Harnessing Advanced Genomic Approaches to Unveil and Enhance Brown Planthopper Resistance in Rice

doi:10.1016/j.rsci.2025.04.003

Abstract

Abstract:

Brown planthopper (BPH) is a highly destructive pest that presents a significant challenge to rice production, particularly in the Asia-Pacific region. Numerous BPH-resistant rice varieties have been successfully bred and released for commercial cultivation across diverse rice-growing ecosystems. However, resistance breakdown in several varieties carrying major resistance genes has been reported, highlighting the urgent need for the development of novel, genetically diverse, and broad-spectrum resistant varieties. To date, more than 45 resistance loci have been identified and mapped from both cultivated and wild rice species. Among these, a subset of genes (including Bph1, Bph3, Bph6, Bph7, Bph9, Bph10, Bph14, Bph15, Bph18, Bph21, Bph26/2, bph29, Bph32, Bph37, and Bph30/Bph40) have been positionally cloned. Most of these genes encode coiled-coil nucleotide-binding leucine-rich repeat proteins, which are central to plant immune responses, along with a few signaling molecules playing pivotal roles. In addition to these core resistance genes, various other genetic components, including miRNAs, protein kinases, and transcription factors, have been functionally characterized for their roles in mediating BPH resistance. The advent of post-genomic tools such as RNA sequencing and single-cell sequencing, along with cutting-edge genomic technologies like CRISPR/Cas gene editing, has significantly accelerated resistance breeding programs. In this context, we provide an overview of genetics, mapping, isolation, and functional characterization of BPH resistance, along with strategies for incorporating resistance using advanced genomics-assisted breeding tools. Furthermore, we present a snapshot of how the integration of genomics and novel breeding technologies holds great promise for dissecting the genetic architecture of pest resistance and accelerating crop improvement.

Key words: brown planthopper, cloning, CRISPR/Cas, genomics, mapping

Lakshmi V. G. Ishwarya, S. Vanisri, P. S. Basavaraj, M. Sreedhar, Lakshmi V. Jhansi, M. Muntazir, C. Gireesh, S. N. C. V. L. Pushpavalli. Harnessing Advanced Genomic Approaches to Unveil and Enhance Brown Planthopper Resistance in Rice[J]. Rice Science, 2025, 32(3): 339-352.

Figures/Tables 4

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[120]	Zhou C, Zhang Q, Chen Y, Huang J, Guo Q, Li Y. 2021. Balancing selection and wild gene pool contribute to resistance in global rice germplasm against planthopper. J Integr Plant Biol, 63(10): 1695-1711.
[121]	Zhou L, Chen Z J, Lang X Y, et al. 2013. Development and validation of a PCR-based functional marker system for the brown planthopper resistance gene Bph14 in rice. Breed Sci, 63(3): 347-352.

