9 (knockout) | Agrobacterium | Regulates fruit ripening | Gao et al., 2021 |
| Fruit ripening | Moneyberg, Ailsa Craig |
FUL1, FUL2 |
Solyc06g069430, Solyc03g114830 | Cas9 (knockout) | Agrobacterium | Independent/overlapping functions in fruit ripening | Wang et al., 2019a |
| Fruit ripening | Micro-Tom | ORRM4 | - | Cas9 (knockout) | Agrobacterium | Involved in RNA editing of transcripts | Yang et al., 2017 |
| Parthenocarpy | Micro-Tom, Ailsa Craig | IAA9 | Solyc04g076850 | Cas9 (knockout) | Agrobacterium | Leaf shape changes and parthenocarpic | Ueta et al., 2017 |
| Parthenocarpy | MP-1 | AGL6 | Solyc01g093960 | Cas9 (knockout) | Agrobacterium | Parthenocarpic fruit and heat stress tolerance | Klap et al., 2017 |
| Fruit flavor | F6 (RILs) | FLORAL4 | Solyc04g063350 | Cas9 (knockout) | Agrobacterium | Increased phenylalanine volatile | Tikunov et al., 2020 |
| Lycopene | – | lncRNA1459 |
KT963310, KT963311 |
Cas9 (knockout) | Agrobacterium | Reduced ethylene and lycopene | Li et al., 2018a |
| Lycopene | Ailsa Craig |
SGR1, LCY-E, Blc, LCY-B1, LCY-B2 |
DQ100158, EU533951, XM_010313794, EF650013, AF254793 |
Cas9 (knockout) | Agrobacterium | Increased lycopene | Li et al., 2018b |
| Lycopene | S. pimpinellifolium | CycB | Solyc04g040190 | Cas9 (knockout) | Agrobacterium | High lycopene content | Zsögön et al., 2018 |
| Carotenoid, chlorophyll, B. cinerea | Moneymaker, San Marzano |
GF | Solyc08g080090 | Cas9 (knockout) | Agrobacterium | high carotenoids, chlorophylls and Botrytis cinerea resistance | Gianoglio et al., 2022 |
| Yellow fruit | Red Setter |
Psy1, CrtR-b2 |
Solyc03g031860, Solyc03g007960 |
Cas9 (knockout) | Agrobacterium | Yellow flesh color fruit | D’Ambrosio et al., 2018 |
| Carotenoid | Periyakulam 1 (PKM1) | CRTISO | AF416727 | Cas9 (knockout) | Agrobacterium | Complete knockout of protein function | Jayaraj et al., 2021 |
| Pink fruit | Inbred lines | MYB12 | Solyc01g079620 | Cas9 (knockout) | Agrobacterium | Pink color fruit | Deng et al., 2018 |
| Orange and yellow fruit | Micro-Tom |
CRISTO, PSY1 |
- | Cas9 (knockin/geminiviral replicon) | Agrobacterium | High editing efficiency | Dahan-Meir et al., 2018 |
| Yellow fruit | M82, S. pimpinellifolium | PSY1 | Solyc03g031860 | Cas9 (knockin) |
Agrobacterium | Yellow fruit | Hayut et al., 2017 |
| Carotenoid, lycopene | Micro-Tom |
DDB1, DET1, CYC-B |
Solyc02g021650, Solyc01g056340, Solyc06g074240 |
Cas9 (Target-AID base editing) |
Agrobacterium | Increased carotenoid, lycopene, and β-carotene content | Hunziker et al. (2020) |
| Anthocyanin | Micro-Tom | ANT1 | Solyc10g086260 | Cas9 (Geminivirus replicon) | Agrobacterium | High anthocyanin content | Cˇermák et al., 2015; Vu et al., 2020b |
| Anthocyanin | Indigo Rose |
AN2, ANT1, ANT1- like, R2R3-MYB TF AN2-like/Anthocyanin fruit (Aft) |
Solyc10g086250, Solyc10g086260, Solyc10g086270, Solyc10g086290 |
Cas9 (knockout) | Agrobacterium | Decreased anthocyanin content | Yan et al., 2020; Zhi et al., 2020 |
| Anthocyanin | Indigo Rose | HY5 | Solyc08g061130 | Cas9 (knockout) | Agrobacterium | Decreased anthocyanin | Qiu et al., 2019 |
| GABA | Micro-Tom |
GAD2, GAD3 |
B1Q3F1 FB1Q3F2 |
Cas9 (knockout) | Agrobacterium | Increased GABA content | Nonaka et al., 2017 |
| GABA | Ailsa Craig and Micro-Tom |
