The pathogenesis-related protein PR10 plays a vital role in plant growth,development,and stress responses.This study systematically identified and analyzed PR10 genes in cultivated peanut(Arachis hypogaea L.),examinin...The pathogenesis-related protein PR10 plays a vital role in plant growth,development,and stress responses.This study systematically identified and analyzed PR10 genes in cultivated peanut(Arachis hypogaea L.),examining their phylogenetic relationships,conserved motifs,gene structures,and syntenic relationships.The analysis identified 54 Ah PR10 genes,which were classified into eight groups based on phylogenetic relationships,supported by gene structure and conserved motif characterization.Analysis of chromosomal distribution and synteny demonstrated that segmental duplications played a crucial role in the expansion of the Ah PR10 gene family.The identified Ah PR10 genes exhibited both constitutive and inducible expression patterns.Significantly,Ah PR10-7,Ah PR10-33,and Ah PR10-41 demonstrated potential importance in peanut resistance to Aspergillus flavus.In vitro fungistatic experiments demonstrated that recombinant Ah PR10-33 effectively inhibited A.flavus mycelial growth.These findings provide valuable insights for future investigations into Ah PR10 functions in protecting peanut from A.flavus infection.展开更多
Peanut kernels rich in oil,particularly those with oleic acid as their primary fatty acid,are in high demand among consumers,the food industry,and farmers due to their superior nutritional content,extended shelf life,...Peanut kernels rich in oil,particularly those with oleic acid as their primary fatty acid,are in high demand among consumers,the food industry,and farmers due to their superior nutritional content,extended shelf life,and health benefits.The oil content and fatty acid composition are governed by multiple genetic factors.Identifying the quantitative trait loci(QTLs)related to these attributes will facilitate marker-assisted selection and genomic selection,thus enhancing quality-focused peanut breeding programs.For this purpose,we developed a population of 521 recombinant inbred lines(RILs)and tested their kernel quality traits across five different environments.We identified two major and stable QTLs for oil content,qOCAh12.1 and qOCAh16.1.The markers linked to these QTLs were designed by Kompetitive allele-specific PCR(KASP)and subsequently validated.Moreover,we found that the superior haplotype of oil content in the qOCAh16.1 region was conserved within the plant introduction(PI)germplasm cluster,as evidenced by a diverse peanut accession panel.In addition,we determined that qAh09 and qAh19.1,which harbor the key gene encoding fatty acid desaturase 2(FAD2),influence all seven fatty acids,palmitic,stearic,oleic,linoleic,arachidic,gadoleic,and behenic acids.Regarding the protein content and the long-chain saturated fatty acid behenic acid,qAh07 emerged as the major and stable QTL,accounting for over 10%of the phenotypic variation explained(PVE).These findings can enhance marker-assisted selection in peanut breeding,with the aim of improving the oil content,and deepen our understanding of the genetic mechanisms that shape fatty acid composition.展开更多
Wild peanut(Arachis)species are promising sources of disease resistance for improving peanut cultivars.The objective of this study was to assess cross-compatibility among cultivated and wild peanuts in crosses between...Wild peanut(Arachis)species are promising sources of disease resistance for improving peanut cultivars.The objective of this study was to assess cross-compatibility among cultivated and wild peanuts in crosses between eight peanut cultivars and 27 wild species carrying the A,B,E,Ex,F,K,P,and H genomes.Embryo culture and chromosome doubling led to polyploids representing hybrids between cultivated peanut and A.stenosperma,A.macedoi,A.duranensis,A.villosa,and A.diogoi.The first two showed greater resistance to bacterial wilt than their cultivated parents.DNA markers were developed for verifying the hybrids and for identifying translocation or introgression lines with alien chromosome fragments.展开更多
Unsteady aerodynamic characteristics at high angles of attack are of great importance to the design and development of advanced fighter aircraft, which are characterized by post-stall maneuverability with multiple Deg...Unsteady aerodynamic characteristics at high angles of attack are of great importance to the design and development of advanced fighter aircraft, which are characterized by post-stall maneuverability with multiple Degrees-of-Freedom(multi-DOF) and complex flow field structure.In this paper, a special kind of cable-driven parallel mechanism is firstly utilized as a new suspension method to conduct unsteady dynamic wind tunnel tests at high angles of attack, thereby providing experimental aerodynamic data. These tests include a wide range of