Gene/QTL	Chr.	Position	Inheritance	Donor	Molecular marker		Reference
Bph1	12	21.20-22.10 Mb	Dominant	IR64	RM28366-RM463		Yang et al, 2019
	12L	-	Dominant	IR28 (from TKM6)	XNpb248-XNpb336		Hirabayashi and Ogawa, 1995
	12L	2.9 cM	Dominant	Gayabyeo	RG634, RG457, RM247		Jeon et al, 1999
	12L	22.81-22.93 Mb	Dominant	Cheongcheongbyeo	STS-pBPH4, STS-pBPH14		Cha et al, 2008
	12L	22.13-23.18 Mb	Recessive	Norin-PL4 (IR1154-243)	G2140-C449		Murata et al, 1998
	12L	22.13-23.18 Mb	Recessive	Norin-PL4 (IR1154-243)	KAM3-KAM5		Murai et al, 2001
	12L	13.21-22.13 Mb	Recessive	ASD7	RM1246-RM463		Sun et al, 2006
Qbph2	2	22.5-24.0 Mb	-	Col.5 Thailand	RM6843-RM3355		Sun et al, 2007
qBph2	2	68.9 cM	-	Oryza rufipogon	RM29-RG157		Huang et al, 2012
Bph3	4S	6.2-7.00 Mb	Dominant	Rathu Heenati	RZ69		Min et al, 2002
			Dominant	Rathu Heenati	RHD9-RHC10		Liu et al, 2015
			Dominant	Rathu Heenati	4M03980, 4M04041, MM28T		Qing et al, 2019
	6	1.21-1.40 Mb	Dominant	Rathu Heenati	RM589, RM588, RM19291-RM8072		Jairin et al, 2007a, b
	6	0.44 Mb	Dominant	Rathu Heenati	RM508		Temnykh et al, 2001
Qbph3	3	48.6 cM	-	Rathu Heenati	RM313-RM7		Sun et al, 2005
QBph3	3L	35.63-35.67 Mb	-	Oryza officinalis	C3-14		Hu et al, 2015a
qBph-3-1	3	15.7-33.5 Mb	-	Rathu Heenati	RM231-RM3872		Shabanimofrad et al, 2017
bph4	6S	1.20-1.76 Mb	Recessive	Babawee	C76A, RM589-RM586		Kawaguchi et al, 2001; Jairin et al, 2010
QBph4	4	4.41-9.37 Mb	-	Rathu Heenati	RM8213-RM5953		Sun et al, 2005
QBph4.1	4S	6.70-6.90 Mb	-	O. officinalis	P17-xc-427		Hu et al, 2015a
QBph4.2	4	6.58-6.89 Mb	-	Oryza australiensis	RM261-S1, XC4-27		Hu et al, 2015b
qBph4.3	4S	0.50-0.70 Mb	-	Salkathi	RM551-RM335		Mohanty et al, 2017
qBph4.4	4S	0.70-13.10 Mb	-	Salkathi	RM335-RM5633		Mohanty et al, 2017
bph5	-	-	Recessive	ARC10550	-		Khush et al, 1985
Bph6	4L	21.24-21.33 Mb	Dominant	Swarnalata	RM16994-RM119		Kabis and Khush, 1988
Bph6	4L	21.36-21.39 Mb	Dominant	Swarnalata	STSY9-STSY19		Qiu et al, 2010
	11	17.23-18.27 Mb	Dominant	O. officinalis	OPA16938a	Jena et al, 2003
Qbph6	6	2.20-2.80 Mb	-	Col.5 Thailand	RM510-RM587	Sun et al, 2007
qBPH6	6	23.00-23.50 Mb	-	ASD7	RM28466-RM7376	Mai et al, 2015
bph7	12L	19.95-20.87 Mb	Recessive	T12	RM3448-RM313	Kabis and Khush, 1988; Qiu et al, 2014
qBph7	7	19.2-25.4 Mb	-	O. rufipogon	RM11-RM234	Huang et al, 2012
Qbph8	8	20.27 Mb	-	Swarnalatha	RM339-RM515	Cheng et al, 2013
Bph9	12L	19.11-22.13 Mb	Dominant	Kaharamana	RM5341-RM463	Su et al, 2006
	12L	19.00-22.50 Mb	Dominant	Pokkali	G2140, OPR04, S2545	Murata et al, 2001
