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Table 1 Improved agricultural traits in transgenic bananas (Musa spp.)

From: Recent advances and future directions in banana molecular biology and breeding

Trait

Banana genotype

Transformation method

Promoter

Transformed gene

Potential molecular mechanism

Efficiency

References

Foc TR4 resistance

Cavendish cv. Grand Nain (AAA)

Agrobacterium + Banana ECS

pNos, pZmUbi

RGA2 from a Foc TR4-resistant wild banana, Ced-9 from nematode

Increasing resistance to Foc TR4 possibly through an R-gene-like cascade pathway (RGA2 lines), and by preventing fungus-induced cell death and maintaining organelle homeostasis (Ced9 lines)

Two transgenic lines (RGA2-3 and Ced9-21) appeared to be immune to Foc TR4 in an infected field trial over a 3-year period

(Dale, et al. 2017)

Foc TR4 resistance

Cavendish cv. Grand Nain (AAA)

Agrobacterium + Banana ECS

pZmUbi

MpbHLH from cold-tolerant Dajiao banana

Strengthen banana cell wall and/or scavenge ROS

Disease index of 2 transgenic plants was significantly lower than control after 2-week inoculation in pots under normal management

(Li, et al. 2022b)

Foc TR4 resistance

Cavendish cv. Grand Nain (AAA)

Agrobacterium + Banana ECS

pZmUbi

Foc TR4 ERG6 double strand RNAs, Foc TR4 ERG11 double strand RNAs

Induced Foc ERG6 and Foc ERG11 genes silencing in banana, inhibited fungal ergosterol synthesis and conidial germination

Only 30% of ERG6-RNAi or 15% of ERG11-RNAi plants were sensitive to Foc TR4, but more than 85% of WT showed apparent Fusarium wilt symptoms in a heavily infected field after 2 years

(Dou, et al. 2020)

Foc TR4 resistance

Furenzhi (AAcv)

Agrobacterium + Banana ECS

pCaMV35S

ThChit42 from Trichoderma harzianum

Antifungal activities by cleaving chitin in the fungal cell wall

A transgenic line T3 showed no disease symptoms and remained healthy after 2 months inoculation in pots

(Hu, et al. 2013)

Foc TR4 resistance

Taijiao (AAA)

Agrobacterium + Particle bombardment + Banana Apical meristem

pCaMV35S

Human lysozyme

Antifungal activity by cleaving chitin in the fungal cell wall

Two transgenic lines H-67 and H-144 remained healthy and were able to fruit in the field

(Pei, et al. 2005)

Foc TR4 resistance

Pisang Nangka (AAB)

Particle bombardment + Banana Single cauliflower-like bodies

pCaMV35S

OsTLP from rice

Antifungal activities by alternating fungus cell membrane integrity leading to inhibition of fungal growth, spore lysis, reduction in spore number, or reduced viability of germinated spores

The average percentage of disease incidence in transgenic plants was 29.4% compared to the control at 89.1% after 4 weeks inoculation in pots

(Mahdavi, et al. 2012)

Foc TR4 resistance

Cavendish cv. Williams (AAA)

Agrobacterium + Banana ECS

pCaMV35S

MaLYK1 from Williams banana

Resistance against Foc TR4 by mediating MAMP-induced ROS generation and defense gene activation

No obvious lesions observed in inoculated MaLYK1-OE lines compared with WT and MaLYK1-RNAi lines in pots

(Zhang, et al. 2019)

Foc Race1 resistance

Lady Finger (AAB)

Agrobacterium + Banana ECS

pZmUbi

Bcl-xL, Ced-9, Bcl-2 3’ UTR

Bcl-xL and ⁄or Ced-9 may prevent cell death and enhance plant resistance characteristics by contributing to the maintenance of organelle homeostasis; Bcl-2 3’ UTR confers resistance to plant cells is unknown

The transgenic line Bcl-2 3’ UTR-6 showed a level of Foc Race1 resistance similar to resistant wild-type ‘Grand Naine’ at least 3 months after inoculation in small-plant bioassays

(Paul, et al. 2011)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi

MusaDAD1, MusaBAG1 and MusaBI1 from Rasthali banana

MusaBAG1 gene plays a far greater role in the control of PCD in banana plants compared to the other two genes studied

MusaBAG1 overexpressing plants demonstrated the best resistance towards Foc Race1 infection in the three groups of transgenic plants derived from the three gene constructs after 6 weeks in greenhouse bioassays

(Ghag, et al. 2014a)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi

PhDef1 and PhDef2 from Petunia

Interact with specific lipids on the fungal membrane and subsequently permeabilize them to inhibit fungus growth