multi-DOF coupled oscillatory motions with various amplitudes and frequencies. Then, for aerodynamic modeling and analysis, a novel data-driven Feature-Level Attention Recurrent neural network(FLAR) is proposed. This model incorporates a specially designed feature-level attention module that focuses on the state variables affecting the aerodynamic coefficients, thereby enhancing the physical interpretability of the aerodynamic model. Subsequently, spin maneuver simulations, using a mathematical model as the baseline, are conducted to validate the effectiveness of the FLAR. Finally, the results on wind tunnel data reveal that the FLAR accurately predicts aerodynamic coefficients, and observations through the visualization of attention scores identify the key state variables that affect the aerodynamic coefficients. It is concluded that the proposed FLAR enhances the interpretability of the aerodynamic model while achieving good prediction accuracy and generalization capability for multi-DOF coupling motion at high angles of attack.展开更多
Oil and protein content and fatty acid composition are quality traits in peanut.Elucidating the genetic mechanisms underlying these traits may help researchers to obtain improved cultivars by molecular breeding.Whole-...Oil and protein content and fatty acid composition are quality traits in peanut.Elucidating the genetic mechanisms underlying these traits may help researchers to obtain improved cultivars by molecular breeding.Whole-genome resequencing of a recombinant inbred population of 318 lines was performed to construct a high-density linkage map and identify QTL for peanut quality.The map,containing 4561 bin markers,covered 2032 c M with a mean marker density of 0.45 c M.A total of 110 QTL for oil and protein content,and fatty acid composition were mapped on the 18 peanut chromosomes.The QTL q A05.1 was detected in four environments and showed a major phenotypic effect on the contents of oil,protein,and six fatty acids.The genomic region spanned by q A05.1,corresponding to a physical interval of approximately 1.5 Mb,contains two SNPs polymorphic between the parents that could cause missense mutations.The two SNP sites were employed as KASP markers and validated using lines with extremely high and low oil contents.These sites may be useful in the marker-assisted breeding of peanut cultivars with high oil contents.展开更多
Fresh-seed germination(FSG)impairs peanut production,especially in areas where the peanut harvest season coincides with rainy weather.Developing FSG-resistant cultivars by molecular breeding is expected to mitigate yi...Fresh-seed germination(FSG)impairs peanut production,especially in areas where the peanut harvest season coincides with rainy weather.Developing FSG-resistant cultivars by molecular breeding is expected to mitigate yield loss and quality impairment caused by FSG.However,the genetic control of FSG awaits elucidation.In this study,FSG at 1,3,5,7,and 9 days post-imbibition in three environments were tested,and quantitative-trait loci(QTL)associated with FSG were mapped in a peanut recombinant inbred line population by leveraging existing high-density peanut genetic maps.Of 24 QTL identified in 13 linkage groups,qFSGA04 was a stable major QTL on linkage group 04(LG04).It was consistently detected in five germination stages and three environments.By designing and validating DNA markers in the confidence interval of qFSGA04,we identified one single-nucleotide polymorphism and one In Del closely associated with FSG that could be used as linked markers for FSG resistance in peanut breeding.展开更多
Oligo probe staining is a low-cost and efficient chromosome identification technique.In this study,oligo genomic in situ hybridization(Oligo-GISH)technology was established in peanut.Peanut A and B subgenome-specific ...Oligo probe staining is a low-cost and efficient chromosome identification technique.In this study,oligo genomic in situ hybridization(Oligo-GISH)technology was established in peanut.Peanut A and B subgenome-specific interspersed repeat(IR)oligo probe sets were developed based on clustering and electronic localization of tandem repeat sequences in the reference genome of Tifrunner.The OligoGISH kit was then used to perform staining of 15 Arachis species.The A-subgenome probe set stained the chromosomes of A-and E-genome Arachis species,the B-subgenome probe set stained those of B-,F-,K-,and E-genome species,and neither set stained those of H-genome species.These results indicate the relationships among the genomes of these Arachis species.The Oligo-GISH kit was also used for batch staining of the chromosomes of 389 seedlings from the irradiated M1generation,allowing 67 translocation and deletion lines to be identified.Subsequent Oligo-FISH karyotyping,FISH using single-copy probe libraries,and trait investigation identified seven homozygous chromosomal variants from the M3generation and suggested that there may be genes on chromosome 4B controlling seed number per pod.These findings demonstrate that the IR probe sets and method developed in this study can facilitate research on distant hybridization and genetic improvement in peanut.展开更多