	12L	22.62-23.20 Mb	Dominant	Pokkali	RM28486-RM28438	Zhao et al, 2016
Bph10	12L	9.00-23.00 Mb	Dominant	O. australiensis	RG457-CDO459, RM260-RM313	Ishii et al, 1994
QBph10	10	21.06-22.43 Mb	-	Rathu Heenati	RM484-RM496	Sun et al, 2005
bph11	3L	35.60-35.80 Mb	Recessive	O. officinalis	G1318	Hirabayashi et al, 1998
Qbph11	11	-	-	DV85	X202-C1172	Su et al, 2005
bph12(t)	4L	20.20-21.20 Mb	Recessive	O. officinalis	G271-R93	Hirabayashi et al, 1999
Bph12	4S	5.21-5.66 Mb	Dominant	B14, Oryza latifolia	RM16459-RM1305	Qiu et al, 2012
Bph12(t)	4S	6.57 Mb	Dominant	B14, O. latifolia	C946-RM261	Yang et al, 2002
qBPH12	12	21.8-24.7 Mb	-	ASD7	RM3326-RM28597	Mai et al, 2015
Bph13(t)	2L	31.40-32.00 Mb	Dominant	Oryza eichingeri	RM240-RM250	Liu et al, 2001
	3S	5.18-5.70 Mb	Dominant	O. officinalis	RG100, RG191, AJ09b, AJ09c	Renganayaki et al, 2002
Bph14	3L	35.70-35.72 Mb	Dominant	B5, O. officinalis	R1925-R2443, R1925-G1318, SM1-G1318, Bph14 P/N	Huang et al, 2001; Ren et al, 2004; Du et al, 2009; Zhou et al, 2013
Bph15	4S	6.68-6.90 Mb	Dominant	B5, O. officinalis	C820, S11182, RG1, RG2, RM261-S16	Huang et al, 2001; Lv et al, 2014
Bph17(t)	4S	6.93-6.97 Mb	Dominant	Rathu Heenati	RM8213-RM5953	Sun et al, 2005
			Dominant	Rathu Heenati	RHD9-RHC10	Liu et al, 2015
			Dominant	Rathu Heenati	LecRK2-SNP, LecRK3-SNP	Kusumawati et al, 2018
Bph18(t)	12L	22.25-23.48 Mb	Dominant	O. australiensis	R10289S-RM6869, S15552-7312.T4A, BIM3-BN162	Jena et al, 2006; Ji et al, 2016
			Dominant	O. australiensis	KC1	Liang et al, 2010
			Dominant	O. australiensis	SNP23-SNP24	Song et al, 2016
			Dominant	O. australiensis	SNP Bph18 marker	Ramkumar et al, 2016
bph18(t)	4L	116.80 cM	Recessive	O. rufipogon	RZ565-RM273	Li et al, 2010
bph19(t)	3S	7.18-7.24 Mb	Recessive	AS20-1	RM6308-RM3134	Chen et al, 2006
Bph20(t)	4S	8.20-9.60 Mb	Dominant	Oryza minuta	B42-B44, MS10-RM5953	Rahman et al, 2009
Bph21(t)	12L	23.28-24.41 Mb	Dominant	O. minuta	B120, B122, RM3726, M5479	Rahman et al, 2009
Bph22(t)	6S	3.40-3.41 Mb	Dominant	O. minuta	RM19429-RM584	Harini et al, 2010
Bph25	6S	0.20-1.71 Mb	Dominant	ADR52	S00310-RM8101	Myint et al, 2012
Bph26	12L	22.13-23.18 Mb	Dominant	IR1154-243	KAM4	Murai et al, 2001
Bph26	12L	21.52-24.46 Mb	Dominant	ADR52	RM5479, RM309, MSSR2, DS-72B4	Myint et al, 2012; Tamura et al, 2014
Bph27	4L	19.12-19.20 Mb	Dominant	O. rufipogon	RM16846-RM16853	Huang et al, 2013
Bph27(t)	4L	20.79-21.33 Mb	Dominant	Balamawee	Q52-Q20	He et al, 2013
Bph28(t)	11L	16.92-16.99 Mb	Dominant	DV85	Indel55-Indel66	Wu et al, 2014
