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Review

Advances in Understanding the Genetic Basis of Fatty Acids Biosynthesis in Perilla: An Update

by
Seon-Hwa Bae
1,
Yedomon Ange Bovys Zoclanclounon
2,
Thamilarasan Senthil Kumar
2,
Jae-Hyeon Oh
3,
Jundae Lee
1,
Tae-Ho Kim
2 and
Ki Young Park
4,*
1
Department of Horticulture, Institute of Agricultural Science & Technology, Jeonbuk National University, Jeonju 54896, Korea
2
Genomics Division, National Institute of Agricultural Sciences, Rural Development Administration, Jeonju 54874, Korea
3
R&D Coordination Division, Rural Development Administration, Jeonju 54875, Korea
4
Department of Practical Arts Education, Gongju National University of Education, Gonju 32553, Korea
*
Author to whom correspondence should be addressed.
Plants 2022, 11(9), 1207; https://doi.org/10.3390/plants11091207
Submission received: 8 March 2022 / Revised: 20 April 2022 / Accepted: 25 April 2022 / Published: 29 April 2022
(This article belongs to the Special Issue Lipid Genes and Biotechnology in Plants)
Browse Figure
Review Reports Versions Notes

Abstract

:
Perilla, also termed as purple mint, Chinese basil, or Perilla mint, is a flavoring herb widely used in East Asia. Both crude oil and essential oil are employed for consumption as well as industrial purposes. Fatty acids (FAs) biosynthesis and oil body assemblies in Perilla have been extensively investigated over the last three decades. Recent advances have been made in order to reveal the enzymes involved in the fatty acid biosynthesis in Perilla. Among those fatty acids, alpha-linolenic acid retained the attention of scientists mainly due to its medicinal and nutraceutical properties. Lipids synthesis in Perilla exhibited similarities with Arabidopsis thaliana lipids’ pathway. The homologous coding genes for polyunsaturated fatty acid desaturases, transcription factors, and major acyl-related enzymes have been found in Perilla via de novo transcriptome profiling, genome-wide association study, and in silico whole-genome screening. The identified genes covered de novo fatty acid synthesis, acyl-CoA dependent Kennedy pathway, acyl-CoA independent pathway, Triacylglycerols (TAGs) assembly, and acyl editing of phosphatidylcholine. In addition to the enzymes, transcription factors including WRINKLED, FUSCA3, LEAFY COTYLEDON1, and ABSCISIC ACID INSENSITIVE3 have been suggested. Meanwhile, the epigenome aspect impacting the transcriptional regulation of FAs is still unclear and might require more attention from the scientific community. This review mainly outlines the identification of the key gene master players involved in Perilla FAs biosynthesis and TAGs assembly that have been identified in recent years. With the recent advances in genomics resources regarding this orphan crop, we provided an updated overview of the recent contributions into the comprehension of the genetic background of fatty acid biosynthesis. The provided resources can be useful for further usage in oil-bioengineering and the design of alpha-linolenic acid-boosted Perilla genotypes in the future.