Peanut(Arachis hypogaea L.)is a globally important oil crop.Web blotch is one of the most important foliar diseases affecting peanut,which results in serious yield losses worldwide.Breeding web blotch-resistant peanut...Peanut(Arachis hypogaea L.)is a globally important oil crop.Web blotch is one of the most important foliar diseases affecting peanut,which results in serious yield losses worldwide.Breeding web blotch-resistant peanut varieties is the most effective and economically viable method for minimizing yield losses due to web blotch.In the current study,a bulked segregant analysis with next-generation sequencing was used to analyze an F2:3 segregating population and identify candidate loci related to web blotch resistance.Based on the fine-mapping of the candidate genomic interval using kompetitive allele-specific PCR(KASP)markers,we identified a novel web blotch resistance-related locus spanning approximately 169 kb on chromosome 16.This region included four annotated genes,of which only Arahy.35VVQ3 had a non-synonymous single nucleotide polymorphism in the coding region between the two parents.Two markers(Chr.16.12872635 and Chr.16.12966357)linked to this gene were shown to be co-segregated with the resistance of peanut web blotch by 72 randomly selected recombinant inbred lines(RIL),which could be used in marker-assisted breeding of resistant peanut varieties.展开更多
Dear Editor,Cultivated peanut(peanut in short)(Arachis hypogaea,2n=4x=40,genome type AABB)is a legume crop serving globally as a crucial source of edible oil and protein.Available peanut reference genome sequences obt...Dear Editor,Cultivated peanut(peanut in short)(Arachis hypogaea,2n=4x=40,genome type AABB)is a legume crop serving globally as a crucial source of edible oil and protein.Available peanut reference genome sequences obtained for the cultivars"Tifrunner,""Shitouqi,"and"Fuhuasheng"were assembled or patched using third-generation sequencing technologies and significantly contributed to advancements in peanut genetics and breeding(Bertioli et al.,2019;Chen et al.,2019;Zhuang et al.,2019).However,unresolved gaps persist in these assemblies,particularly in regions of biological importance such as centromeres and telomeres(Supplemental Figure 1),hindering comprehensive exploration of their functional and evolutionary significance.展开更多
Peanut (Arachis hypogaea L.) is an important food and oil crop valued for its oil-rich kernel, high protein content, and diverse nutritional components, making it a staple raw material in the food industry. Fragrance ...Peanut (Arachis hypogaea L.) is an important food and oil crop valued for its oil-rich kernel, high protein content, and diverse nutritional components, making it a staple raw material in the food industry. Fragrance is an essential trait influencing both the initial appeal and flavor quality of food products. For instance, fragrant rice has gained global popularity due to its appealing aroma. The fragrance trait in rice is regulated by the BADH2 gene, which encodes betaine aldehyde dehydrogenase 2 (BADH) (Chen et al., 2008). In non-fragrant rice, BADH2 catalyzes the conversion of γ-aminobutyraldehyde (GABald) to γ-aminobutyric acid. When BADH2 function is disrupted, GABald is redirected to form Δ-1-pyrroline and the subsequent 2-acetyl-1-pyrroline (2-AP), the key compound contributing to rice fragrance (Okpala et al., 2019).展开更多
基金supported by the National Key R&D Program of China(2022YFD1200400)the National Natural Science Foundation of China(32301851)。
摘要The pathogenesis-related protein PR10 plays a vital role in plant growth,development,and stress responses.This study systematically identified and analyzed PR10 genes in cultivated peanut(Arachis hypogaea L.),examining their phylogenetic relationships,conserved motifs,gene structures,and syntenic relationships.The analysis identified 54 Ah PR10 genes,which were classified into eight groups based on phylogenetic relationships,supported by gene structure and conserved motif characterization.Analysis of chromosomal distribution and synteny demonstrated that segmental duplications played a crucial role in the expansion of the Ah PR10 gene family.The identified Ah PR10 genes exhibited both constitutive and inducible expression patterns.Significantly,Ah PR10-7,Ah PR10-33,and Ah PR10-41 demonstrated potential importance in peanut resistance to Aspergillus flavus.In vitro fungistatic experiments demonstrated that recombinant Ah PR10-33 effectively inhibited A.flavus mycelial growth.These findings provide valuable insights for future investigations into Ah PR10 functions in protecting peanut from A.flavus infection.
