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====== Plant Resistance Genes in Banana (//Musa balbisiana// and Pisang Awak cultivar) against //Xanthomonas// Infection ======

Author: Rita Fernandes\\
Internal reviewer: [[https://www.researchgate.net/profile/Noemi_Casarin2|Noemi Casarin]]\\

===== Pathogen: //Xanthomonas vasicola// pv. //musacearum// (//Xvm//) =====

==== Resistance gene: germin-like protein 8-14 gene ====

=== Synonyms ===

GER1, GLP1, Ma02_g20530.1, [[https://banana-genome-hub.southgreen.fr/jbrowse_ma2/?loc=chr09:10823178..10825807|Ma09_p15510.1]], LOC103998053, LOC103976190.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.
=== Status (identified, mapped, cloned, sequenced) ===

Identified (Tripathi //et al//., 2019), mapped on chromosome 2 and 9, sequenced (D'Hont //et al//., 2012; Hubert //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Stress associated gene, similar to germin 3 of //Arabidopsis thaliana// (Tripathi //et al//., 2019), codes for germin-like proteins with functions such as [[https://www.ebi.ac.uk/QuickGO/term/GO:0030145|manganese ion binding]] (GO:0030145), [[https://www.ebi.ac.uk/QuickGO/term/GO:0045735|nutrient reservoir activity]] (GO:0045735), metal ion binding (GO:0046872) and which play role in oxidation-reduction process. Upregulated in //Musa balbisiana// and cultivar Pisang Awak after both 12 hpi (hours post-inoculation) with //Xvm// (Tripathi //et al//., 2019).

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==== Resistance gene: //RPM1// gene ====

=== Synonyms ===

Putative disease resistance RPM1, Ma09_g28690, GSMUA_Achr9G28020_001, LOC103999252, XM_009420945.2.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.

=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 9, sequenced (D'Hont //et al//., 2012).

=== Molecular markers ===

//NA//

=== Brief description ===

Gene involved in plant-pathogen interaction, coding for disease resistance protein RPM1, which is an ADP binding (GO:0043531) and which confers resistance to //Pseudomonas syringae// pv. //maculicola// 1. Activated in //Musa balbisiana// as early response to //Xvm// at 12 hpi (hours post-inoculation) (Tripathi //et al//., 2019). Organism-specific biosystem.

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==== Resistance gene: BAG family molecular chaperone regulator 6-like ====

=== Synonyms ===

Ma10_g14770, XM_009422955.2, LOC104000818, GSMUA_Achr10G14650_001.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.

=== Status (identified, mapped, cloned, sequenced) ===

Identified (Tripathi //et al//., 2019), mapped on chromosome 2 and 9, sequenced (D'Hont //et al//., 2012; Hubert //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Protein binding (GO:0005515), chaperone binding (GO:0051087), apoptosis regulator. Activated in //Musa balbisiana// as early response to //Xvm// at 12 hpi (hours post-inoculation) (Tripathi //et al//., 2019).

----

==== Resistance gene: MYB transcription factor gene ====

=== Synonyms ===

Ma05_g25630, GSMUA_Achr5G23240_001, LOC103985880, XM_018826157.1.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.

=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 5, sequenced (Medina-Suárez //et al//., 1997; D'Hont //et al//., 2012; Hubert //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Gene coding for MYB transcription factor, involved in DNA binding (GO:0003677), activated in Pisang Awak at 12 hpi (hours post-inoculation) with //Xvm// (Tripathi //et al//., 2019).

----

==== Resistance gene: Ninja-family protein AFP3-like gene ====

=== Synonyms ===

Ma04_t39350.1, GSMUA_Achr5G23240_001, LOC103982144, XM_009398965.2.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.

=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 4, sequenced (D'Hont //et al//., 2012).

=== Molecular markers ===

//NA//

=== Brief description ===

Coding for Ninja-family protein AFP3-like. Signal transduction (GO:0007165), role in Abscisic acid (ABA) metabolism, activated in //Musa balbisiana// at both 12 and 48 hpi (hours post-inoculation) with //Xvm// (Tripathi //et al//., 2019).

----

==== Resistance gene: lipase-like //PAD4// ====

=== Synonyms ===

Ma06_t14360.1 (Tripathi //et al//., 2019), LOC103987676, XM_009406048.2, GSMUA_Achr6G12900_001.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.

=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 6, sequenced (Medina-Suárez //et al//., 1997; D'Hont //et al//., 2012; Hubert //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Gene coding for lipase-like PAD4 protein, which is involved in SA signalling, in hydrolase activity (GO:0016787) and in lipid metabolic process (GO:0006629). R-gene-mediated and basal plant disease resistance. Activated in //Musa balbisiana// 48 hpi (hours post-inoculation) with //Xvm// (Tripathi //et al//., 2019).

