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. 2018 Aug 14;9:1089. doi: 10.3389/fpls.2018.01089

Table 2.

Genes involved in the biosynthesis of health-promoting secondary metabolites, which are upregulated in above- or belowground cell tissues of mycorrhizal food plants.

Plant Effective AMF Plant tissue Assay method Upregulated gene/enzyme family Involved pathway References
Citrus sinensis Glomus versiforme (BGC HUN02B) Leaves RNA-Seq 12-oxophytodienoate reductase
Acyl-CoA oxidase
Enoyl-CoA hydratase
Jasmonate O-methyltransferase
Linoleate 13S-lipoxygenase
OPC-8:CoA ligase
Secondary metabolites biosynthesis Gao et al., 2016
Cucumis sativus Funneliformis mosseae Leaves RT-qPCR Caffeoyl CoA 3-O-methyltransferase
Cinnamyl alcohol dehydrogenase-like protein
Cytochrome P450 C4H cinnamate-4-hydroxylase
Phenylalanine ammonia-lyase
Phenylpropanoid biosynthesis Chen et al., 2013
Cucumis sativus Rhizophagus irregularis (PH5, formerly Glomus intraradices) Roots RNA-Seq Allene oxide synthase 1

Cinnamyl alcohol dehydrogenase (6-like)
Cytochrome P450 (734A6-like)
Gibberellin 2-beta-dioxygenase
Secondary metabolites biosynthesis
Phenylpropanoid biosynthesis

Phenylpropanoid biosynthesis
Diterpenoid biosynthesis
Ma et al., 2018
Glycine max Rhizophagus irregularis (49, formerly Glomus intraradices) Roots Microarray 9-cis-epoxycarotenoid dioxygenase
Cytochrome P450 monooxygenase (CYP 701)
Geranylgeranyl diphosphate or pyrophosphate synthase
Gibberellin 2-beta-dioxygenase
Gibberellin 3-beta (20)-dioxygenase
Isoflavone-O-methyltransferase
Trihydroxyisoflavanone-O-methyltransferase
Tropinone reductase
Carotenoid biosynthesis

Terpenoid backbone biosynthesis
Terpenoid backbone biosynthesis
Diterpenoid biosynthesis
Diterpenoid biosynthesis

Isoflavonoid biosynthesis
Isoflavonoid biosynthesis

Alkaloids biosynthesis
Schaarschmidt et al., 2013
Cytochrome p450 (CYP93A-like)
Ent-kaurene oxidase cyp701a5
Flavanone 3-hydroxylase
Flavonoid glucosyltransferase
Flavonoid peroxidase 1
Geraniol 8-hydroxylase-like
Gibberellin 2-beta-dioxygenase
Gibberellin 3-beta (20)-dioxygenase
Hydroxycinnamoyl transferase
Hyoscyamine 6-dioxygenase
Isoleucine N-monooxygenase
Pelargonidin 3-o-(6-caffeoylglucoside) 5-o-(6-o-malonylglucoside) 4-malonyltransferase-like
Shikimate o-hydroxycinnamoyltransferase-likeTetrahydrocannabinolic acid synthase-like
Tropinone reductase homolog
Valine N-monooxygenase (CYP79D1-2)
Zeatin-O-xylosyltransferase-like
Isoflavonoid biosynthesis
Diterpenoid biosynthesis
Flavonoid biosynthesis
Flavonoid biosynthesis
Flavonoid biosynthesis
Monoterpenoid biosynthesis
Diterpenoid biosynthesis
Diterpenoid biosynthesis

Phenylpropanoid biosynthesis

Alkaloid biosynthesis Glucosinolate biosynthesis
Anthocyanin biosynthesis

Flavonoid biosynthesis


Cannabinoid biosynthesis

Alkaloids biosynthesis
Glucosinolate biosynthesis

Secondary metabolites biosynthesis
Vangelisti et al., 2018
Litchi chinensis native AMF community Roots RNA-Seq Anthocyanidin reductase
Bifunctional dihydroflavonol 4-reductase/flavanone 4-reductase
Caffeic acid 3-O-methyltransferase
Carotenoid cleavage dioxygenase 7
Chalcone synthase
Coumarate-CoA ligase 2
Cytochrome P450 (CYP73A)
Flavanone 3-dioxygenase
Flavonoid 3′-monooxygenase