GABA-TP1, GABA-TP2, GABA-TP3, CAT9, SSADH |
AY240229, AY240230, AY240231, XM_004248503, NM_001246912 |
Cas9 (knockout) | Agrobacterium | Increased GABA content | Li et al., 2018c |
| Total soluble solid (TSS) | M82 |
INVINH1, VPE5 |
Solyc12g099200, Solyc12g095910 | Cas9 (knockout) | Agrobacterium | Increased glucose, fructose, and TSS contents | Wang et al., 2021a |
| Sugar | Suzukoma | INVINH1 | Solyc12g099200 | Cas9 (knockout, Target-AID base editing) |
Agrobacterium | High sugar content | Kawaguchi et al., 2021 |
| Vitamin D | Moneymaker | 7-DR2 | Solyc06g074090 | Cas9 (knockout) | Agrobacterium | Increased 7-dehydrocholesterol (7-DHC) level | Li et al., 2022a |
| Ascorbate | Micro-Tom | APX4 | Solyc06g005150 | Cas9 (knockout) | Agrobacterium | Increased ascorbate content | Do et al., 2022 |
| Malate | S. lycopersicum, S. pimpinellifolium, S.l. var cerasiforme |
TFM6, ALMT |
Solyc06g072910, Solyc06g072920 | Cas9 (knockout) | Agrobacterium | Reduced malate content | Ye et al., 2017 |
| Shelf-life | M82 | ALC | FJ404469 | Cas9 (knockin) | Agrobacterium | Extended shelf-life | Yu et al., 2017 |
| Fruit softening | Micro-Tom | PG | solyc10g080210 | Cas9 (knockout) | A. rhizogenes | Delayed fruit softening | Nie et al., 2022 |
| Biotic stress resistance | |||||||
| TYLCV | Moneymaker |
CP, Rep
(virus gene) |
- | Cas9 (knockout) | Agrobacterium | Resistance | Tashkandi et al., 2018 |
| TYLCV | Moneymaker | rgsCaM promoter | - | Cas9 (knockout) | Agrobacterium | Efficiency of an inducible promoter | Faal et al., 2020 |
| TYLCV, Powdery mildew | BN-86 |
Pelo, Mlo1 |
Solyc04g009810, Solyc04g049090 |
Cas9 (knockout) | Agrobacterium | High resistance | Pramanik et al., 2021 |
| ToMV | Ailsa Craig | DCL2b | – | Cas9 (knockout) | Agrobacterium | Resistance | Wang et al., 2018 |
| Bacterial speck (Pseudomonas syringae) | Moneymaker | JAZ2 | Solyc12g009220 | Cas9 (knockout) | Agrobacterium | Resistance | Ortigosa et al., 2019 |
| Late blight (Phytophthora infestans) | Alisa Craig | MYBS2 | Solyc04g008870 | Cas9 (knockout) | Agrobacterium | High resistance | Liu et al., 2021b |
| Powdery mildew (Oidium sp.) | Moneymaker | Mlo1 | Solyc04g049090 | Cas9 (knockout) | Agrobacterium | Resistance | Nekrasov et al., 2017 |
| Powdery mildew | Moneymaker | PMR4 | Solyc07g053980 | Cas9 (knockout) | Agrobacterium | Reduced resistance | Martínez et al., 2020 |
| Downy mildew | FL8000 | DMR6-1, DMR6-2 |
Solyc03g080190, Solyc06g073080 |
Cas9 (knockout) | Agrobacterium | High resistance | Thomazella et al., 2021 |
| Multiple diseases | FL8000 | DMR6-1 | Solyc03g080190 | Cas9 (knockout) | Agrobacterium | Disease resistance to P. syringae, P. capsici, and Xanthomonas spp. | Thomazella et al., 2016 |
| Fusarium wilt | 76R, rmc | - | Solyc08g075770 | Cas9 (knockout) | Agrobacterium | Resistance | Prihatna et al., 2018 |
| Gray mold (Botrytis cinerea) |
Micro-Tom, Ailsa Craig | MYC2, MAPK3 | - | Cas9 (knockout) | Agrobacterium | Decreased disease resistance to B. cinerea | Zhang et al., 2018; Shu et al., 2020 |
| PVY, CMV | Inbred line S8 |
eIF4E1
(host gene) |
– | Cas9 (knockout) | Agrobacterium | PVY and CMV resistant mutants | Atarashi et al., 2020 |
| Abiotic stress tolerance | |||||||
| Multiple stresses | Rubion | GRXS14/15/16/17 |
Solyc02g082200, Solyc06g067960, Solyc09g005620, Solyc02g078360 |