1. Introduction

Perilla frutescens var. frutescens is an oil crop from the mint family that is widely distributed in East Asia including India, Vietnam, China, and Korea [1]. The Perilla genetic resource encompasses the oil crop type P. frutescens var. frutescens, the weedy/wild type P. frutescens, and wild species Perilla setoyensis, Perilla hirtella, and Perilla citriodora [2]. While P. citriodora is known as one of the diploid progenitors [3] of tetraploid P. frutescens, the second diploid donor has not yet been elucidated. In Korean dietary habits, P. frutescens var. frutescens is used for its oil and as leafy vegetable. The fresh leaves can serve as a wrap for meat and boiled rice and are also prepared in a pickled form [2]. In China, where it originated [1,2], Perilla is used secularly as a traditional herbal medicine and fragrance [2]. The health-promoting properties of this plant are attributable to its wide panel of phytochemical compounds [4]. Among them, fatty acids including omega-3, -6, and -9 have been reported as anti-cancer agents [5,6,7], coronary heart-disease protectants [8], anti-diabetic agents [9], insulin-resistant [10], anti-cardiovascular disease agents [11], and anti-depressive agents [12,13,14]. In addition, preclinical tests revealed the positive effect of Perilla for mitigating moderate dementia [15]. However, further investigations are required to confirm its role before a recommendation for its use as an antioxidative complement for patients with dementia [4,15]. In addition, Perilla is also used as a supplement in animal feeding [16,17]. Due to the numerous applications of fatty acids from Perilla in the health industry, the oil industry, and for animal breeding, a comprehensive background underpins fatty acid biosynthesis as a fundamental prerequisite for proper utilization in the biomedical, bioengineering, and animal industries.
Recently, Perilla entered into the genomics era with the sequencing of tetraploid P. frutescence and one diploid donor P. citriodora [3], laying a foundation for unraveling the genetic basis of its multiple health and nutraceutical benefits. In the present review, we will examine recent breakthroughs on the genetic basis of fatty acid biosynthesis in Perilla.

2. Earlier Identification and Cloning of Fatty Acid Encoding Gene in Perilla

The genetic characterization interest for Perilla as an oil crop with numerous health beneficial attributes started as early as the 1900s. Several fatty acid genes have been cloned and functionally characterized. Lee et al. (https://www.ncbi.nlm.nih.gov/nuccore/U59477.1/, accessed on 12 February 2021) first characterized a ω-3 fatty acid desaturase PfrFAD7 (Genbank accession: U59477.1) extracted from a Korean cultivar “Okdong” seedling. Subsequently, a cloning of a second gene PrFAD3 was conducted by Chung et al. [18]. PrFAD3 exhibited a seed-specific expression when compared to other organs including the leaf, stem, and root, suggesting a preferential accumulation of alpha-linolenic acid (ALA) in the seed.
Hwang et al. [19,20] also reported four 3-ketoacyl-acyl carrier protein synthases (KAS) encoding genes, PfKAS3a (KAS III) and PfKAS3b (KAS III), PfFAB1 (KAS I), and PfFAB24 (KAS II/IV), which were responsible in the high accumulation of alpha-linolenic synthesis in P. frutescens seeds. Another alpha-linolenic acid-related gene, the microsomal oleate 12-desaturase (PfFAD2) gene, was functionally characterized for the first time in P. frutescens var. frutescens seed [21] in later studies. In addition to the previously identified FAD3 and FAD7 type genes, Xue et al. [22] isolated two FAD8 alpha-linoleic-related genes (PrFAD8a and PrFADb) harboring two pyrimidine stretches. Interestingly, the expression of PrFAD8 genes was predominantly observed in the Perilla bud while its accumulation increased under injury, Methyl jasmonate (MeJA), Salicylic Acid (SA), and Abscisic acid (ABA) effects; highlighting their implications in plant defense, growth, and development.