Four transgenic plants displayed a high degree of resistance to Foc Race1 challenge after 3 months inoculation in pots

(Ghag, et al. 2012)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pCaMV35S

Ace-AMP1 from onion

The activity of this AMP is mainly on the structural components of the cell wall attacking multiple targets

Six transgenic plants root challenged with Foc Race1 showed VDIs (vascular disease index) ranging from 38 to 48% compared to the control at 100%, after 6 months inoculation in pots

(Mohandas, et al. 2013)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pCaMV35S, pZmUbi

Ace-AMP1 from onion and PFLP from sweet pepper

Higher tolerance to oxidative stress caused by Foc Race1 infection

Two transgenic plants root challenged with Foc Race1 showed VDIs ranging from 10 to 20% compared to the control at 96%, after 6 months inoculation in pots

(Sunisha, et al. 2020a)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi

Sm-AMP-D1 from Stellaria media

Antifungal activity by destabilizing the microbial membranes

Two transgenic lines, Sm-D1 and Sm-D2, were without external wilt symptoms after 6 months inoculation in pots

(Ghag, et al. 2014c)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi

Foc Race1 VEL intron hairpin RNAs, Foc Race1 FTF1 intron hairpin RNAs

Induced Foc VEL and Foc FTF1 genes silencing in banana, inhibited fungal growth, development and pathogenesis

Disease severity scale of 7 ihpRNA-VEL and 5 ihpRNA-VEL transgenic plants was less than 1 (1 = no symptoms) in a 6-week-long bioassay in pots

(Ghag, et al. 2014b)

Foc Race1 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana Single buds

pCaMV35S

GmEg from soybean

Antifungal activity by cleaving β-glucan, a component of mycelial cell walls

Performance of the two transgenic lines was better than the control after 4 weeks inoculation in pots

(Maziah, et al. 2007)

Foc Race2 resistance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pAtUbq3

MSI99 (a magainin analogue gene)

Interacted with the microbial membrane, resulting in loss of essential metabolites and membrane depolarization and uncoupling of respiration in bacteria

Five plants expressing the peptide in the cytoplasm and 11 transgenic plants expressing the peptide in the intercellular spaces were resistant to Foc Race2 in pots

(Chakrabarti, et al. 2003)

Banana Xanthomonas wilt

Sukali Ndiizi and Nakinyika bananas

Agrobacterium + Banana ECS

pCaMV35S

Plant ferredoxin-like protein (Pflp) from sweet pepper

Disease resistance through formation of hypersensitivity response-like necrosis

All the transgenic lines except one showed absolute resistance to BXW after 60 days inoculation in pots

(Namukwaya, et al. 2012)

Banana Xanthomonas wilt

Sukali Ndiizi and Mpologoma bananas

Agrobacterium + Banana ECS

pCaMV35S

Hypersensitivity response-assisting protein (Hrap) gene from sweet pepper

Disease resistance resulting from enhanced hypersensitive cell death

Six transgenic lines showed absolute resistance to BXW after 60 days inoculation in pots

(Tripathi, et al. 2010)

Banana Xanthomonas wilt

Gonja manjaya banana

Agrobacterium + Banana ECS

pCaMV35S

Pflp and Hrap from sweet pepper

Stacking might provide of durable resistance to BXW

Seven transgenic lines with stacked genes showed complete resistance to BXW after 60 days inoculation in pots

(Muwonge, et al. 2016)

Banana Xanthomonas wilt

Dwarf Cavendish (AAA)

Agrobacterium + Banana ECS

pCaMV35S

ELONGATION FACTOR-TU RECEPTOR (AtEFR) gene from Arabidopsis

Enhanced resistance through activation of early immune outputs (e.g., ROS, defense gene expression) mediated by recognition of Xcm EF-Tu by EFR

Eighteen transgenic lines exhibited partial resistance (50–75%) compared to control after 60 days inoculation in pots

(Adero, et al. 2023a)

Banana Xanthomonas wilt

Sukali Ndiizi banana

Agrobacterium + Banana ECS

CRISPR/Cas9-mediated editing

Downy mildew resistance 6 (DMR6) gene from banana

Enhanced resistance through knockout of a susceptibility gene during pathogen infection

One inoculated plant of D15 was without BXW symptoms after 60 days inoculation in pots

(Tripathi, et al. 2021)

Banana Xanthomonas wilt

Gonja manjaya banana

Agrobacterium + Banana ECS

pZmUbi

Xa21 pattern-recognition receptor from wild rice

Xa21 receptor may recognize a microbial determinant that is conserved in Xcm

Twelve inoculated plants were without BXW symptoms after 60 days inoculation in pots