Leucoanthocyanidin reductase
Naringenin,2-oxoglutarate 3-dioxygenase
Peroxidase 53
Tropinone reductase
Flavonoid biosynthesis
Flavonoid biosynthesis


Phenylpropanoid biosynthesis

Carotenoid biosynthesis

Flavonoid biosynthesis
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis
Flavonoid biosynthesis
Flavonoid, flavone, flavonol biosynthesis
Flavonoid biosynthesis
Flavonoid biosynthesis

Phenylpropanoid biosynthesis
Alkaloid biosynthesis
Shu et al., 2016
Ocimum basilicum Rhizoglomus irregulare (IMA6, formerly Glomus intraradices) Leaves RT-qPCR Tyrosine amino-transferase Tyrosine metabolism Battini et al., 2016a
Oryza sativa Glomus intraradices (DAOM197198) Leaves Macroarray Ascorbate peroxidase (APX8)

Cytochrome P450
Dehydroascorbate reductase

Squalene monooxygenase
Ascorbate and aldarate metabolism
Phenylpropanoid biosynthesis
Ascorbate and aldarate metabolism
Secondary metabolites biosynthesis
Campos-Soriano et al., 2012
Rhizophagus irregularis (DAOM197198) Roots RNA-Seq Anthocyanidin 3-O-glucosyltransferase
Cinnamoyl-CoA reductase
cytochrome P450
Ent-kaurene synthase
Flavonol synthase/flavanone 3-hydroxylase
Laccase precursor protein

Mannose-6-phosphate isomerase
Oxidoreductase, aldo/keto reductase family protein
Phytoene synthase
Terpene synthase
Anthocyanin biosynthesis

Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis
Diterpenoid biosynthesis
Flavonoid biosynthesis

Ascorbate and aldarate metabolism
Ascorbate and aldarate metabolism
Retinoic acid biosynthesis

Carotenoid biosynthesis
Diterpenoid biosynthesis
Fiorilli et al., 2015
Glomus intraradices Roots Microarray Cinnamoyl-CoA reductase
4-coumarate CoA ligase
Cycloartenol synthase

Cytochrome p450
Flavonoid 3′,5′-hydroxylase
Geranylgeranyl diphosphate synthase
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis
Secondary metabolites biosynthesis
Phenylpropanoid biosynthesis
Flavonoid biosynthesis
Terpenoid backbone biosynthesis
Güimil et al., 2005
Rhizophagus irregularis Roots Microarray 4-coumarate-CoA ligase
Caffeoyl CoA 3-O-methyltransferase
Gibberellin 2-beta-dioxygenase
p-coumaroyl-CoA:caffeoyl-CoA 3-hydroxylase
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis

Diterpenoid biosynthesis
Phenylpropanoid biosynthesis
Gutjahr et al., 2015
Pisum sativum Glomus mosseae (BB-E-Sc-02; Biorize, Dijon, FR) Roots SSH Beta-cyanoalanine synthase

Neoxanthin (clavage enzyme) synthase
Secondary metabolites biosynthesis
Carotenoid biosynthesis
Grunwald et al., 2004
Solanum lycopersicum Rhizophagus irregularis Leaves RNA-Seq 4-coumarate CoA ligase
Acyltransferase-like protein
Caffeoyl CoA 3-O-methyltransferase
Cinnamoyl CoA-reductase-like
Cytochrome P450 NADPH-reductase
Hydroxycinnamoyl transferase
Hydroxycinnamoyl-CoA shikimate/quinate hydroxycinnamoyltransferase
Mevalonate kinase

Phenylalanine ammonia-lyase
Undecaprenyl pyrophosphate synthase
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis

Flavonoid biosynthesis
Phenylpropanoid biosynthesis

Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis


Terpenoid backbone biosynthesis
Phenylpropanoid biosynthesis
Terpenoid backbone biosynthesis
Cervantes-Gámez et al., 2016
Glomus intraradices (BEG141) Roots Microarray Carotenoid cleavage dioxygenase
Cytochrome P450
Mevalonate disphosphate decarboxylase
Phytoene dehydrogenase
Carotenoid biosynthesis

Phenylpropanoid biosynthesis
Carotenoid biosynthesis

Carotenoid biosynthesis
Dermatsev et al., 2010
Glomus mosseae (BEG12) Roots RNA-Seq (−)-a-terpineol synthase

(+)-delta-cadinene synthase

4-coumarate CoA ligase
Anthocyanidin synthase
Carotenoid cleavage dioxygenase 1
Cytochrome P450
Cytochrome p450 monooxygenase cyp72a59
Monoterpene glucosyltransferase
Phenylalanine ammonia-lyase
Phytoene synthase
Sesquiterpene synthase
Terpenoid backbone biosynthesis
Terpenoid backbone biosynthesis
Phenylpropanoid biosynthesis
Flavonoid biosynthesis
Carotenoid biosynthesis

Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis

Terpenoid backbone biosynthesis
Phenylpropanoid biosynthesis
Carotenoid biosynthesis
Terpenoid backbone biosynthesis
Fiorilli et al., 2009
Glomus mosseae (BEG12) Shoots RNA-Seq Caffeic acid 3-O-methyltransferase
Cytochrome p450
Tropinone reductase
Phenylpropanoid biosynthesis

Phenylpropanoid biosynthesis
Alkaloid biosynthesis
Glomus irregulare (DAOM 197198) Roots Microarray Allene oxide synthase 3

Gibberellin 3-beta-dioxygenase
Phytoene desaturase
Secondary metabolites biosynthesis
Carotenoid biosynthesis
Carotenoid biosynthesis
Garrido et al., 2010
Glomus intraradices Roots Microarray Deoxy-D-xylulose-5-phosphate reductoisomerase
Gibberellin 2-beta-dioxygenase
Hydroxycinnamoyl/benzoyl transferase
Lycopene b-cyclase
Phytoene desaturase
z-carotene desaturase
Terpenoid backbone biosynthesis
Diterpenoid biosynthesis
Phenylpropanoid biosynthesis

Carotenoid biosynthesis
Carotenoid biosynthesis
Carotenoid biosynthesis
López-Ráez et al., 2010b
Glomus intraradices/Glomus mosseae Roots Microarray Allene oxide synthase 1-3

Antocyanin acyltrasferase
Cytochrome P450 monooxygenase (CYP 81, CYP721)
1-deoxy-D-xylulose 5-phosphate synthase
Gibberellin 3-beta (20)-dioxygenase
Secondary metabolites biosynthesis
Anthocyanidin biosynthesis
Terpenoid backbone biosynthesis

Terpenoid backbone biosynthesis
Diterpenoid biosynthesis
Glomus mosseae Roots Microarray Polyphenol oxidase Secondary metabolites biosynthesis
Glomus mosseae Fruit Microarray Histidine decarboxylase Secondary metabolites biosynthesis Salvioli et al., 2012
Rhizophagus irregularis (DAOM 197198) Roots RNA-Seq Carotenoid cleavage dioxygenase 8
Cytochrome P450 (CYP93A1)
Carotenoid biosynthesis

Flavonoid biosynthesis
Sugimura and Saito, 2017
Funneliformis mosseae (BEG12) Fruit RNA-Seq Terpene synthase Diterpenoid biosynthesis Zouari et al., 2014
Solanum tuberosum Glomus sp. MUCL 41833 Roots Microarray Anthocyanidin-3-glucoside rhamnosyltransferase
Anthranilate N-hydroxycinnamoyl/benzoyltransferase
Benzyl alcohol benzoyl transferase
Carotenoid isomerase
Catechol oxidase
Chalcone reductase
Coumarate-CoA ligase (2-4)
Cytochrome p450 (CYP71-like)
Flavanone 3-hydroxylase
Flavonoid 1-2 rhamnosyltransferase
Flavonoid 3′-monooxygenase

