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Article: Overexpression of soybean

Mazarei, Mitra / Routray, Pratyush / Piya, Sarbottam / Stewart, C Neal / Hewezi, Tarek

2023 Volume 14, Page(s) 1186292

Abstract: Soybean ( ...

Abstract	Soybean (
Language	English
Publishing date	2023-05-31
Publishing country	Switzerland
Document type	Journal Article
ZDB-ID	2613694-6
ISSN	1664-462X
ISSN	1664-462X
DOI	10.3389/fpls.2023.1186292
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Article ; Online: miR778 mediates gene expression, histone modification, and DNA methylation during cyst nematode parasitism.

Bennett, Morgan / Piya, Sarbottam / Baum, Thomas J / Hewezi, Tarek

Plant physiology

2022 Volume 189, Issue 4, Page(s) 2432–2453

Abstract: Despite the known critical regulatory functions of microRNAs, histone modifications, and DNA methylation in reprograming plant epigenomes in response to pathogen infection, the molecular mechanisms underlying the tight coordination of these components ... ...

Abstract	Despite the known critical regulatory functions of microRNAs, histone modifications, and DNA methylation in reprograming plant epigenomes in response to pathogen infection, the molecular mechanisms underlying the tight coordination of these components remain poorly understood. Here, we show how Arabidopsis (Arabidopsis thaliana) miR778 coordinately modulates the root transcriptome, histone methylation, and DNA methylation via post-transcriptional regulation of the H3K9 methyltransferases SU(var)3-9 homolog 5 (SUVH5) and SUVH6 upon infection by the beet cyst nematode Heterodera schachtii. miR778 post-transcriptionally silences SUVH5 and SUVH6 upon nematode infection. Manipulation of the expression of miR778 and its two target genes significantly altered plant susceptibility to H. schachtii. RNA-seq analysis revealed a key role of SUVH5 and SUVH6 in reprograming the transcriptome of Arabidopsis roots upon H. schachtii infection. In addition, chromatin immunoprecipitation (ChIP)-seq analysis established SUVH5 and SUVH6 as the main enzymes mediating H3K9me2 deposition in Arabidopsis roots in response to nematode infection. ChIP-seq analysis also showed that these methyltransferases possess distinct DNA binding preferences in that they are targeting transposable elements under noninfected conditions and protein-coding genes in infected plants. Further analyses indicated that H3K9me2 deposition directed by SUVH5 and SUVH6 contributes to gene expression changes both in roots and in nematode feeding sites and preferentially associates with CG DNA methylation. Together, our results uncovered multi-layered epigenetic regulatory mechanisms coordinated by miR778 during Arabidopsis-H. schachtii interactions.
MeSH term(s)	Animals ; Arabidopsis/metabolism ; Arabidopsis Proteins/genetics ; Arabidopsis Proteins/metabolism ; Cysts/genetics ; Cysts/metabolism ; DNA Methylation/genetics ; Gene Expression ; Gene Expression Regulation, Plant ; Histone Code ; Methyltransferases/metabolism ; Plant Diseases/genetics ; Plant Roots/genetics ; Plant Roots/metabolism ; Tylenchoidea
Chemical Substances	Arabidopsis Proteins ; Methyltransferases (EC 2.1.1.-)
Language	English
Publishing date	2022-05-17
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't ; Research Support, U.S. Gov't, Non-P.H.S.
ZDB-ID	208914-2
ISSN	1532-2548 ; 0032-0889
ISSN (online)	1532-2548
ISSN	0032-0889
DOI	10.1093/plphys/kiac228
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Article ; Online: The soybean immune receptor GmBIR1 regulates host transcriptome, spliceome, and immunity during cyst nematode infection.

Hawk, Tracy E / Piya, Sarbottam / Zadegan, Sobhan Bahrami / Li, Peitong / Rice, John H / Hewezi, Tarek

The New phytologist

2023 Volume 239, Issue 6, Page(s) 2335–2352

Abstract: BAK1-INTERACTING RECEPTOR LIKE KINASE1 (BIR1) is a negative regulator of various aspects of disease resistance and immune responses. Here, we investigated the functional role of soybean (Glycine max) BIR1 (GmBIR1) during soybean interaction with soybean ... ...