bph20(t)/29	6S	0.48-0.49 Mb	-	O. rufipogon	RM435-RM540, BYL8-BID2	Yang et al, 2012; Wang et al, 2015
bph21(t)/30	10S	12.90-15.00 cM	Recessive	O. rufipogon	RM222-RM244	Yang et al, 2012; Wang et al, 2015
Bph30/40	4S	0.90-0.94 Mb	Dominant	AC-1613	RM16294, RM16299	Wang et al, 2018
Bph31	3L	26.26-26.74 Mb	Dominant	CR2711-76	PA26, RM2334	Prahalada et al, 2017
Bph32	6S	1.21-1.40 Mb	Dominant	PTB33	RM19291, RM8072, PASH6	Ren et al, 2016
Bph32	6S	1.21-1.40 Mb	Dominant	Rathu Heenati	Bph32-SNP, SSR23	Kusumawati et al, 2018
Bph33	4S	0.91-0.97 Mb	Dominant	Kolayal/Poliyal	H99, H101	Hu et al, 2018a
Bph33(t)	1	24.80-28.00 Mb	Dominant	RP2068-18-3-5	RM488-RM11522	Naik et al, 2018
Bph34	4L	21.15-21.30 Mb	Dominant	Oryza nivara	RM16994-RM17007	Kumar et al, 2018
Bph35	4	6.28-6.93 Mb	Dominant	O. rufipogon	PSM16, RM413	Zhang et al, 2020
Bph36	4S	5.57-6.66 Mb	Dominant	O. rufipogon	S13, X48, RM16465-RM16502	Li et al, 2019
Bph37	1	33.00 Mb	Dominant	IR64	RM302, Indel YM35	Yang et al, 2019
Bph38	4	3.72-15.83 Mb	Dominant	O. rufipogon	RM16563-RM16763	Yang et al, 2020
Bph38(t)	1L	20.71-21.23 Mb	Dominant	Khazar	SNP-693369-SNP-id10112165	Balachiranjeevi et al, 2019
BPH41	4S	0.40-1.50 Mb	-	SWD10	SWRm_01617, SWRm_01522	Tan et al, 2022
BPH42	4L	12.5-14.8 Mb	-	SWD10	SWRm_01695, SWRm_00328	Tan et al, 2022
bph42	4S	0.4-16.9 Mb	Recessive	CR100441, O. rufipogon	RM16282-RM16335	Kaur et al, 2022
Bph43	11	16.2-17.6 Mb	Dominant	IRGC 8678	6-22-16-3	Kim et al, 2022
Bph44	4L	4.76-5.42 cM	Dominant	Balamwee	Q31, RM17007	Kiswanto et al, 2022
Bph45	4	13.67-13.70 Mb	Dominant	IRGC102165, O. nivara	RM3317-RM16655	Li et al, 2023
bph46	4S	6.54-18.60 Mb	Recessive	IRGC93198, O. nivara	RM16285, M6314, RM6659	Kaur et al, 2023

Gene/QTL	Chr.	Position	Inheritance	Donor	Molecular marker		Reference
Bph1	12	21.20-22.10 Mb	Dominant	IR64	RM28366-RM463		Yang et al, 2019
	12L	-	Dominant	IR28 (from TKM6)	XNpb248-XNpb336		Hirabayashi and Ogawa, 1995
	12L	2.9 cM	Dominant	Gayabyeo	RG634, RG457, RM247		Jeon et al, 1999
	12L	22.81-22.93 Mb	Dominant	Cheongcheongbyeo	STS-pBPH4, STS-pBPH14		Cha et al, 2008
	12L	22.13-23.18 Mb	Recessive	Norin-PL4 (IR1154-243)	G2140-C449		Murata et al, 1998
	12L	22.13-23.18 Mb	Recessive	Norin-PL4 (IR1154-243)	KAM3-KAM5		Murai et al, 2001
	12L	13.21-22.13 Mb	Recessive	ASD7	RM1246-RM463		Sun et al, 2006
Qbph2	2	22.5-24.0 Mb	-	Col.5 Thailand	RM6843-RM3355		Sun et al, 2007
qBph2	2	68.9 cM	-	Oryza rufipogon	RM29-RG157		Huang et al, 2012
Bph3	4S	6.2-7.00 Mb	Dominant	Rathu Heenati	RZ69		Min et al, 2002
			Dominant	Rathu Heenati	RHD9-RHC10		Liu et al, 2015
