Rice Aroma: A Natural Gift Comes with Price and the Way Forward

doi:10.1016/j.rsci.2020.01.001

摘要/Abstract

. [J]. Rice Science, 2020, 27(2): 86-100.

Fig. 1. Integration of previously reported QTLs for aroma with respective candidate genes in rice. I, Luriex et al (1996); II, Singh et al (2007); III, Amaranthi et al (2008); IV, Pachuri et al (2014); V, Talukder et al (2017); Chr, Chromosome; Mb, Mega base; cM, Centimorgan.The map has been constructed using the Chromosome Map Tool of Oryzabase from http://viewer.shigen.info/oryzavw/maptool/MapTool.do; http://archive.gramene.org/qtl/ and http://archive.gramene.org/markers/ (information retrieved on 20 May, 2019).

Table 1 Description of some potential genes related to aroma in rice.

Gene	ID	Tissue specificity (Expression)	Response to stress	Splicing	Subcellular localization	Protein interaction
P5CS gene
OsP5CS1	Os05g0455500 LOC_Os05g38150	Flower buds, milk grains	Osmoregulation, salinity, anoxia	Introns 3, 19; Termination	Chl, ER, Nu, Cyt, Mit, Pl, Extr, Vac	Ferredoxin-dependent glutamate synthase
OsP5CS2	Os01g0848200 LOC_Os01g62900	Flowers, flower buds	Osmoregulation, salinity, anoxia	All introns; Termination	Chl, Extr, Nu, Vac	Glutamate synthase
GLY gene
OsGlyI	Os05g0295800 LOC_Os05g22970	Leaves before flowering, flower buds	Salinity, anoxia	‒	Chl, Cyt	‒
OsGlyII	Os03g0332400 LOC_Os03g21460	Flower buds, flower	Salinity, anoxia	‒	Cyt, Chl, Nu, Extr	Glyoxalase
OsGlyIII	Os01g0667200 LOC_Os01g47690	Roots and leaves before flowering	Salinity, anoxia	Introns 7, 8	Chl, Per, Cyt, Golg	Ferredoxin-nitrite reductase
BADH gene
OsBadh1	Os04g0464200 LOC_Os04g39020	Roots before flowering, flowers	Salinity, anoxia, submergence	Intron 4	Per, Chl, Cyt, Nu	Glutamate synthase
OsBadh2	Os08g0424500 LOC_Os08g32870	Flowers, flower buds	Salinity, anoxia, submergence	‒	Chl, Per, Cyt, Nu	Glutamate synthase

Fig. 2. Haplotypes of OsBadh1 gene in rice based on 15 single nucleotide polymorphisms (SNPs) with no missing data genotyped using Sequenom MassARRAY system. Protein haplotypes are based on three exonic SNPs (S6, S18 and S19). It was redrawn from Singh et al (2010).

Fig. 3. Comprehensive 2-acetyl-1-phrroline (2AP) pathway related to other pathways in aromatic rice plant cells. 1,3-BPG, 1,3-biphosphoglycerate; ArgD, Arginine decarboxylase; ArgU, Argamatine ureodehydrolase; Asp, Aspartic acid; Asp-AT, Aspartate aminotransferase; CaM, Calmodulin; CAT9, Cationic amino acid transporter 9; DAO, Diamine oxidase; DhAP, Dihydroxyacetone phosphate; Fru-1,6BP, Fructose-1,6-biphosphate; GABA, γ-aminobutyric acid; GAB-ald, γ-aminobutyraldehyde; GABA-P, γ-aminobutyric acid permease; GABA-TK, α-ketoglutarate dependent GABA transaminase; GABA-TP, Pyruvate dependent GABA transaminase; GAD, Glutamate decarboxylase; GAP-A,B,C, Glyceraldehyde-3-phosphate dehydrogenase homologues (A,B,C); GHB, Gamma hydroxybutyrate; Glu, Glutamate; Gly3P, Glyceraldehyde-3-phosphate; GSA, Gamma glutamyl semialdehyde; GSAR, Gamma glutamyl semialdehyde reductase; Meth, Methylglyoxal; MGD, Methylglyoxal dehydrogenase; OAT, Ornithine aminotransferase; OCD, Ornithine cyclodeaminase; ODC, Ornithine decarboxylase; P5C, Δ1-pyrroline-5-carboxylate; P5CS, Δ1-pyrroline-5-carboxylate synthetase; P5CR, Δ1-pyrroline-5-carboxylate reductase; PAO, Polyamine oxidase; PLP, Pyridoxal 5′-phosphate; PRODH, Proline dehydrogenase; SPDS, Spermidine synthase; SSA, Succinic semialdehyde; SSADH, Succinic semialdehyde dehydrogenase; SSAR, Succinic semialdehyde reductase; TCA, Tricarboxylic acid; TPI, Triosephosphate isomerase; αKT, α-ketoglutarate; αKTDH, α-ketoglutarate dehydrogenase; γ-GK, γ-glutamyl kinase; Δ1PDH, Δ1-pyrroline dehydrogenase; Δ1-pyrr, Δ1-pyrroline.

Table 2 Variation in 2-acetyl-1-phrroline (2AP) concentration under different environmental conditions.

Treatment/Environment	Variety	2AP concentration (µg/kg)	Reference
Shading			Mo et al, 2015
Control	Yuxiangyouzhan	85.10
Treated	Yuxiangyouzhan	175.86
Control	Nongxiang 18	94.84
Treated	Nongxiang 18	135.02
Salinity			Gay et al, 2010
2006	Aychade	715.00
2007	Aychade	1 440.00
2006	Fidji	560.00
2007	Fidji	1 715.00
2006	Giano	301.00
2007	Giano	741.00
Growing condition			Yoshihashi et al, 2004
Sandy soil, dry during ripening	KDML105	532.00
Submerged during ripening	KDML105	218.00
Clay soil	KDML105	388.00
Growth stage			Hinge et al, 2016
Vegetative stage	AM157	310.00
Reproductive stage	AM157	400.00
Grain filling stage	AM157	580.00
Mature grain stage	AM157	662.00
Vegetative stage	BA370	380.00
Reproductive stage	BA370	420.00
Grain filling stage	BA370	260.00
Mature grain stage	BA370	451.00

Fig. 4. Roles of GABA and proline in plant stress responses and environmental changes. 2AP, 2-acetyl-1-phrroline; ACS, 1-aminocyclopropane-1-carboxylic acid synthase; ACO, 1-aminocyclopropane-1-carboxylic acid oxidase; CaM, Calmodulin; GABA, γ-aminobutyric acid; GAD, Glutamate decarboxylase; Glu, Glutamate; P5CS, Δ1-pyrroline-5-carboxylate synthetase; PDH, Proline dehydrogenase.GABA and proline may function as a cellular barometer and transducer of environmental stress signals (Modified from Kinnersley and Turano, 2000).

Fig. 5. Regulation of aroma related compounds during normal or optimum environmental condition.2AP, 2-acetyl-1-pyrroline; GABA, γ-aminobutyric acid; GAB-ald, Gamma aminobutyraldehyde; P5C, Δ1-pyrroline-5-carboxylate; P5CR, Δ1-pyrroline-5-carboxylate reductase; PRODH, Proline dehydrogenase; TCA, Tricarboxylic acid cycle; α-KTDH, α-ketoglutarate dehydrogenase.

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