Abstract	BAK1-INTERACTING RECEPTOR LIKE KINASE1 (BIR1) is a negative regulator of various aspects of disease resistance and immune responses. Here, we investigated the functional role of soybean (Glycine max) BIR1 (GmBIR1) during soybean interaction with soybean cyst nematode (SCN, Heterodera glycines) and the molecular mechanism through which GmBIR1 regulates plant immunity. Overexpression of wild-type variant of GmBIR1 (WT-GmBIR1) using transgenic soybean hairy roots significantly increased soybean susceptibility to SCN, whereas overexpression of kinase-dead variant (KD-GmBIR1) significantly increased plant resistance. Transcriptome analysis revealed that genes oppositely regulated in WT-GmBIR1 and KD-GmBIR1 upon SCN infection were enriched primarily in defense and immunity-related functions. Quantitative phosphoproteomic analysis identified 208 proteins as putative substrates of the GmBIR1 signaling pathway, 114 of which were differentially phosphorylated upon SCN infection. In addition, the phosphoproteomic data pointed to a role of the GmBIR1 signaling pathway in regulating alternative pre-mRNA splicing. Genome-wide analysis of splicing events provided compelling evidence supporting a role of the GmBIR1 signaling pathway in establishing alternative splicing during SCN infection. Our results provide novel mechanistic insights into the function of the GmBIR1 signaling pathway in regulating soybean transcriptome and spliceome via differential phosphorylation of splicing factors and regulation of splicing events of pre-mRNA decay- and spliceosome-related genes.
MeSH term(s)	Animals ; Transcriptome/genetics ; Glycine max/genetics ; Glycine max/metabolism ; Gene Expression Profiling ; Nematode Infections ; Plant Diseases/genetics ; Tylenchoidea/physiology
Language	English
Publishing date	2023-06-20
Publishing country	England
Document type	Journal Article ; Research Support, U.S. Gov't, Non-P.H.S.
ZDB-ID	208885-x
ISSN	1469-8137 ; 0028-646X
ISSN (online)	1469-8137
ISSN	0028-646X
DOI	10.1111/nph.19087
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Article ; Online: The soybean immune receptor GmBIR1 regulates host transcriptome, spliceome, and immunity during cyst nematode infection

Hawk, Tracy E. / Piya, Sarbottam / Zadegan, Sobhan Bahrami / Li, Peitong / Rice, John H. / Hewezi, Tarek

New Phytologist. 2023 Sept., v. 239, no. 6 p.2335-2352

2023

Abstract: BAK1‐INTERACTING RECEPTOR LIKE KINASE1 (BIR1) is a negative regulator of various aspects of disease resistance and immune responses. Here, we investigated the functional role of soybean (Glycine max) BIR1 (GmBIR1) during soybean interaction with soybean ... ...

Abstract	BAK1‐INTERACTING RECEPTOR LIKE KINASE1 (BIR1) is a negative regulator of various aspects of disease resistance and immune responses. Here, we investigated the functional role of soybean (Glycine max) BIR1 (GmBIR1) during soybean interaction with soybean cyst nematode (SCN, Heterodera glycines) and the molecular mechanism through which GmBIR1 regulates plant immunity. Overexpression of wild‐type variant of GmBIR1 (WT‐GmBIR1) using transgenic soybean hairy roots significantly increased soybean susceptibility to SCN, whereas overexpression of kinase‐dead variant (KD‐GmBIR1) significantly increased plant resistance. Transcriptome analysis revealed that genes oppositely regulated in WT‐GmBIR1 and KD‐GmBIR1 upon SCN infection were enriched primarily in defense and immunity‐related functions. Quantitative phosphoproteomic analysis identified 208 proteins as putative substrates of the GmBIR1 signaling pathway, 114 of which were differentially phosphorylated upon SCN infection. In addition, the phosphoproteomic data pointed to a role of the GmBIR1 signaling pathway in regulating alternative pre‐mRNA splicing. Genome‐wide analysis of splicing events provided compelling evidence supporting a role of the GmBIR1 signaling pathway in establishing alternative splicing during SCN infection. Our results provide novel mechanistic insights into the function of the GmBIR1 signaling pathway in regulating soybean transcriptome and spliceome via differential phosphorylation of splicing factors and regulation of splicing events of pre‐mRNA decay‐ and spliceosome‐related genes.
Keywords	Glycine max ; Heterodera glycines ; cyst nematodes ; disease resistance ; genetically modified organisms ; genome-wide association study ; immunity ; immunologic receptors ; nematode infections ; phosphorylation ; proteomics ; soybeans ; transcriptome ; transcriptomics
Language	English
Dates of publication	2023-09
Size	p. 2335-2352.
Publishing place	John Wiley & Sons, Ltd
Document type	Article ; Online
Note	JOURNAL ARTICLE
ZDB-ID	208885-x
ISSN	1469-8137 ; 0028-646X
ISSN (online)	1469-8137
ISSN	0028-646X
DOI	10.1111/nph.19087
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Soybean gene co-expression network analysis identifies two co-regulated gene modules associated with nodule formation and development.