3. Transcriptomics Sheds Lights into Key Master Player Enzymes of Perilla Fatty Acid Biosynthesis

Although some genes have been investigated earlier, the fully resolved biosynthesis pathway of fatty acids in Perilla was still unclear. To fill this gap, the RNA sequencing approach has been extensively used because it helps in uncovering expressed genes related to a biological process. By deciphering the transcriptome of Perilla using diverse organs, scientists were able to identify key genes related to fatty acid biosynthesis via de novo transcripts assembly and functional gene prediction. Thus, extensive transcriptome studies have been initiated using different materials, including P. fruescens var. frutescens, Perilla frutescens var. crispa f. purpurea (red Perilla), and P. frutescens var. crispa f. viridis (green Perilla) [23,24,25,26]. The uncovered key genes involved in fatty acid biosynthesis in Perilla have been summarized in Figure 1. Briefly, based on Perilla’s fatty acid desaturase subcellular localization prediction [27] and the well-studied Arabidopsis fatty acid biosynthesis model [28], most fatty acids, including palmitic acid (C16:0), stearic acid (C18:0), and oleic acid (C18:1), were exclusively synthesized in plastids and conveyed into the cytoplasm where they entered into an acyl-CoA pool for the esterification process at sn-2 position resulting in phosphatidylcholine under the acyl-CoA:lysophosphatidylcholine acyltransferase (LPCAT) enzyme effect.
Oleic acid was then desaturated in the endoplastic rediculum (ER) to become consecutively linoleic acid (LA) and alpha-linolenic acid (ALA) under FAD2 and FAD3 genes, respectively. The resulting polyunsaturated fatty acids were transacylated onto the sn-3 position of diacylglycerol by phospholipid:diacylglycerol acyltransferase (PDAT) or returned to the acyl-CoA pool via LPCAT to be incorporated into TAG through the Kennedy pathway, inducing the production of triacylglycerols (TAGs) [29].
Using Perilla as a plant model, numerous fatty acid-related genes have been identified. From a time-course seed transcriptome analysis, Kim et al. [25] identified 43 acyl-lipid related genes in P. frutescens var. frutescens cv. Dayudeulkkae (Table 1). The identified genes via Arabidopsis orthologs detection covered the de novo fatty acid biosynthetic key enzymes present in the plastid, endoplasmic reticulum desaturases, oil body proteins, acyl-CoA-, and phosphatidylcholine-mediated TAG synthesis.
Transcriptome mining revealed five sub-unit genes (α-PDH, β-PDH, EMB3003, LTA2, and LPD1) of the precursor enzyme plastidial pyruvate dehydrogenase complex (PDHC) involved in the synthesis of acetyl-CoA from pyruvate. Afterward, acetyl-CoA carboxylase (ACCase) transformed acetyl-CoA ito malonyl-CoA [30]. The ACCase in Perilla encompassed two ACCases subunits alpha (α-CTa and α-CTb), one ACCase subunit beta (β-CT), two isoforms of biotin carboxyl-carrier protein (BCCP1 and BCCP2), and one biotin carboxylase (BC).
Furthermore, the malonyl-CoA ACP transacylase, an acyl carrier protein transacylase, catalyzed malonyl-CoA to form malonyl-ACP, paving the way for fatty acid elongation under the action of acyl-chain enzymes, i.e., 3-keto-acyl-ACP synthase (KAS), 3-ketoacyl-ACP reductase (KAR), 3-hydroxylacyl-ACP dehydratase (HAD), and Trans-∆2-enoyl-ACP reductase (EAR), respectively [23,24,31]. It is worth mentioning that WR1 is well conserved in plant species. For instance, homologous genes have been identified in Brachypodium distachyon [32], Camelina sativa [33], Solanum tuberosum [34], Cocos nucifera [35], Brassica napus [36], Elaeis guineensis [37], and Jatropha curcas [38]. In A. thaliana, through the promoter binding element AW-box, WRI1 targets upstream genes encoding for malonyl-CoA:ACP malonyl transferase, enoyl-ACP reductase, pyruvate dehydrogenase, oleoyl-ACP thioesterase, biotin carboxyl carrier protein 2, ketoacyl-ACP synthase, and hydroxyacyl-ACP dehydrase [39,40,41,42,43,44,45,46]. The homologous sequence of WR1 has been demonstrated in augmentation from 10 to 40% of seed oil in transgenic maize [47] and Brassica napus [36], suggesting that Perilla’s WR1 gene might be a promising candidate for oil-oriented bioengineering in Perilla.
Through carbon chain elongation, palmitoyl-ACP (C16:0) is converted into stearoyl-ACP (C18:0). The latter is transformed into oleic acid (C18:1)-ACP under the catalysis of stearoyl-acyl carrier protein desaturase (SAD). In Perilla, two SAD genes have been identified, including PfFAB2 and PfDES6 [25]. Using a red Perilla (Perilla frutescens var. crispa F. purpurea) seed transcriptome, Liao et al. [23] identified fatty acid desaturases PfFAD6 and PfFAD7/8 that act on the vector glycerolipid, i.e., monogalactosyldiacylglycerol (MGDG), in order to process (C18:1) into (C18:2) and (C18:2) to (C18:3), respectively (Figure 1).
To terminate fatty acids synthesis in Perilla plastids, fatty acyl-ACP thioesterase (FATA), palmitoyl/stearoyl-acyl carrier protein thioesterase (FATB), and palmitoyl-CoA hydrolase (PCH) were solicitated. PCH specifically induced C18:1- and C18:2-synthesis, while FATA was a C18:1-exclusive catalyst. Meanwhile, FATB transformed only C16:0-ACP or C18:0-ACP to C16:0 or C18:0, respectively (Figure 1). Representative gene coding for these enzyme has been pinpointed by de novo transcriptome analysis and comparative transcripts with regard to the well characterized A. thaliana fatty acid-related gene [23,24]. Free FAs were then moved into the cytoplasm where they were esterified to form an Acyl-CoA pool under the action of long-chain acyl-COA synthesis (LACS). Liao et al. [23] reported the important expression of LACS genes in Perilla seeds ten days after flowering, indicating an initiation of TAGs synthesis pathway in the endoplasmic reticulum (ER).
In the ER, esterified fatty acids are translated into phosphatidylcholines via lysophosphatidylcholine acyltransferase (LPCAT). Based on the Arabidopsis plant model, mainly two fatty acid desaturases have been identified in the ER: an FAD2 that converts PC-C18:1 into PC-18:2 and an FAD3 that catalyzes PC-C18:2 into PC-C18:3 [48,49,50]. Homologous sequences in Perilla seed (PfFAD2 and PfFAD3) transcriptome [23,24,25] have also been identified (Table 1).
Recently, the transcriptome assessment of Chinese cultivar PF40 highlighted 33 candidate genes involved in TAG biosynthesis-covering transcription factors (Supplementary Table S1), and fatty acids were exported from plastid, acyl editing of phospatidylcholine, acyl-CoA dependent Kennedy pathway, acyl-CoA independent pathway, and TAGs assembly into oil bodies (Table 1). The identified genes corroborated with previous findings [23,24,25], except for the first identification of fatty export1 (FAX1) as an additional enzyme to long-chain acyl-CoA synthetase (LACS) that mediated plastid fatty acid export.
In the absence of a whole genome representative resources, the detection of potential genes isoforms and the full FADs gene repertoire is difficult to predict, and diverse gene targets for functional validation and bio-engineering purposes are not provided. Due to the fact that Perilla has entered into the genomics era, the next section covers genomics-based advances in the detection of fatty acids in Perilla via genome-wide identification and genome-wide association study strategies.