基金supported by the National Key R&D Program of China(2022YFD1200400)the earmarked fund for CARS-13,the Major Science and Technology Projects of Henan Province,China(221100110300)the Henan Provincial R&D Program of Interregional Cooperation for Local Scientific and Technological Development Guided by Central Government,China(YDZX20214100004191)。
摘要Peanut kernels rich in oil,particularly those with oleic acid as their primary fatty acid,are in high demand among consumers,the food industry,and farmers due to their superior nutritional content,extended shelf life,and health benefits.The oil content and fatty acid composition are governed by multiple genetic factors.Identifying the quantitative trait loci(QTLs)related to these attributes will facilitate marker-assisted selection and genomic selection,thus enhancing quality-focused peanut breeding programs.For this purpose,we developed a population of 521 recombinant inbred lines(RILs)and tested their kernel quality traits across five different environments.We identified two major and stable QTLs for oil content,qOCAh12.1 and qOCAh16.1.The markers linked to these QTLs were designed by Kompetitive allele-specific PCR(KASP)and subsequently validated.Moreover,we found that the superior haplotype of oil content in the qOCAh16.1 region was conserved within the plant introduction(PI)germplasm cluster,as evidenced by a diverse peanut accession panel.In addition,we determined that qAh09 and qAh19.1,which harbor the key gene encoding fatty acid desaturase 2(FAD2),influence all seven fatty acids,palmitic,stearic,oleic,linoleic,arachidic,gadoleic,and behenic acids.Regarding the protein content and the long-chain saturated fatty acid behenic acid,qAh07 emerged as the major and stable QTL,accounting for over 10%of the phenotypic variation explained(PVE).These findings can enhance marker-assisted selection in peanut breeding,with the aim of improving the oil content,and deepen our understanding of the genetic mechanisms that shape fatty acid composition.
基金supported by National Natural Science Foundation of China(32272153)Henan Province Science and Technology R&D Joint Fund(232301420025)+4 种基金National Key Research and Development Program of China(2023YFD1200200)the Key Research Project of the Shennong Laboratory(SN01-2022-03)Independent Innovation Foundation of Henan Academy of Agricultural Sciences(2024ZC024)China Agriculture Research System(CARS-13)Henan Provincial Agriculture Research System(S2012-5).
摘要Wild peanut(Arachis)species are promising sources of disease resistance for improving peanut cultivars.The objective of this study was to assess cross-compatibility among cultivated and wild peanuts in crosses between eight peanut cultivars and 27 wild species carrying the A,B,E,Ex,F,K,P,and H genomes.Embryo culture and chromosome doubling led to polyploids representing hybrids between cultivated peanut and A.stenosperma,A.macedoi,A.duranensis,A.villosa,and A.diogoi.The first two showed greater resistance to bacterial wilt than their cultivated parents.DNA markers were developed for verifying the hybrids and for identifying translocation or introgression lines with alien chromosome fragments.
基金supported by the National Natural Science Foundation of China(Nos.12172315,12072304,11702232)the Fujian Provincial Natural Science Foundation,China(No.2021J01050)the Aeronautical Science Foundation of China(No.20220013068002).
摘要Unsteady aerodynamic characteristics at high angles of attack are of great importance to the design and development of advanced fighter aircraft, which are characterized by post-stall maneuverability with multiple Degrees-of-Freedom(multi-DOF) and complex flow field structure.In this paper, a special kind of cable-driven parallel mechanism is firstly utilized as a new suspension method to conduct unsteady dynamic wind tunnel tests at high angles of attack, thereby providing experimental aerodynamic data. These tests include a wide range of multi-DOF coupled oscillatory motions with various amplitudes and frequencies. Then, for aerodynamic modeling and analysis, a novel data-driven Feature-Level Attention Recurrent neural network(FLAR) is proposed. This model incorporates a specially designed feature-level attention module that focuses on the state variables affecting the aerodynamic coefficients, thereby enhancing the physical interpretability of the aerodynamic model. Subsequently, spin maneuver simulations, using a mathematical model as the baseline, are conducted to validate the effectiveness of the FLAR. Finally, the results on wind tunnel data reveal that the FLAR accurately predicts aerodynamic coefficients, and observations through the visualization of attention scores identify the key state variables that affect the aerodynamic coefficients. It is concluded that the proposed FLAR enhances the interpretability of the aerodynamic model while achieving good prediction accuracy and generalization capability for multi-DOF coupling motion at high angles of attack.