Piya, Sarbottam / Pantalone, Vince / Zadegan, Sobhan Bahrami / Shipp, Sarah / Lakhssassi, Naoufal / Knizia, Dounya / Krishnan, Hari B / Meksem, Khalid / Hewezi, Tarek

Molecular plant pathology

2023 Volume 24, Issue 6, Page(s) 628–636

Abstract: Gene co-expression network analysis is an efficient systems biology approach for the discovery of novel gene functions and trait-associated gene modules. To identify clusters of functionally related genes involved in soybean nodule formation and ... ...

Abstract	Gene co-expression network analysis is an efficient systems biology approach for the discovery of novel gene functions and trait-associated gene modules. To identify clusters of functionally related genes involved in soybean nodule formation and development, we performed a weighted gene co-expression network analysis. Two nodule-specific modules (NSM-1 and NSM-2, containing 304 and 203 genes, respectively) were identified. The NSM-1 gene promoters were significantly enriched in cis-binding elements for ERF, MYB, and C2H2-type zinc transcription factors, whereas NSM-2 gene promoters were enriched in cis-binding elements for TCP, bZIP, and bHLH transcription factors, suggesting a role of these regulatory factors in the transcriptional activation of nodule co-expressed genes. The co-expressed gene modules included genes with potential novel roles in nodulation, including those involved in xylem development, transmembrane transport, the ethylene signalling pathway, cytoskeleton organization, cytokinesis and regulation of the cell cycle, regulation of meristem initiation and growth, transcriptional regulation, DNA methylation, and histone modifications. Functional analysis of two co-expressed genes using TILLING mutants provided novel insight into the involvement of unsaturated fatty acid biosynthesis and folate metabolism in nodule formation and development. The identified gene co-expression modules provide valuable resources for further functional genomics studies to dissect the genetic basis of nodule formation and development in soybean.
MeSH term(s)	Glycine max/genetics ; Gene Regulatory Networks ; Gene Expression Regulation, Plant/genetics ; Gene Expression Profiling ; Transcription Factors/genetics
Chemical Substances	Transcription Factors
Language	English
Publishing date	2023-03-28
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2020755-4
ISSN	1364-3703 ; 1364-3703
ISSN (online)	1364-3703
ISSN	1364-3703
DOI	10.1111/mpp.13327
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Article: Canonical and noncanonical ethylene signaling pathways that regulate Arabidopsis susceptibility to the cyst nematode Heterodera schachtii

Piya, Sarbottam / Brad M. Binder / Tarek Hewezi

new phytologist. 2019 Jan., v. 221, no. 2

2019

Abstract: Plant‐parasitic cyst nematodes successfully exploit various phytohormone signaling pathways to establish a new hormonal equilibrium that facilitates nematode parasitism. Although it is largely accepted that ethylene regulates plant responses to nematode ... ...