4. Whole-Genome-Driven Fatty Acid Genes Discovery

With the advent of long-reads and chromosome conformation capture technologies, a high-quality chromosome scale genome of tetraploid P. frutescens var. frutescens has recently been assembled [3]. The genome spanned 1.203 Gb, along with 20 chromosomes with an N50 of 62.64 Mb and a total of 38,941 predicted gene models.
From a panel of 191 accessions, a genome-wide association study for seed alpha-linolenic acid content enabled the identification of an LPCAT encoding region located in chromosome 2. This finding corroborates previous observations, suggesting the role of LPCAT in FAs and TAGs synthesis in B. napus [51] and A. thaliana [52]. Interestingly, a deletion of this gene was noted in some individuals of the studied panel corresponding to a loss of around 6% of seed oil ALA content. This suggests that the transcriptional regulation of LPCAT might be responsible for ALA content variations in Perilla.
Taking advantage of the PF40-generated high-quality genome, in silico genome-wide analysis identified a repertoire of 42 fatty acid desaturases clustered into five families including omega-3 desaturase, ∆7/∆9 desaturase, FAD4 desaturase, ∆12 desaturase, and front-end desaturase [27]. The heterologous validation of candidate fatty acid desaturase genes using A. thaliana revealed a positive impact (increase of 18–37% alpha-linolenic acid content) of the PfFAD3.1 gene.
Furthermore, the upregulation of WRINKLED (WRI1), FUSCA3 (FUS3), LEAFY COTYLEDON1 (LEC1 and LCE2), and ABSCISIC ACID INSENSITIVE3 (ABI3) transcription factors was noted in PfFAD3.1 Arabidopsis transgenic lines [3] and Perilla seed expression profiles [23], suggesting their regulation roles in the Perilla FAs synthesis pathway.