			Dominant	Rathu Heenati	4M03980, 4M04041, MM28T		Qing et al, 2019
	6	1.21-1.40 Mb	Dominant	Rathu Heenati	RM589, RM588, RM19291-RM8072		Jairin et al, 2007a, b
	6	0.44 Mb	Dominant	Rathu Heenati	RM508		Temnykh et al, 2001
Qbph3	3	48.6 cM	-	Rathu Heenati	RM313-RM7		Sun et al, 2005
QBph3	3L	35.63-35.67 Mb	-	Oryza officinalis	C3-14		Hu et al, 2015a
qBph-3-1	3	15.7-33.5 Mb	-	Rathu Heenati	RM231-RM3872		Shabanimofrad et al, 2017
bph4	6S	1.20-1.76 Mb	Recessive	Babawee	C76A, RM589-RM586		Kawaguchi et al, 2001; Jairin et al, 2010
QBph4	4	4.41-9.37 Mb	-	Rathu Heenati	RM8213-RM5953		Sun et al, 2005
QBph4.1	4S	6.70-6.90 Mb	-	O. officinalis	P17-xc-427		Hu et al, 2015a
QBph4.2	4	6.58-6.89 Mb	-	Oryza australiensis	RM261-S1, XC4-27		Hu et al, 2015b
qBph4.3	4S	0.50-0.70 Mb	-	Salkathi	RM551-RM335		Mohanty et al, 2017
qBph4.4	4S	0.70-13.10 Mb	-	Salkathi	RM335-RM5633		Mohanty et al, 2017
bph5	-	-	Recessive	ARC10550	-		Khush et al, 1985
Bph6	4L	21.24-21.33 Mb	Dominant	Swarnalata	RM16994-RM119		Kabis and Khush, 1988
Bph6	4L	21.36-21.39 Mb	Dominant	Swarnalata	STSY9-STSY19		Qiu et al, 2010
	11	17.23-18.27 Mb	Dominant	O. officinalis	OPA16938a	Jena et al, 2003
Qbph6	6	2.20-2.80 Mb	-	Col.5 Thailand	RM510-RM587	Sun et al, 2007
qBPH6	6	23.00-23.50 Mb	-	ASD7	RM28466-RM7376	Mai et al, 2015
bph7	12L	19.95-20.87 Mb	Recessive	T12	RM3448-RM313	Kabis and Khush, 1988; Qiu et al, 2014
qBph7	7	19.2-25.4 Mb	-	O. rufipogon	RM11-RM234	Huang et al, 2012
Qbph8	8	20.27 Mb	-	Swarnalatha	RM339-RM515	Cheng et al, 2013
Bph9	12L	19.11-22.13 Mb	Dominant	Kaharamana	RM5341-RM463	Su et al, 2006
	12L	19.00-22.50 Mb	Dominant	Pokkali	G2140, OPR04, S2545	Murata et al, 2001
	12L	22.62-23.20 Mb	Dominant	Pokkali	RM28486-RM28438	Zhao et al, 2016
Bph10	12L	9.00-23.00 Mb	Dominant	O. australiensis	RG457-CDO459, RM260-RM313	Ishii et al, 1994
QBph10	10	21.06-22.43 Mb	-	Rathu Heenati	RM484-RM496	Sun et al, 2005
bph11	3L	35.60-35.80 Mb	Recessive	O. officinalis	G1318	Hirabayashi et al, 1998
Qbph11	11	-	-	DV85	X202-C1172	Su et al, 2005
bph12(t)	4L	20.20-21.20 Mb	Recessive	O. officinalis	G271-R93	Hirabayashi et al, 1999
Bph12	4S	5.21-5.66 Mb	Dominant	B14, Oryza latifolia	RM16459-RM1305	Qiu et al, 2012
Bph12(t)	4S	6.57 Mb	Dominant	B14, O. latifolia	C946-RM261	Yang et al, 2002
qBPH12	12	21.8-24.7 Mb	-	ASD7	RM3326-RM28597	Mai et al, 2015
Bph13(t)	2L	31.40-32.00 Mb	Dominant	Oryza eichingeri	RM240-RM250	Liu et al, 2001
	3S	5.18-5.70 Mb	Dominant	O. officinalis	RG100, RG191, AJ09b, AJ09c	Renganayaki et al, 2002