----

==== Resistance gene: putative 26.5 kDa heat shock ====

=== Synonyms ===

GSMUA_Achr9G28520_001, LOC103999445, Ma09_t29150.1.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.
=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 6 (D'Hont //et al//., 2012; Hubert //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Gene coding for 26.5 kDa heat shock protein, mitochondrial-like. Protein processing in endoplasmic reticulum, conserved biosystem, response to heat (GO:0009408), response to high light intensity (GO:0009644), response to hydrogen peroxide (GO:0042542). Activated in //Musa balbisiana// 12 hpi (hours post-inoculation) with //Xvm// (Tripathi //et al//., 2019).

----

==== Resistance gene: vicilin-like seed storage At2g18540 ====

=== Synonyms ===

Ma03_t31180.1, LOC103980070, GSMUA_Achr3G29960_001.

=== Source ===

//Musa alcuminata// subsp. //malaccensis//.

=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 3, sequenced (D'Hont //et al//., 2012; Hubert //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Gene coding for signal peptide vicilin-like seed storage protein At2g18540, plant-derived inhibitors known for having antibacterial and antifungal activity, nutrient reservoir activity (GO:0045735). Activated in //Musa balbisiana// 12 hpi (hours post-inoculation) with //Xvm// (Tripathi //et al//., 2019).

----

==== Resistance gene: //hrap// ====

=== Synonyms ===

Extracellular ribonuclease LE-like, AF168415.1, LOC107877517.

=== Source ===

Transformed into banana cultivars ‘Sukali Ndiizi’ and ‘Nakinyika’, but isolated from sweet pepper (//Capsicum annuum//) (Tripathi //et al//., 2019).

=== Status (identified, mapped, cloned, sequenced) ===

Identified, mapped on chromosome 7 (of sweet pepper) and sequenced (Chen //et al//., 2000; Qin //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Extracellular ribonuclease LE-like, encodes for hypersensitive response assisting protein, ribonuclease T2 activity, RNA binding. Role in intensifying the hypersensitive response mediated by harpinPSS (harpin derived from //Pseudomonas syringae// pv. //syringae//) by dissociating multimeric forms of the hairpin into dimers and monomers that trigger stronger hypersensitive cell death necrosis (Tripathi //et al//., 2014).

----

==== Resistance gene: //Pflp// ====

=== Synonyms ===

ap1, LOC107839912, AF039662.1.

=== Source ===

Transformed into banana cultivars ‘Sukali Ndiizi’ and ‘Nakinyika’, but isolated from sweet pepper (//Capsicum annuum//) (Namukwaya //et al//., 2012).

=== Status (identified, mapped, cloned, sequenced) ===

Mapped on chromosome 8 of sweet pepper (//Capsicum annuum//), sequenced (Dayakar //et al//., 2003; Qin //et al//., 2014).

=== Molecular markers ===

//NA//

=== Brief description ===

Encodes for ferredoxin-like protein (member of photosynthesis proteins), chloroplastic. Known resistance against various bacterial pathogens like //Erwinia//, //Pseudomonas//, //Ralstonia//, and //Xanthomonas// spp. in other plants. Over-expression of this gene leads to intensified production of active oxygen species (AOS) and activation of the hypersensitive response (HR) when plants are challenged with bacterial pathogens (Namukwaya //et al//., 2012).

----

==== Resistance gene: //MusaPUB22/23// ====

=== Synonyms ===

//NA//

=== Source ===

Susceptibility genes in the BXW-susceptible 'Sukali Ndiizi' cultivar (Tripathi //et al//., 2025).

=== Status (identified, mapped, cloned, sequenced) ===

Identified and sequenced (Tripathi //et al//., 2025).

=== Molecular markers ===

Edited genes (Tripathi //et al//., 2025)

=== Brief description ===

Genes //MusaPUB22// and //MusaPUB23// were identified through prior comparative transcriptomics analysis, which revealed their upregulation in susceptible versus resistant genotypes during early infection with //Xvm//. Several gene edited lines displayed complete resistance to //Xvm// infection, while others showed partial resistance, in contrast to the full susceptibility of wild-type controls. Resistance correlated with hydrogen peroxide accumulation and activation of key defense-related genes, indicating enhanced immune responses (Tripathi //et al//., 2025).