5. Concluding Remarks and Outlook

Fatty acids play an important role in the lipid supply of plants and have valuable medicinal properties for humans. Here, we summarized the breakthroughs that shed light into the genetic and molecular determinants of FA and TAG synthesis in Perilla. Transcriptomics and genomics studies revealed the key master player enzymes responsible for FAs synthesis in Perilla, including polyunsaturated fatty acids desaturases, acyl-related enzymes, and transcription factors. However, the evidence of their role is still elusive since strong functional validation has not yet been provided.
The mechanism of the regulation of FA synthesis by TFs in Perilla is still elusive. Meanwhile, the recent work from Moreno-Perez et al. [53] suggested histone methylation (H3K4me3) implication into fatty acid biosynthesis in sunflowers with interactions with TFs. Moreover, acetyl-CoA, which is involved in fatty acid synthesis in plants, has been found to be correlated with histone acetylation and DNA methylation in A. thaliana through the beta-oxidation process [54]. Therefore, an in-depth investigation of identified TFs, such as ABI3, FUS3, LEC1, and LEC2, and the epigenome landmark of Perilla will pave a new avenue in deciphering the full landscape of fatty-acid biosynthesis in Perilla.
Functional validation using Perilla as a material instead of A. thaliana would drastically shape the validation efficiency of the identified genes. For this purpose, Agrobacterium-based protocols [55,56] have been tested and can serve as further functional validation. Moreover, in the current era of gene and genome editing with applicable cases in plants [57,58,59,60], designing appropriate gene editing strategies that fit into the Perilla system will surely expedite the production of enriched alpha-linolenic acid-Perilla genotypes. Furthermore, considering the species diversity within the Perilla genus, systematic fatty acid content evaluation within the Perilla species will help reveal potential alpha-linolenic acid-enriched species donors and characterize their respective biosynthetic pathways.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/plants11091207/s1, Table S1: Identified transcription factors from Perilla through trancriptome, whole genome, and in silico co-expression analyses.

Author Contributions

Conceptualization, S.-H.B.; methodology, S.-H.B. and Y.A.B.Z.; writing—original draft preparation, S.-H.B., Y.A.B.Z. and T.S.K.; writing—review and editing, S.-H.B., J.-H.O., J.L., T.-H.K. and K.Y.P.; visualization, Y.A.B.Z.; supervision, J.L., T.-H.K. and K.Y.P.; funding acquisition, K.Y.P. All authors have read and agreed to the published version of the manuscript.

Funding

This study received no external funding.

Data Availability Statement

The data presented in this study are available in Table S1.

Conflicts of Interest

The authors declare no conflict of interest.