(Tripathi, et al. 2017)

Shelf life

Cavendish cv. Brazilian (AAA)

Agrobacterium + Banana ECS

CRISPR/Cas9-mediated editing

MaACO1 from banana

Promotes the shelf life of banana fruit by inhibiting ethylene biosynthesis

MaACO1-disrupted fruit remained yellow or green 60 days postinoculation vs. WT fruit with brown spots at day 21

(Hu, et al. 2021)

Cold tolerance

Dajiao banana (ABB) and Cavendish cv. Grand Nain (AAA)

Agrobacterium + Banana ECS

pZmUbi

MAPK3 (same as MAPK5 in this review) and ICE1 from cold-tolerant Dajiao banana

MaMAPK5-MaICE1-MaPOD P7 pathway, a positive regulator of cold tolerance in banana

Cold tolerance of MAPK3-RNAi plants decreased at 10 ℃ for 7 days in pots in an ambient environment. Cold tolerance of MaICE1-overexpressing plants increased at 10 ℃ for 2 days followed by recovery for 3 days

(Gao, et al. 2021)

Cold tolerance

Cavendish cv. Brazilian (AAA)

Agrobacterium + Banana ECS

pZmUbi

MYBS3 from cold-tolerant Dajiao banana

MpMYBS3-overexpressing lines had a higher proline content, accumulated less malondialdehyde and displayed lower levels of electrolyte leakage

Cold tolerance of MpMYBS-overexpressing plants increased at 10 °C for 2 days followed by recovery for 3 days

(Dou, et al. 2016)

Drought and salinity tolerance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi

DHN-1 from banana

Increased the protective antioxidative capacity under drought- and salt-stress conditions, reducing free radical-induced damage to the cellular membranes of transgenic plants

Transgenic lines responded significantly better after the initiation of drought or salt stress

(Shekhawat, et al. 2011)

Drought and salinity tolerance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi

SAP1 from banana

Improves capacity to scavenge free radicals under drought/salt stress conditions, reducing free radical-induced damage to the cellular membranes of transgenic plants

Small, uniform, in vitro transgenic shoots overexpressing MusaSAP1 tolerated simulated drought and salt stress (after 10 days in rooting medium supplemented with 100 mM mannitol or 100 mM NaCl) better than the controls

(Sreedharan, et al. 2012)

Drought and salinity tolerance

Silk cv. Rasthali (AAB) and Cavendish cv. Grand Nain (AAA)

Agrobacterium + Banana ECS

pZmUbi

NAC29-like from banana

Increases JA and SA content, further elevating the antioxidant capacity of transgenic plants

Transgenic cv. Rasthali and cv. Grand Naine overexpressing MusaNAC29-like had superior tolerance of drought and salinity stress, but overexpression retarded growth and yield of the transgenic banana fruits

(Negi, et al. 2023)

Cold, drought and salinity tolerance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi, pMusaDHN-1

PIP1;2 from banana

Lower malondialdehyde levels, elevated proline and relative water content, and higher photosynthetic efficiency in transgenic lines under different abiotic stress conditions

Transgenic banana plants overexpressing MusaPIP1;2 had better abiotic stress survival characteristics

(Sreedharan, et al. 2013)

Salinity tolerance

Silk cv. Rasthali (AAB)

Agrobacterium + Banana ECS

pZmUbi, pMusaDHN-1

PIP2;6 from banana

Better photosynthetic efficiency and less membrane damage in transgenic lines under salt stress conditions

Transgenic banana plants overexpressing MusaPIP2;6 used constitutive or inducible promoter led to higher salt tolerance

(Sreedharan, et al. 2015)

Drought, cold and salinity tolerance

Mas cv. Gongjiao (AA)

Particle bombardment + Agrobacterium + buds of immature banana male flower

pCaMV35S

PIP2;7 from banana

Conferred tolerance by maintaining an osmotic balance, reducing membrane injury, and improving ABA levels

Transgenic banana plants overexpressing MusaPIP2;7 improved tolerance to multiple stresse, including drought, cold, and salt

(Xu, et al. 2020)

Cold and drought tolerance

Mas cv. Gongjiao (AA)

Particle bombardment + Agrobacterium + buds of immature banana male flower

pCaMV35S

DREB1F from banana

Conferred tolerance by common modulation of the protectant metabolite levels of soluble sugar and proline, activating the antioxidant system, and promoting jasmonate and ethylene syntheses

Transgenic banana plants overexpressing MaDREB1F increased banana resistance to cold and drought stress

(Xu, et al. 2023)