----

===== References =====

Chen CH, Lin HJ, Ger MJ, Chow D, Feng TY (2000). cDNA cloning and characterization of a plant protein that may be associated with the harpinPSS-mediated hypersensitive response. Plant Mol. Biol. 43: 429-438. DOI: [[https://doi.org/10.1023/a:1006448611432|10.1023/a:1006448611432]]

D’Hont A, Denoeud F, Aury JM, Baurens FC, Carreel F, Garsmeur O, Noel B, Bocs S, Droc G, Rouard M (2012). The banana (//Musa acuminata//) genome and the evolution of monocotyledonous plants. Nature 488: 213. DOI: [[https://doi.org/10.1038/nature11241|10.1038/nature11241]]

Dayakar BV, Lin HJ, Chen CH, Ger MJ, Lee BH, Pai CH, Chow D, Huang HE, Hwang SY, Chung MC (2003). Ferredoxin from sweet pepper (//Capsicum annuum// L.) intensifying harpin pss-mediated hypersensitive response shows an enhanced production of active oxygen species (AOS). Plant Mol. Biol. 51: 913-924. DOI: [[https://doi.org/10.1023/a:1023061303755|10.1023/a:1023061303755]]

Hubert O, Piral G, Galas C, Baurens F-C, Mbéguié-A-Mbéguié D (2014). Changes in ethylene signaling and MADS box gene expression are associated with banana finger drop. Plant Sci. 223: 99-108. DOI: [[https://doi.org/10.1016/j.plantsci.2014.03.008|10.1016/j.plantsci.2014.03.008]]

Medina-Suárez R, Manning K, Fletcher J, Aked J, Bird CR, Seymour GB (1997). Gene expression in the pulp of ripening bananas (two-dimensional sodium dodecyl sulfate-polyacrylamide gel electrophoresis of //in vitro// translation products and cDNA cloning of 25 different ripening-related mRNAs). Plant Physiol. 115: 453-461. DOI: [[https://doi.org/10.1104/pp.115.2.453|10.1104/pp.115.2.453]]

Namukwaya B, Tripathi L, Tripathi J, Arinaitwe G, Mukasa S, Tushemereirwe W (2012). Transgenic banana expressing //Pflp// gene confers enhanced resistance to //Xanthomonas// wilt disease. Transgenic Res. 21: 855-865. DOI: [[https://doi.org/10.1007/s11248-011-9574-y|10.1007/s11248-011-9574-y]]

Qin C, Yu C, Shen Y, Fang X, Chen L, Min J, Cheng J, Zhao S, Xu M, Luo Y (2014). Whole-genome sequencing of cultivated and wild peppers provides insights into //Capsicum// domestication and specialization. Proc. Natl. Acad. Sci. USA 111: 5135-5140. DOI: [[https://doi.org/10.1073/pnas.1400975111|10.1073/pnas.1400975111]]

Tripathi L, Tripathi JN, Kiggundu A, Korie S, Shotkoski F, Tushemereirwe WK (2014). Field trial of //Xanthomonas// wilt disease-resistant bananas in East Africa. Nat. Biotech. 32: 868-870. DOI: [[https://doi.org/10.1038/nbt.3007|10.1038/nbt.3007]]

Tripathi L, Tripathi JN, Shah T, Muiruri KS, Katari M (2019). Molecular basis of disease resistance in banana progenitor //Musa balbisiana// against //Xanthomonas campestris// pv. //musacearum//. Sci. Rep. 9: 7007. DOI: [[https://doi.org/10.1038/s41598-019-43421-1|10.1038/s41598-019-43421-1]]

Tripathi L, Ntui VO, Muiruri S, Shah T, Tripathi JN (2025). Loss of function of //MusaPUB// genes in banana can provide enhanced resistance to bacterial wilt disease. Commun. Biol. 8: 708. DOI: [[https://doi.org/10.1038/s42003-025-08093-w|10.1038/s42003-025-08093-w]]

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===== Further reading =====

Biruma M, Pillay M, Tripathi L, Blomme G, Abele S, Mwangi M, Bandyopadhyay R, Muchunguzi P, Kassim S, Nyine M, Turyagyenda L, Eden-Green S (2007). Banana //Xanthomonas// wilt: a review of the disease, management strategies and future research directions. African J. Biotechnol. 6: 953-962. PDF: [[https://www.ajol.info/index.php/ajb/article/view/56989|www.ajol.info/index.php/ajb/article/view/56989]]

Cox KL Jr, Babilonia K, Wheeler T, He P, Shan L (2019). Return of old foes - recurrence of bacterial blight and Fusarium wilt of cotton. Curr. Opin. Plant Biol. 50: 95-103. DOI: [[https://doi.org/10.1016/j.pbi.2019.03.012|10.1016/j.pbi.2019.03.012]]

Endah R, Coutinho T, Chikwamba R (2009). //Xanthomonas campestris// pv. //musacearum// induces sequential expression of two NPR-1 like genes in banana. Aspects Appl. Biol. 96: 325-330.