Abstract	Plant‐parasitic cyst nematodes successfully exploit various phytohormone signaling pathways to establish a new hormonal equilibrium that facilitates nematode parasitism. Although it is largely accepted that ethylene regulates plant responses to nematode infection, a mechanistic understanding of how ethylene shapes plant–nematode interactions remains largely unknown. In this study, we examined the involvement of various components regulating ethylene perception and signaling in establishing Arabidopsis susceptibility to the cyst nematode Heterodera schachtii using a large set of well‐characterized single and higher order mutants. Our analyses revealed the existence of two pathways that separately engage ethylene with salicylic acid (SA) and cytokinin signaling during plant response to nematode infection. One pathway involves the canonical ethylene signaling pathway in which activation of ethylene signaling results in suppression of SA‐based immunity. The second pathway involves the ethylene receptor ETR1, which signals independently of SA acid to affect immunity, instead altering cytokinin‐mediated regulation of downstream components. Our results reveal important mechanisms through which cyst nematodes exploit components of ethylene perception and signaling to affect the balance of hormonal signaling through ethylene interaction with SA and cytokinin networks. This hormonal interaction overcomes plant defense and provokes a susceptible response.
Keywords	Arabidopsis ; cyst nematodes ; cytokinins ; ethylene ; Heterodera schachtii ; immunity ; mutants ; nematode infections ; parasitism ; plant response ; salicylic acid ; signal transduction
Language	English
Dates of publication	2019-01
Size	p. 946-959.
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	208885-x
ISSN	1469-8137 ; 0028-646X
ISSN (online)	1469-8137
ISSN	0028-646X
DOI	10.1111/nph.15400
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Soybean gene co‐expression network analysis identifies two co‐regulated gene modules associated with nodule formation and development

Piya, Sarbottam / Pantalone, Vince / Zadegan, Sobhan Bahrami / Shipp, Sarah / Lakhssassi, Naoufal / Knizia, Dounya / Krishnan, Hari B. / Meksem, Khalid / Hewezi, Tarek

Molecular Plant Pathology. 2023 Mar. 28, v. 24, no. 6 p.628-636

2023

Abstract	Gene co-expression network analysis is an efficient systems biology approach for the discovery of novel gene functions and trait-associated gene modules. To identify clusters of functionally related genes involved in soybean nodule formation and development, we performed a weighted gene co-expression network analysis. Two nodule-specific modules (NSM-1 and NSM-2, containing 304 and 203 genes, respectively) were identified. The NSM-1 gene promoters were significantly enriched in cis-binding elements for ERF, MYB, and C2H2-type zinc transcription factors, whereas NSM-2 gene promoters were enriched in cis-binding elements for TCP, bZIP, and bHLH transcription factors, suggesting a role of these regulatory factors in the transcriptional activation of nodule co-expressed genes. The co-expressed gene modules included genes with potential novel roles in nodulation, including those involved in xylem development, transmembrane transport, the ethylene signalling pathway, cytoskeleton organization, cytokinesis and regulation of the cell cycle, regulation of meristem initiation and growth, transcriptional regulation, DNA methylation, and histone modifications. Functional analysis of two co-expressed genes using TILLING mutants provided novel insight into the involvement of unsaturated fatty acid biosynthesis and folate metabolism in nodule formation and development. The identified gene co-expression modules provide valuable resources for further functional genomics studies to dissect the gene
Keywords	DNA methylation ; biosynthesis ; cytokinesis ; cytoskeleton ; ethylene ; genes ; genomics ; histones ; meristems ; nodulation ; plant pathology ; soybeans ; transcription (genetics) ; transcriptional activation ; unsaturated fatty acids ; vitamin metabolism ; xylem ; zinc
Language	English
Dates of publication	2023-0328
Size	p. 628-636.
Publishing place	Wiley
Document type	Article ; Online
Note	JOURNAL ARTICLE ; Resource is Open Access
ZDB-ID	2020755-4
ISSN	1364-3703 ; 1464-6722
ISSN (online)	1364-3703
ISSN	1464-6722
DOI	10.1111/mpp.13327
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Article: Kinase‐dead mutation: A novel strategy for improving soybean resistance to soybean cyst nematode Heterodera glycines

Piya, Sarbottam / Hawk, Tracy / Patel, Bhoomi / Baldwin, Logan / Rice, John H. / Stewart, C. Neal, Jr / Hewezi, Tarek

Molecular plant pathology. 2022 Mar., v. 23, no. 3

2022

Abstract: Protein kinases phosphorylate proteins for functional changes and are involved in nearly all cellular processes, thereby regulating almost all aspects of plant growth and development, and responses to biotic and abiotic stresses. We generated two ... ...