Geranylgeranyl pyrophosphate synthase 1
Gibberellin 2-beta-dioxygenase
Hydroxycinnamoyl transferase
Leucoanthocyanidin dioxygenase-like
Orcinol O-methyltransferase

Peroxidase (Class III)
Phenylalanine ammonia-lyase
Phytoene desaturase
Hyoscyamine 6 beta-hydroxylase-likeSesquiterpene synthase 2

Tropinone reductase I
Tyramine hydroxycinnamoyl transferase
Vetispiradiene synthase

z-carotene desaturase
Anthocyanin biosynthesis

Secondary metabolites biosynthesis

Phenylpropanoid biosynthesis

Carotenoid biosynthesisIsoquinoline alkaloid biosynthesisFlavonoid biosynthesis
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis
Flavonoid biosynthesis
Flavonoid biosynthesis

Flavonoid, flavone, flavonol biosynthesis
Terpenoid backbone biosynthesis
Diterpenoid biosynthesis
Phenylpropanoid biosynthesis
Anthocyanidin biosynthesis

Secondary metabolites biosynthesis
Phenylpropanoid biosynthesis
Phenylpropanoid biosynthesis
Carotenoid biosynthesis
Alkaloid biosynthesis

Terpenoid backbone biosynthesis
Alkaloids biosynthesis
Phenylpropanoid biosynthesis

Terpenoid backbone biosynthesis
Carotenoid biosynthesis
Gallou et al., 2012
Vitis vinifera Funneliformis mosseae (BEG12) Roots RNA-Seq ABA 8′-hydroxylase CYP707A1
Alliin lyase precursor
Carotenoid biosynthesis
Secondary metabolites biosynthesis
Balestrini et al., 2017
Funneliformis mosseae (BEG12)/MICOSAT F® Vite (CCS-Aosta) Roots RNA-Seq 4-coumarate-CoA ligase
ABA 8'-hydroxylase CYP707A1
Cinnamyl alcohol dehydrogenaseTaxane 10-beta-hydroxylase
Tropinone reductase
Urophorphyrin III methylase
Terpenoid-quinone biosynthesis
Carotenoid biosynthesis
Phenylpropanoid biosynthesis
Carotenoid biosynthesis
Alkaloids biosynthesis
Secondary metabolites biosynthesis
MICOSAT F® Vite (CCS-Aosta) Roots RNA-Seq Methyl jasmonate esterase Secondary metabolites biosynthesis
Zea mays Rhizophagus irregularis Leaves Microarray 1-aminocyclopropane-1-carboxylate oxidase
3-hydroxy-3-methylglutaryl- reductase
Alliin lyase
Cysteine and methionine metabolism
Terpenoid backbone biosynthesis
Secondary metabolites biosynthesis
Gerlach et al., 2015
Anthocyanidin 3-O-glucosyltransferase 2
Beta-carotene hydroxylase 1
Cytochrome p450 71a1
Farnesyl pyrophosphate synthetase
Gibberellin 2-beta-dioxygenase
Homogentisate phytyltransferase vte2-1
Mevalonate kinase

Short-chain dehydrogenase reductase family protein
Anthocyanin biosynthesis

Carotenoid biosynthesis
Phenylpropanoid biosynthesis
Terpenoid backbone biosynthesis
Diterpenoid biosynthesis
Terpenoid-quinone biosynthesis

Terpenoid backbone biosynthesis
Retinoic acid biosynthesis

The binomial nomenclature of arbuscular mycorrhizal fungi (AMF) reported in the quoted papers has been maintained. Assay methods: RNA sequencing (RNA-Seq), quantitative reverse transcription PCR (RT-qPCR), suppression subtractive hybridization (SSH).