Jin Y, Goodman RE, Tetteh AO, Lu M, Tripathi L (2017). Bioinformatics analysis to assess potential risks of allergenicity and toxicity of HRAP and PFLP proteins in genetically modified bananas resistant to //Xanthomonas// wilt disease. Food Chem. Toxicol. 109: 81-89. DOI: [[https://doi.org/10.1016/j.fct.2017.08.024|10.1016/j.fct.2017.08.024]]

Merga IF, Tripathi L, Hvoslef-Eide AK, Gebre E (2019). Application of genetic engineering for control of bacterial wilt disease of enset, Ethiopia's sustainability crop. Front. Plant Sci. 10: 133. DOI: [[https://doi.org/10.3389/fpls.2019.00133|10.3389/fpls.2019.00133]]

Muwongeab A, Tripathi J, Kunert K, Tripathi L (2016). Expressing stacked //HRAP// and //PFLP// genes in transgenic banana has no synergistic effect on resistance to //Xanthomonas// wilt disease. South African J. Botany 104: 125-133. DOI: [[https://doi.org/10.1016/j.sajb.2015.09.017|10.1016/j.sajb.2015.09.017]]

Nakato G, Christelová P, Were E, Nyine M, Coutinho TA, Doležel J, Uwimana B, Swennen R, Mahuku G (2019). Sources of resistance in //Musa// to //Xanthomonas campestris// pv. //musacearum//, the causal agent of banana //xanthomonas// wilt. Plant Pathol. 68: 49-59. DOI: [[https://doi.org/10.1111/ppa.12945|10.1111/ppa.12945]]

Nakato V, Mahuku G, Coutinho T 2018). //Xanthomonas campestris// pv. //musacearum//: a major constraint to banana, plantain and enset production in central and east Africa over the past decade. Mol. Plant Pathol. 19: 525‐536. DOI: [[https://doi.org/doi:10.1111/mpp.12578|doi:10.1111/mpp.12578]]

Smith J, Jones D, Karamura E, Blomme G, Turyagyenda F (2008). An analysis of the risk from //Xanthomonas campestris// pv. //musacearum// to banana cultivation in Eastern, Central and Southern Africa. Bioversity International, Montpellier, France ISBN: 978-972.

Ssekiwoko F, Kiggundu A, Tushemereirwe W, Karamura E, Kunert K (2015). //Xanthomonas vasicola// pv. //musacearum// down-regulates selected defense genes during its interaction with both resistant and susceptible banana. Physiol. Mol. Plant Pathol. 90: 21-26. DOI: [[https://doi.org/10.1016/j.pmpp.2015.02.007|10.1016/j.pmpp.2015.02.007]]

Tripathi L, Atkinson H, Roderick H, Kubiriba J, Tripathi JN (2017). Genetically engineered bananas resistant to //Xanthomonas// wilt disease and nematodes. Food Energy Secur. 6: 37-47. DOI: [[https://doi.org/10.1002/fes3.101|10.1002/fes3.101]]

Tripathi JN, Lorenzen J, Bahar O, Ronald P, Tripathi L (2014). Transgenic expression of the rice //Xa21// pattern-recognition receptor in banana (//Musa// sp.) confers resistance to //Xanthomonas// //campestris// pv. //musacearum//. Plant Biotechnol. J. 12: 663-673. doi: [[https://doi.org/10.1111/pbi.12170|10.1111/pbi.12170]]

Tripathi L, Mwaka H, Tripathi JN, Tushemereirwe WK (2010). Expression of sweet pepper //Hrap// gene in banana enhances resistance to //Xanthomonas campestris// pv. //musacearum//. Mol. Plant Pathol. 11: 721-731. DOI: [[https://doi.org/10.1111/j.1364-3703.2010.00639.x|10.1111/j.1364-3703.2010.00639.x]]

Tripathi L, Ntui VO, Tripathi JN (2022). Control of bacterial diseases of banana using CRISPR/Cas-based gene editing. Int. J. Mol. Sci. 23: 3619. DOI: [[https://doi.org/10.3390/ijms23073619|10.3390/ijms23073619]]

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===== Acknowledgements =====

This fact sheet is based upon work from COST Action CA16107 EuroXanth, supported by COST (European Cooperation in Science and Technology).
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