Bph14	3L	35.70-35.72 Mb	Dominant	B5, O. officinalis	R1925-R2443, R1925-G1318, SM1-G1318, Bph14 P/N	Huang et al, 2001; Ren et al, 2004; Du et al, 2009; Zhou et al, 2013
Bph15	4S	6.68-6.90 Mb	Dominant	B5, O. officinalis	C820, S11182, RG1, RG2, RM261-S16	Huang et al, 2001; Lv et al, 2014
Bph17(t)	4S	6.93-6.97 Mb	Dominant	Rathu Heenati	RM8213-RM5953	Sun et al, 2005
			Dominant	Rathu Heenati	RHD9-RHC10	Liu et al, 2015
			Dominant	Rathu Heenati	LecRK2-SNP, LecRK3-SNP	Kusumawati et al, 2018
Bph18(t)	12L	22.25-23.48 Mb	Dominant	O. australiensis	R10289S-RM6869, S15552-7312.T4A, BIM3-BN162	Jena et al, 2006; Ji et al, 2016
			Dominant	O. australiensis	KC1	Liang et al, 2010
			Dominant	O. australiensis	SNP23-SNP24	Song et al, 2016
			Dominant	O. australiensis	SNP Bph18 marker	Ramkumar et al, 2016
bph18(t)	4L	116.80 cM	Recessive	O. rufipogon	RZ565-RM273	Li et al, 2010
bph19(t)	3S	7.18-7.24 Mb	Recessive	AS20-1	RM6308-RM3134	Chen et al, 2006
Bph20(t)	4S	8.20-9.60 Mb	Dominant	Oryza minuta	B42-B44, MS10-RM5953	Rahman et al, 2009
Bph21(t)	12L	23.28-24.41 Mb	Dominant	O. minuta	B120, B122, RM3726, M5479	Rahman et al, 2009
Bph22(t)	6S	3.40-3.41 Mb	Dominant	O. minuta	RM19429-RM584	Harini et al, 2010
Bph25	6S	0.20-1.71 Mb	Dominant	ADR52	S00310-RM8101	Myint et al, 2012
Bph26	12L	22.13-23.18 Mb	Dominant	IR1154-243	KAM4	Murai et al, 2001
Bph26	12L	21.52-24.46 Mb	Dominant	ADR52	RM5479, RM309, MSSR2, DS-72B4	Myint et al, 2012; Tamura et al, 2014
Bph27	4L	19.12-19.20 Mb	Dominant	O. rufipogon	RM16846-RM16853	Huang et al, 2013
Bph27(t)	4L	20.79-21.33 Mb	Dominant	Balamawee	Q52-Q20	He et al, 2013
Bph28(t)	11L	16.92-16.99 Mb	Dominant	DV85	Indel55-Indel66	Wu et al, 2014
bph20(t)/29	6S	0.48-0.49 Mb	-	O. rufipogon	RM435-RM540, BYL8-BID2	Yang et al, 2012; Wang et al, 2015
bph21(t)/30	10S	12.90-15.00 cM	Recessive	O. rufipogon	RM222-RM244	Yang et al, 2012; Wang et al, 2015
Bph30/40	4S	0.90-0.94 Mb	Dominant	AC-1613	RM16294, RM16299	Wang et al, 2018
Bph31	3L	26.26-26.74 Mb	Dominant	CR2711-76	PA26, RM2334	Prahalada et al, 2017
Bph32	6S	1.21-1.40 Mb	Dominant	PTB33	RM19291, RM8072, PASH6	Ren et al, 2016
Bph32	6S	1.21-1.40 Mb	Dominant	Rathu Heenati	Bph32-SNP, SSR23	Kusumawati et al, 2018
Bph33	4S	0.91-0.97 Mb	Dominant	Kolayal/Poliyal	H99, H101	Hu et al, 2018a
Bph33(t)	1	24.80-28.00 Mb	Dominant	RP2068-18-3-5	RM488-RM11522	Naik et al, 2018
Bph34	4L	21.15-21.30 Mb	Dominant	Oryza nivara	RM16994-RM17007	Kumar et al, 2018
Bph35	4	6.28-6.93 Mb	Dominant	O. rufipogon	PSM16, RM413	Zhang et al, 2020
Bph36	4S	5.57-6.66 Mb	Dominant	O. rufipogon	S13, X48, RM16465-RM16502	Li et al, 2019