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Plants 11 01207 g001 550
Figure 1. A simplified putative diagram view of fatty acids biosynthetic pathway in Perilla and triacylglycerols (TAGs) assembly. The schematic view involved bio-chemical interactions occurring in plastid, cytoplasm, and endoplasmic reticulum, respectively. The resulting TAGs are indicated in yellow. Purple circles indicate transcription factors, including WRINKLED (WRI1), FUSCA3 (FUS3), LEAFY COTYLEDON1 (LEC1, LCE2), and ABSCISIC ACID INSENSITIVE3 (ABI3). The transcriptional regulation of FUS3, LCE1, LCE2, and ABI3 with PfFAD3.1 is not yet uncovered. PDHC: plastidial pyruvate dehydrogenase complex; ACCase: acetyl-CoA carboxylase; MCMT: malonyl-CoA ACP transacylase; KASIII: ketoacyl-ACP synthase type III; KAR: 3-ketoacyl-ACP reductase; HAD: 3-hydroxyacyl-ACP dyhydratase; EAR: 2-enoyl-ACP reductase; KASII: ketoacyl-ACP synthase type II; KASI: ketoacyl-ACP synthase type I; SAD: stearoyl-acyl carrier protein desaturase; FATB: acyl-ACP thioesterase B; FATA: acyl-ACP thioesterase A; MGDG: monogalactosyldiacylglycerol; PfFAD: Perilla frutescens fatty acid desaturase; PC Pool: phosphatidylcholines pool; PCH: palmitoyl-CoA hydrolase; LACS: long-chain acyl-CoA synthetase; PDCT: phosphatidylcholinediacylglycerol cholinephosphotransferase; FAX: fatty acid export; LPCAT: lysophosphatidylcholine acyltransferase; PDAT: phospholipid diacylglycerol acyltransferase; DGAT: diacylglycerolacyltransferase; GPAT: glycerol-3-phosphate acyltransferase; LPAT: 1-acylglycerol-3-phosphate acyltransferase; DHAP: dihydroxyacetone phosphate; PAH: phosphatidic acid phosphatase; OLEO: Oleosin.
Figure 1. A simplified putative diagram view of fatty acids biosynthetic pathway in Perilla and triacylglycerols (TAGs) assembly. The schematic view involved bio-chemical interactions occurring in plastid, cytoplasm, and endoplasmic reticulum, respectively. The resulting TAGs are indicated in yellow. Purple circles indicate transcription factors, including WRINKLED (WRI1), FUSCA3 (FUS3), LEAFY COTYLEDON1 (LEC1, LCE2), and ABSCISIC ACID INSENSITIVE3 (ABI3). The transcriptional regulation of FUS3, LCE1, LCE2, and ABI3 with PfFAD3.1 is not yet uncovered. PDHC: plastidial pyruvate dehydrogenase complex; ACCase: acetyl-CoA carboxylase; MCMT: malonyl-CoA ACP transacylase; KASIII: ketoacyl-ACP synthase type III; KAR: 3-ketoacyl-ACP reductase; HAD: 3-hydroxyacyl-ACP dyhydratase; EAR: 2-enoyl-ACP reductase; KASII: ketoacyl-ACP synthase type II; KASI: ketoacyl-ACP synthase type I; SAD: stearoyl-acyl carrier protein desaturase; FATB: acyl-ACP thioesterase B; FATA: acyl-ACP thioesterase A; MGDG: monogalactosyldiacylglycerol; PfFAD: Perilla frutescens fatty acid desaturase; PC Pool: phosphatidylcholines pool; PCH: palmitoyl-CoA hydrolase; LACS: long-chain acyl-CoA synthetase; PDCT: phosphatidylcholinediacylglycerol cholinephosphotransferase; FAX: fatty acid export; LPCAT: lysophosphatidylcholine acyltransferase; PDAT: phospholipid diacylglycerol acyltransferase; DGAT: diacylglycerolacyltransferase; GPAT: glycerol-3-phosphate acyltransferase; LPAT: 1-acylglycerol-3-phosphate acyltransferase; DHAP: dihydroxyacetone phosphate; PAH: phosphatidic acid phosphatase; OLEO: Oleosin.
Plants 11 01207 g001
Table
Table 1. Summary of Identified Major Genes Involved in Fatty Acid and Triacylglycerols Biosynthesis in Perilla.
Table 1. Summary of Identified Major Genes Involved in Fatty Acid and Triacylglycerols Biosynthesis in Perilla.