基金supported by the National Basic Research Program of ChinaSpecial Project for National Supercomputing Zhengzhou Center Innovation Ecosystem Construction(201400210600)+4 种基金Outstanding Young Scientists of Henan Academy of Agricultural Sciences(2020YQ08)Fund for Distinguished Young Scholars from Henan Academy of Agricultural Sciences(2019JQ02)China Agriculture Research System(CARS-13)Henan Provincial Agriculture Research System,China(S2012-5)Henan Provincial Young Talents Supporting Project(2020HYTP044)。
摘要Oil and protein content and fatty acid composition are quality traits in peanut.Elucidating the genetic mechanisms underlying these traits may help researchers to obtain improved cultivars by molecular breeding.Whole-genome resequencing of a recombinant inbred population of 318 lines was performed to construct a high-density linkage map and identify QTL for peanut quality.The map,containing 4561 bin markers,covered 2032 c M with a mean marker density of 0.45 c M.A total of 110 QTL for oil and protein content,and fatty acid composition were mapped on the 18 peanut chromosomes.The QTL q A05.1 was detected in four environments and showed a major phenotypic effect on the contents of oil,protein,and six fatty acids.The genomic region spanned by q A05.1,corresponding to a physical interval of approximately 1.5 Mb,contains two SNPs polymorphic between the parents that could cause missense mutations.The two SNP sites were employed as KASP markers and validated using lines with extremely high and low oil contents.These sites may be useful in the marker-assisted breeding of peanut cultivars with high oil contents.
基金supported by China Agriculture Research System(CARS-13)Henan Provincial Agriculture Research System,China(S2012-5)+1 种基金Major Science and Technology Projects of Henan Province(201300111000)the Henan Provincial R&D Projects of Interregional Cooperation for Local Scientific and Technological Development Guided by Central Government(YDZX20214100004191)。
摘要Fresh-seed germination(FSG)impairs peanut production,especially in areas where the peanut harvest season coincides with rainy weather.Developing FSG-resistant cultivars by molecular breeding is expected to mitigate yield loss and quality impairment caused by FSG.However,the genetic control of FSG awaits elucidation.In this study,FSG at 1,3,5,7,and 9 days post-imbibition in three environments were tested,and quantitative-trait loci(QTL)associated with FSG were mapped in a peanut recombinant inbred line population by leveraging existing high-density peanut genetic maps.Of 24 QTL identified in 13 linkage groups,qFSGA04 was a stable major QTL on linkage group 04(LG04).It was consistently detected in five germination stages and three environments.By designing and validating DNA markers in the confidence interval of qFSGA04,we identified one single-nucleotide polymorphism and one In Del closely associated with FSG that could be used as linked markers for FSG resistance in peanut breeding.
基金supported by National Natural Science Foundation of China(31801397)Fund for Distinguished Young Scholars of Henan Academy of Agricultural Sciences(2020JQ03)+3 种基金Independent Innovation Foundation of Henan Academy of Agricultural Sciences,China(2022ZC69)China Agriculture Research System(CARS-13)Key Scientific and Technological Project of Henan Province(201300111000)Henan Provincial Agriculture Research System(S2012-5)。
摘要Oligo probe staining is a low-cost and efficient chromosome identification technique.In this study,oligo genomic in situ hybridization(Oligo-GISH)technology was established in peanut.Peanut A and B subgenome-specific interspersed repeat(IR)oligo probe sets were developed based on clustering and electronic localization of tandem repeat sequences in the reference genome of Tifrunner.The OligoGISH kit was then used to perform staining of 15 Arachis species.The A-subgenome probe set stained the chromosomes of A-and E-genome Arachis species,the B-subgenome probe set stained those of B-,F-,K-,and E-genome species,and neither set stained those of H-genome species.These results indicate the relationships among the genomes of these Arachis species.The Oligo-GISH kit was also used for batch staining of the chromosomes of 389 seedlings from the irradiated M1generation,allowing 67 translocation and deletion lines to be identified.Subsequent Oligo-FISH karyotyping,FISH using single-copy probe libraries,and trait investigation identified seven homozygous chromosomal variants from the M3generation and suggested that there may be genes on chromosome 4B controlling seed number per pod.These findings demonstrate that the IR probe sets and method developed in this study can facilitate research on distant hybridization and genetic improvement in peanut.