Bph37	1	33.00 Mb	Dominant	IR64	RM302, Indel YM35	Yang et al, 2019
Bph38	4	3.72-15.83 Mb	Dominant	O. rufipogon	RM16563-RM16763	Yang et al, 2020
Bph38(t)	1L	20.71-21.23 Mb	Dominant	Khazar	SNP-693369-SNP-id10112165	Balachiranjeevi et al, 2019
BPH41	4S	0.40-1.50 Mb	-	SWD10	SWRm_01617, SWRm_01522	Tan et al, 2022
BPH42	4L	12.5-14.8 Mb	-	SWD10	SWRm_01695, SWRm_00328	Tan et al, 2022
bph42	4S	0.4-16.9 Mb	Recessive	CR100441, O. rufipogon	RM16282-RM16335	Kaur et al, 2022
Bph43	11	16.2-17.6 Mb	Dominant	IRGC 8678	6-22-16-3	Kim et al, 2022
Bph44	4L	4.76-5.42 cM	Dominant	Balamwee	Q31, RM17007	Kiswanto et al, 2022
Bph45	4	13.67-13.70 Mb	Dominant	IRGC102165, O. nivara	RM3317-RM16655	Li et al, 2023
bph46	4S	6.54-18.60 Mb	Recessive	IRGC93198, O. nivara	RM16285, M6314, RM6659	Kaur et al, 2023

Gene	Locus name	Gene ID	Annotation
Bph3	LOC_Os04g12540	Os04g0201900	Plasma membrane-localized lectin receptor kinases
Bph3	LOC_Os04g12580	Os04g0202500	Plasma membrane-localized lectin receptor kinases
Bph15	LOC_Os04g12390	Os04g0200400	Lectin receptor kinases
Bph15	LOC_Os04g12460	Os04g0201000	Lectin receptor kinases
Bph18	LOC_Os12g37290	Os12g0559400	Resistance protein T10rga2-1A, putative, expressed
Bph18	LOC_Os12g37280	Os12g0559400	Resistance protein T10rga2-1A, putative, expressed
Bph27	LOC_Os04g31924	Os04g0388700	Putative nodulin protein
	LOC_Os04g32000	Os04g0389600	Putative MYB transcription factor
	LOC_Os04g53190.2	Os04g0623300	TGF beta receptor associated protein
bph29	LOC_Os06g01850	Os06g0107700	Ferredoxin NADP reductase, chloroplast precursor
bph29	LOC_Os06g01860	Os06g0107800	Expressed protein
Bph30	LOC_Os04g08390	Os04g0166000	LRR family protein, expressed
Bph31	LOC_Os03g46550	Os03g0668300	LRR domain protein
	LOC_Os03g46454	Os03g0667300	Metal cation transporter, putative, expressed
	LOC_Os03g46440	Os03g0667100	BTB domain with Ankyrin repeat region, expressed
Bph32	LOC_Os06g03660	Os06g0127000	Peroxisomal biogenesis factor 11, putative
Bph32	LOC_Os06g03890	Os06g0129600	Plasma membrane receptor proteins
Bph33	LOC_Os04g02520	Os04g0115650	LRR family protein
Bph33(t)	LOC_Os01g42190.1	Os01g0606900	Heat shock protein DnaJ, putative, expressed
Bph34	LOC_Os04g35210	Os04g0431700	LRR family protein
Bph36	LOC_Os04g11800	Os04g0194400	Hypothetical protein
	LOC_Os04g11810	Os04g0194466	Acyltransferase family protein
	LOC_Os04g11820	Os04g0194500	White-brown complex homolog protein
	LOC_Os04g11830	Os04g0194600	TCP family transcription factor