Abstract	Protein kinases phosphorylate proteins for functional changes and are involved in nearly all cellular processes, thereby regulating almost all aspects of plant growth and development, and responses to biotic and abiotic stresses. We generated two independent co‐expression networks of soybean genes using control and stress response gene expression data and identified 392 differentially highly interconnected kinase hub genes among the two networks. Of these 392 kinases, 90 genes were identified as “syncytium highly connected hubs”, potentially essential for activating kinase signalling pathways in the nematode feeding site. Overexpression of wild‐type coding sequences of five syncytium highly connected kinase hub genes using transgenic soybean hairy roots enhanced plant susceptibility to soybean cyst nematode (SCN; Heterodera glycines) Hg Type 0 (race 3). In contrast, overexpression of kinase‐dead variants of these five syncytium kinase hub genes significantly enhanced soybean resistance to SCN. Additionally, three of the five tested kinase hub genes enhanced soybean resistance to SCN Hg Type 1.2.5.7 (race 2), highlighting the potential of the kinase‐dead approach to generate effective and durable resistance against a wide range of SCN Hg types. Subcellular localization analysis revealed that kinase‐dead mutations do not alter protein cellular localization, confirming the structure–function of the kinase‐inactive variants in producing loss‐of‐function phenotypes causing significant decrease in nematode susceptibility. Because many protein kinases are highly conserved and are involved in plant responses to various biotic and abiotic stresses, our approach of identifying kinase hub genes and their inactivation using kinase‐dead mutation could be translated for biotic and abiotic stress tolerance.
Keywords	Heterodera glycines ; abiotic stress ; giant cells ; growth and development ; loss-of-function mutation ; plant growth ; plant pathology ; protein kinases ; soybeans ; stress response ; stress tolerance
Language	English
Dates of publication	2022-03
Size	p. 417-430.
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	2020755-4
ISSN	1364-3703 ; 1464-6722
ISSN (online)	1364-3703
ISSN	1464-6722
DOI	10.1111/mpp.13168
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Article ; Online: Soybean MKK2 establishes intricate signalling pathways to regulate soybean response to cyst nematode infection.

Hawk, Tracy E / Piya, Sarbottam / Sultana, Mst Shamira / Zadegan, Sobhan Bahrami / Shipp, Sarah / Coffey, Nicole / McBride, Natalie B / Rice, John H / Hewezi, Tarek

Molecular plant pathology

2024 Volume 25, Issue 5, Page(s) e13461

Abstract: Mitogen-activated protein kinase (MPK) cascades play central signalling roles in plant immunity and stress response. The soybean orthologue of MPK kinase2 (GmMKK2) was recently identified as a potential signalling node whose expression is upregulated in ... ...

Abstract	Mitogen-activated protein kinase (MPK) cascades play central signalling roles in plant immunity and stress response. The soybean orthologue of MPK kinase2 (GmMKK2) was recently identified as a potential signalling node whose expression is upregulated in the feeding site induced by soybean cyst nematode (SCN, Heterodera glycines). To investigate the role of GmMKK2 in soybean-SCN interactions, we overexpressed a catabolically inactive variant referred to as kinase-dead variant (KD-GmMKK2) using transgenic hairy roots. KD-GmMKK2 overexpression caused significant reduction in soybean susceptibility to SCN, while overexpression of the wild-type variant (WT-GmMKK2) exhibited no effect on susceptibility. Transcriptome analysis indicated that KD-GmMKK2 overexpressing plants are primed for SCN resistance via constitutive activation of defence signalling, particularly those related to chitin, respiratory burst, hydrogen peroxide and salicylic acid. Phosphoproteomic profiling of the WT-GmMKK2 and KD-GmMKK2 root samples upon SCN infection resulted in the identification of 391 potential targets of GmMKK2. These targets are involved in a broad range of biological processes, including defence signalling, vesicle fusion, chromatin remodelling and nuclear organization among others. Furthermore, GmMKK2 mediates phosphorylation of numerous transcriptional and translational regulators, pointing to the presence of signalling shortcuts besides the canonical MAPK cascades to initiate downstream signalling that eventually regulates gene expression and translation initiation. Finally, the functional requirement of specific phosphorylation sites for soybean response to SCN infection was validated by overexpressing phospho-mimic and phospho-dead variants of two differentially phosphorylated proteins SUN1 and IDD4. Together, our analyses identify GmMKK2 impacts on signalling modules that regulate soybean response to SCN infection.
MeSH term(s)	Glycine max/parasitology ; Glycine max/genetics ; Animals ; Signal Transduction ; Plant Diseases/parasitology ; Plant Diseases/genetics ; Tylenchoidea/physiology ; Tylenchoidea/pathogenicity ; Gene Expression Regulation, Plant ; Plants, Genetically Modified ; Plant Roots/parasitology ; Plant Roots/metabolism ; Plant Roots/genetics ; Plant Proteins/metabolism ; Plant Proteins/genetics ; Disease Resistance/genetics
Chemical Substances	Plant Proteins
Language	English
Publishing date	2024-05-02
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2020755-4
ISSN	1364-3703 ; 1364-3703
ISSN (online)	1364-3703
ISSN	1364-3703
DOI	10.1111/mpp.13461
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Article: Soil inoculation with Trichoderma asperellum, T. harzianum or Streptomyces griseoviridis prior to anaerobic soil disinfestation (ASD) does not increase ASD efficacy against Sclerotium rolfsii germination