基金supported by the Key Research Project of the Shennong Laboratory,China(SN01-2022-03)the Henan Provincial Science and Technology R&D Program Joint Fund(Superiority Discipline Cultivation)Project,China(222301420100)+4 种基金the Major Science and Technology Projects of Henan Province,China(221100110300)the China Agriculture Research System of MOF and MARA(CARS-13)the Henan Provincial Agriculture Research System,China(S2012-5)the Outstanding Young Scientists of Henan Academy of Agricultural Sciences,China(2022YQ16)the Independent Innovation Project of the Henan Academy of Agricultural Sciences,China(2023ZC093)。
摘要Peanut(Arachis hypogaea L.)is a globally important oil crop.Web blotch is one of the most important foliar diseases affecting peanut,which results in serious yield losses worldwide.Breeding web blotch-resistant peanut varieties is the most effective and economically viable method for minimizing yield losses due to web blotch.In the current study,a bulked segregant analysis with next-generation sequencing was used to analyze an F2:3 segregating population and identify candidate loci related to web blotch resistance.Based on the fine-mapping of the candidate genomic interval using kompetitive allele-specific PCR(KASP)markers,we identified a novel web blotch resistance-related locus spanning approximately 169 kb on chromosome 16.This region included four annotated genes,of which only Arahy.35VVQ3 had a non-synonymous single nucleotide polymorphism in the coding region between the two parents.Two markers(Chr.16.12872635 and Chr.16.12966357)linked to this gene were shown to be co-segregated with the resistance of peanut web blotch by 72 randomly selected recombinant inbred lines(RIL),which could be used in marker-assisted breeding of resistant peanut varieties.
基金supported by the National Key Research and Development Program(2023YFD1202800)the Key Research Project of the Shennong Laboratory(SN01-2022-03)+4 种基金the China Agriculture Research System of MOF and MARA(CARS-13)Key Projects of Henan Province(222301420026)Major Science and Technology Projects of Henan Province(221100110300)the Fund for Distinguished Young Scholars from Henan Academy of Agricultural Sciences(2025ZC13,2023JQ02 and 2024JQ06)the Henan Provincial Agriculture Research System,China(S2012-5).
摘要Dear Editor,Cultivated peanut(peanut in short)(Arachis hypogaea,2n=4x=40,genome type AABB)is a legume crop serving globally as a crucial source of edible oil and protein.Available peanut reference genome sequences obtained for the cultivars"Tifrunner,""Shitouqi,"and"Fuhuasheng"were assembled or patched using third-generation sequencing technologies and significantly contributed to advancements in peanut genetics and breeding(Bertioli et al.,2019;Chen et al.,2019;Zhuang et al.,2019).However,unresolved gaps persist in these assemblies,particularly in regions of biological importance such as centromeres and telomeres(Supplemental Figure 1),hindering comprehensive exploration of their functional and evolutionary significance.
基金supported by grants from the National Key Research and Development Program of China (2022YFD1200400)the Key Project of Science and Technology of Henan Province (201300111000, 221100110300)+1 种基金the Earmarked Fund (CARS-13)the Henan Province Agriculture Research System (S2012-5).
摘要Peanut (Arachis hypogaea L.) is an important food and oil crop valued for its oil-rich kernel, high protein content, and diverse nutritional components, making it a staple raw material in the food industry. Fragrance is an essential trait influencing both the initial appeal and flavor quality of food products. For instance, fragrant rice has gained global popularity due to its appealing aroma. The fragrance trait in rice is regulated by the BADH2 gene, which encodes betaine aldehyde dehydrogenase 2 (BADH) (Chen et al., 2008). In non-fragrant rice, BADH2 catalyzes the conversion of γ-aminobutyraldehyde (GABald) to γ-aminobutyric acid. When BADH2 function is disrupted, GABald is redirected to form Δ-1-pyrroline and the subsequent 2-acetyl-1-pyrroline (2-AP), the key compound contributing to rice fragrance (Okpala et al., 2019).