Bph37	LOC_Os06g03500	Os06g0125000	NBS-LRR disease resistance protein, putative
Bph38(t)	LOC_Os01g37260	Os01g0553400	FBXL class of F-box protein possessing LRR
Bph43	LOC_Os11g29000	Os11g0479400	Pollen signaling protein with adenylyl cyclase activity
	LOC_Os11g29050	Os11g0480000	NBS-LRR type disease resistance protein
	LOC_Os11g29090	Os11g0481000	NB-ARC/LRR disease resistance protein
	LOC_Os11g29110	Os11g0481150	LRR family protein, expressed

Gene	Locus name	Gene ID	Annotation
Bph3	LOC_Os04g12540	Os04g0201900	Plasma membrane-localized lectin receptor kinases
Bph3	LOC_Os04g12580	Os04g0202500	Plasma membrane-localized lectin receptor kinases
Bph15	LOC_Os04g12390	Os04g0200400	Lectin receptor kinases
Bph15	LOC_Os04g12460	Os04g0201000	Lectin receptor kinases
Bph18	LOC_Os12g37290	Os12g0559400	Resistance protein T10rga2-1A, putative, expressed
Bph18	LOC_Os12g37280	Os12g0559400	Resistance protein T10rga2-1A, putative, expressed
Bph27	LOC_Os04g31924	Os04g0388700	Putative nodulin protein
	LOC_Os04g32000	Os04g0389600	Putative MYB transcription factor
	LOC_Os04g53190.2	Os04g0623300	TGF beta receptor associated protein
bph29	LOC_Os06g01850	Os06g0107700	Ferredoxin NADP reductase, chloroplast precursor
bph29	LOC_Os06g01860	Os06g0107800	Expressed protein
Bph30	LOC_Os04g08390	Os04g0166000	LRR family protein, expressed
Bph31	LOC_Os03g46550	Os03g0668300	LRR domain protein
	LOC_Os03g46454	Os03g0667300	Metal cation transporter, putative, expressed
	LOC_Os03g46440	Os03g0667100	BTB domain with Ankyrin repeat region, expressed
Bph32	LOC_Os06g03660	Os06g0127000	Peroxisomal biogenesis factor 11, putative
Bph32	LOC_Os06g03890	Os06g0129600	Plasma membrane receptor proteins
Bph33	LOC_Os04g02520	Os04g0115650	LRR family protein
Bph33(t)	LOC_Os01g42190.1	Os01g0606900	Heat shock protein DnaJ, putative, expressed
Bph34	LOC_Os04g35210	Os04g0431700	LRR family protein
Bph36	LOC_Os04g11800	Os04g0194400	Hypothetical protein
	LOC_Os04g11810	Os04g0194466	Acyltransferase family protein
	LOC_Os04g11820	Os04g0194500	White-brown complex homolog protein
	LOC_Os04g11830	Os04g0194600	TCP family transcription factor
Bph37	LOC_Os06g03500	Os06g0125000	NBS-LRR disease resistance protein, putative
Bph38(t)	LOC_Os01g37260	Os01g0553400	FBXL class of F-box protein possessing LRR
Bph43	LOC_Os11g29000	Os11g0479400	Pollen signaling protein with adenylyl cyclase activity
	LOC_Os11g29050	Os11g0480000	NBS-LRR type disease resistance protein
	LOC_Os11g29090	Os11g0481000	NB-ARC/LRR disease resistance protein
	LOC_Os11g29110	Os11g0481150	LRR family protein, expressed