Cas9 (knockout) | Agrobacterium | Increased tolerance to heat, chilling, drought, heavy metal toxicity and nutrient deficiency | Kakeshpour et al., 2021 |
| Drought | Ailsa Craig | NPR1 | KX198701 | Cas9 (knockout) | Agrobacterium | Reduced drought tolerance | Li et al., 2019 |
| Drought | Condine Red |
ALD1, FMO1 |
Solyc11g044840, Solyc07g04243 |
Cas9 (knockout) | Agrobacterium | Increased drought tolerance | Wang et al., 2021b |
| Drought | Micro-Tom | LBD40 | Solyc02g085910 | Cas9 (knockout) | Agrobacterium | Increased drought tolerance | Liu et al., 2020a |
| Drought | Ailsa Craig | MAPK3 | AY261514 | Cas9 (knockout) | Agrobacterium | Reduced drought tolerance | Wang et al., 2017 |
| Heat | Condine Red | BZR1 | Solyc04g079980 | Cas9 (knockout) | Agrobacterium | Reduced heat tolerance | Yin et al., 2018 |
| Heat | Condine Red |
CPK28, APX2 |
Solyc02g083850, Solyc06g005150 | Cas9 (knockout) | Agrobacterium | Improved thermotolerance | Hu et al., 2021 |
| Heat | MP-1 | AGL6 | Solyc01g093960 | Cas9 (knockout) | Agrobacterium | Seedless fruit under heat stress | Klap et al., 2017 |
| Cold | Ailsa Craig | CBF1 | AAS77820 | Cas9 (knockout) | Agrobacterium | Reduced chilling tolerance | Li et al., 2018e |
| Salinity | S. lycopersicum, S. pimpinellifolium | HAK20, SOS1 | Solyc04g008450, Solyc11g044540 | Cas9 (knockout) | Agrobacterium | Increased salt sensitivity | Wang et al., 2020; Wang et al., 2021c |
| Salinity | Hongkwang |
HKT1;2
RAD51/54 |
Solyc07g017540, Solyc04g056400 |
Cas12a/ LbCpf1 (Geminivirus replicon) |
Agrobacterium | Increased salt tolerance | Vu et al., 2020b |
| Salinity | Hongkwang | HyPRP1 | Solyc12g009650 | Cas9 (knockout) | Agrobacterium | Increased salt tolerance | Tran et al., 2021 |
| C-N metabolism | Micro-Tom | SBPase | Solyc05g052600 | Cas9 (knockout) | Agrobacterium | Optimal growth, carbon assimilation and nitrogen metabolism | Ding et al., 2018 |
| Phosphate transporter | Micro-Tom |
PHO1;1, PHO1;2, PHO1;3, PHO1;4, PHO1;5, PHO1;6 |
Solyc09g090360
Solyc08g068240, Solyc05g010060, Solyc05g013180, Solyc02g088220, Solyc02g088230 |
Cas9 (knockout) | Agrobacterium | High anthocyanin and phosphate content | Zhao et al., 2019 |
| Herbicide resistance | WVA106 | ALS | Solyc03g044330 | Cas9 (base editing) |
Agrobacterium | Resistance (Chlorsulfuron) | Veillet et al., 2019 |
| Herbicide resistance | Micro-Tom |
PDS, ALS, EPSPS |
Solyc03g123760, Solyc06g059880, Solyc03g044330, Solyc01g091190 |
Cas9 (knockout) | Agrobacterium | Resistance | Yang et al., 2022 |
| Herbicide resistance | – |
ALS1, ALS2, ALS3 |
Solyc03g044330, Solyc07g061940, Solyc06g059880 |
Cas9 (knockin) | Agrobacterium | Resistance (Chlorsulfuron) | Danilo et al., 2019 |
7-DR2 (7-Dehydrocholesterol reductase), AGL6 (Agamous-like 6), AGO7 (Argonaute7), ALC (Alcobaca), ALD1 (AGD2-like defense response protein), ALMT (Al-activated malate transporter),ALS1 (Acetolactate synthase 1), AMS (Aborted microspores), AN2 (Anthocyanin2), ANT1 (Anthocyanin 1), AP2a (Apetala2a TF), APX2/4 (Ascorbate peroxidase2/4), Blc (Beta-lycopene cyclase),BOPs (Blade-on-petiole), BZR1 (Brassinazole-resistant 1), CRTISO (Carotenoid isomerise), CAT9 (Cationic amino acid transporter 9), CBF1 (C-repeat/dehydration responsive element binding factor1), CLV3 (Clavata3), CMT4 (Chromomethylase), CNR (SBP-box colorless non-ripening), CPK28 (Calcium-dependent protein kinase28), CP (Coat protein), CrtR-b2 (Beta-carotene hydroxylase 2),CRTISO (Central role of carotenoid isomerase), CYC-B/CycB (Lycopene beta cyclase), DDB1 (DNA damage UV binding protein 1), DELLA (Aspartic acid–glutamic acid–leucine–leucine–alanine),DCL2b (Dicer-like 2b), DET1 (Deetiolated1), DMR6 (Downy mildew resistance 6), EJ2 (Enhancer-of-jointless2), ENO (Excessive number of floral organs), EPSPS (5-Enolpyruvylshikimate-3-phosphate synthase), ETR1 (Ethylene receptor 1), FASCIATED (FAS), FMO1 (Flavin-dependent monooxygenase), FUL1/2 (Fruitfull), GABA-TP1 (Pyruvate-dependent g-aminobutyric acidtransaminase 1), GAD2 (Glutamate decarboxylase 2), GAD3 (Glutamate decarboxylase 3), GF (Greenflesh/Staygreen), GGP1 (GDP-l-galactose phosphorylase1), GRXS (CGFS-type glutaredoxin),GSTAA (Glutathione S-transferase), HAK20 (High-affinity K+ 20), HKT1;2 (High-affinity potassium transporter 1;2), HY5 (elongated hypocotyl5), HyPRP1 (Hybrid proline-rich protein 1), IAA9(Auxin-induced 9), INVINH1 (Invertase inhibitor 1), J2 (Jointless-2), JAZ2 (Jasmonate zim domain), LBD40 (Lateral organ boundaries domain40), LOCULE NUMBER (LC), LCY-B1 (Lycopene bcyclase1), LCY-B2 (Lycopene b-cyclase 2), LCY-E (Lycopene e-cyclase), LIN (Long inflorescence), MAPK3 (Mitogen activated protein kinase 3), MIR164A (MicroRNA164A), TFAM1/TFAM2(Mitochondrial transcription factor A), Mlo1 (Mildew resistance locus o 1), MS10 (Male sterile 10), Ms1035 (Male sterile 1035), MULT (Multiflora), MYBS2 (MYB transcription factor S2), MYC2(Basic helix–loop–helix transcription factor), NAC (NAM-ATAF-CUC), NAM1/2/3 (No apical meristem1/2/3), NAC-NOR (NAC TF non-ripening), NOR (Non-ripening), NOR-like1 (Non-ripeninglike1), NPR1 (Nonexpressor of pathogenesis-related gene 1), O (ovate), ORRM4 (organelle RNA recognition motif-containing protein4), PDS (Phytoene desaturase), OFP (OVATE family protein),PG (Polygalacturonase), PG2a (polygalacturonase 2a), PHO1 (Phosphate 1), PIF4 (Phytochrome interacting factor 4), PL (pectate lyase), PMR4 (Powdery mildew resistance 4), PR-1 (Pathogenesisrelatedprotein-1), PRO (Procera), ProSys (Prosystemin), PSY1 (Phytoene synthase 1), RAD51/54 (DNA repair and recombination protein51/54), RBOH/RBOHE (Respiratory burst oxidasehomolog), RDR6 (RNA ald1dependent RNA polymerase 6), REP (Replicase), RIN (Ripening inhibitor), RMC (Reduced mycorrhizal colonization), SBPase (Sedoheptulose-1,7-bisphosphatase), S(Compound inflorescence), SCR (Scarecrow), SHR (Shortroot), SGR1 (Stay-green 1), SGS3 (Suppressor of gene silencing 3), SOS1 (Salt overly sensitive 1), SP (Self pruning), SP5G (Self pruning 5G),SSADH (Succinate semialdehyde dehydrogenase), Target-AID (Target activation-induced cytidine deaminase), TBG4 (b-galactanase), TFM6 (Tomato fruit malate on chromosome6), TMF(Terminating flower), ToMV (Tomato Mosaic virus), TRM3/4/5 (TONNEAU1 Recruiting Motif3/4/5), VPE5 (Vacuolar processing enzyme5), and WUS (Wuschel).RNPs (ribonucleoproteins), PEG (polyethylene glycol), Target-AID (Target Activation Induced Cytidine Deaminase), NHEJ (Homologous-End-Joining), KO (gene knock-out), KI (gene knock-in), HDR (homology-directed repair), HR (homologous recombination), HKI (HR-based KI).