Enzyme IDEnzyme NameGeneIDHomologousPathways InvolvedField of StudyReferences
PF40 *Dayudeulkkae **PC ***A. Thaliana
PDH(E1α)Pyruvate Dehydrogenase E1 Subunit Alpha 1 Locus_2112 AT1G01090.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
PDH(E1ß)Pyruvate Dehydrogenase E1 Subunit beta 1 Locus_25208 AT2G34590.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
EMB3003(E2)Pyruvate dehydrogenase e2 component (dihydrolipoamide acetyltransferase) Locus_33306 AT1G34430.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
LTA2 (E2)Plastid E2 Subunit of Pyruvate Decarboxylase, PLE2 Locus_5104 AT3G25860.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
LPD1 (E3)Lipoamide dehydrogenase Locus_7407 AT3G16950.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
α-CTaAlpha-carboxyltransferase
Isoform a		 Locus_8492 AT2G38040.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
α-CTbApha-carboxyltransferase
Isoform b		 Locus_2178 AT2G38040.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
ß-CTBeta-carboxyltransferase Locus_53041 ATCG00500.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
BCBiotin carboxylase Locus_22078 AT5G35360.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
BCCP1Biotin carboxyl carrier protein of acetyl-CoA carboxylase 1 Locus_29162 AT5G16390.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
BCCP2Biotin carboxyl carrier protein of acetyl-CoA carboxylase 2 Locus_17340 AT5G15530.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
MCMTMalonyl-CoA ACP transacylase Locus_14579 AT2G30200.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
KASIII3-Ketoacyl-ACP synthase Locus_10821 AT1G62640.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
KAR3-ketoacyl-ACP reductase Locus_1445 AT1G24360.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
HAD3-hydroxyacyl-ACP dyhydratase Locus_19332 AT5G10160.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
EAR2-enoyl-ACP reductase Locus_25443 AT2G05990.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
FATAFatty acyl-ACP thioesterase A Locus_29919 AT3G25110.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
FATBFatty acyl-ACP thioesterase B Locus_6603 AT1G08510.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
FAB2Fatty acid biosynthesis2 Locus_13564 AT2G43710.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
DES6Stearoyl-acyl carrier protein desaturase Locus_9486 AT1G43800.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
KASIKetoacyl-ACP Synthase I Locus_26341 AT5G46290.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
KASIIKetoacyl-ACP Synthase II Locus_1373 AT1G74960.1FA de novo biosynthesis and export from plastidTranscriptomics[25]
LACS8Long-chain acyl-CoA synthetase 8chr07_36292788_36299197
chr19_22302145_22308533		Locus_3838chr06_37084362_37090768AT2G04350.1FA de novo biosynthesis and export from plastidGenome
Assembly, Transcriptomics		[3,25]
LACS9Long-chain acyl-CoA synthetase 9chr03_70622879_70627324
chr09_58852417_58856892
chr01_02424545_02428997		Locus_23636chr01_02424545_02428997AT1G77590.1FA de novo biosynthesis and export from plastidGenome
Assembly, Transcriptomics		[3,25]
FAX1Fatty acid export 1chr05_24282740_24284950
chr01_71691539_71693779		 chr02_42552603_42554830 FA de novo biosynthesis and export from plastid [25]
FAX2Fatty acid export 2chr07_10626150_10628000 chr06_11381976_11383822 FA de novo biosynthesis and export from plastid [25]