Shrestha, Utsala / Dee, Mary E / Piya, Sarbottam / Ownley, Bonnie H / Butler, David M

Applied soil ecology. 2020 Mar., v. 147

2020

Abstract: Studies on anaerobic soil disinfestation (ASD), a non-chemical alternative to soil fumigants for controlling plant diseases caused by soilborne pathogens, have shown that ASD increases populations of beneficial microorganisms with activity against plant ... ...

Abstract	Studies on anaerobic soil disinfestation (ASD), a non-chemical alternative to soil fumigants for controlling plant diseases caused by soilborne pathogens, have shown that ASD increases populations of beneficial microorganisms with activity against plant pathogens, including species of Trichoderma and Streptomyces, which are known to parasitize sclerotia of Sclerotium rolfsii. However, studies on ASD effectiveness paired with inoculation of Trichoderma and Streptomyces are lacking. This study compared the effects of ASD, soil incorporation of antagonists [Trichoderma asperellum, T. harzianum (RootShield®), Streptomyces griseoviridis (Mycostop®) and T. harzianum + S. griseoviridis], and ASD + soil incorporation of antagonists at the initiation of ASD treatment, on sclerotial germination and colonization. The effect of ASD and soil incorporation of antagonists on abundance of Trichoderma spp., actinomycetes, and Bacillus spp. in soil were assessed. In contrast to the negative effect of ASD on sclerotial germination, we observed positive or no effect of ASD alone on populations of beneficial microorganisms evaluated. ASD treatment that included addition of T. asperellum or T. harzianum did not increase colonization of sclerotia by Trichoderma spp. compared to ASD alone, whereas ASD treatment with inoculation of S. griseoviridis did increase sclerotia colonization by actinomycetes. Indigenous soil actinomycete populations were suppressed by incorporation of T. asperellum and T. harzianum during ASD treatment. Soil abundance of T. harzianum was increased by ASD treatment, and further increased by T. harzianum inoculation at ASD treatment initiation. Inoculation of T. asperellum, T. harzianum, or S. griseoviridis at ASD treatment did not further reduce germination of sclerotia of S. rolfsii, indicating that addition of these microorganisms to soil prior to ASD treatment is unlikely to improve ASD treatment efficacy against pathogens with resistant sclerotial survival structures.
Keywords	Athelia rolfsii ; Bacillus (bacteria) ; Streptomyces griseoviridis ; Trichoderma asperellum ; disinfestation ; germination ; sclerotia ; soil ; soil ecology ; soil inoculation
Language	English
Dates of publication	2020-03
Publishing place	Elsevier B.V.
Document type	Article
Note	NAL-AP-2-clean
ZDB-ID	1196758-4
ISSN	0929-1393
ISSN	0929-1393
DOI	10.1016/j.apsoil.2019.103383
Database	NAL-Catalogue (AGRICOLA)

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