FAX3Fatty acid export 3chr04_00857340_00859552 chr03_67540865_67543081 FA de novo biosynthesis and export from plastid [25]
FAX5Fatty acid export 5chr04_65527957_65529911
chr07_22534802_22537586
chr06_00746938_00748860
chr19_10735560_10738363		 chr03_02347871_02349825
chr06_23562111_23564893		 FA de novo biosynthesis and export from plastid [25]
FAD2Omega-6 fatty acid desaturasechr12_56933298_56934446
chr11_05592060_05593208
chr11_05575254_05576393		Locus_733chr08_55538081_55539229AT3G12120.1Acyl editing of phospatidylcholineGenome
Assembly, Transcriptomics		[3,25]
chr12_56948107_56949167 chr08_55558209_55559348
FAD3Omega-3 fatty acid desaturasechr12_04645208_04647776
chr11_54194712_54197265		Locus_22029chr08_04030082_04032640AT2G29980.1Acyl editing of phospatidylcholineGenome
Assembly, Transcriptomics		[3,25]
FAD8Omega-8 fatty acid desaturase Locus_5107 AT5G05580.2Acyl editing of phospatidylcholineTranscriptomics[25]
GPAT9Glycerol-3-phosphate acyltransferase 9chr12_33733527_33737891
chr11_26255533_26259881		Locus_10180chr08_33038421_33042132AT5G60620.1Acyl-CoA-dependent TAG synthesis in Kennedy pathwayGenome
Assembly, Transcriptomics		[3,25]
LPAT21-acyl-sn-glycerol-3-phosphate acyltransferase 2chr05_23583386_23588593
chr05_34400913_34404444
chr01_72114246_72119454		Locus_6587chr02_43313059_43318262
chr02_32585727_32589258		AT3G57650.1Acyl-CoA-dependent TAG synthesis in Kennedy pathwayGenome
Assembly, Transcriptomics		[3,25]
PAH1Phenylalanine hydrolase 1chr01_61567423_61570965
chr14_08597119_08602056
chr15_37103964_37108907
chr03_61656532_61661875
chr18_09154357_09159306
chr17_34575710_34580664
chr09_50343045_50349360		 chr10_43830659_43835596
chr01_11516392_11522733		 Acyl-CoA-dependent TAG synthesis in Kennedy pathway
DGAT1Diacylglycerol O-acyltransferase 1chr01_09730655_09741367
chr01_48275733_48286173		Locus_14696chr05_08797620_08808333AT2G19450.1Acyl-CoA-dependent TAG synthesis in Kennedy pathwayGenome
Assembly, Transcriptomics		[3,25]
DGAT2Diacylglycerol O-acyltransferase 2chr14_26782964_26787941
chr18_25811826_25816791		Locus_12629chr10_25785382_25790335AT3G51520.1Acyl-CoA-dependent TAG synthesis in Kennedy pathwayGenome
Assembly, Transcriptomics		[3,25]
DGAT3Diacylglycerol O-acyltransferase 3 Locus_1560 AT1G48300.1Acyl-CoA-dependent TAG synthesis in Kennedy pathwayTranscriptomics[25]
LPCATLysophosphatidylcholine acyltransferasechr01_06996630_07001595
chr05_56678891_56685081
chr01_03079195_03084058
chr07_53028425_53034567
chr01_43224061_43229071
chr02_66141068_66147271
chr02_04634020_04638876
chr19_35211932_35217537		Locus_43749PC00000058_00436672_00441634
chr02_10454190_10460391
chr05_03185967_03190829
chr06_54113419_54119561		AT1G12640.1PC-mediated TAG synthesisTranscriptomics[3,25]
CPT1Diacylglycerol cholinephosphotransferase Locus_7821 AT1G13560.1PC-mediated TAG synthesisTranscriptomics[25]
CPT2Diacylglycerol cholinephosphotransferase Locus_22567 AT3G25585.1PC-mediated TAG synthesisTranscriptomics[25]
PDAT1Phospholipid:diacylglycerol acyltransferase 1chr05_44104376_44108847Locus_7255chr02_22969948_22974420AT5G13640.1Acyl-CoA independent pathwayTranscriptomics[3,25]
chr03_00447151_00451507 PC00002899_00154872_00159184
chr02_52135886_52140327
chr09_00376677_00380564
PDAT2Phospholipid:diacylglycerol acyltransferase 2chr05_38922115_38924735Locus_29208chr02_28050267_28052887AT3G44830.1Acyl-CoA independent pathwayTranscriptomics[3,25]
chr02_45992086_45994691
PDCTPhosphatidylcholine:diacylglycerol cholinephosphotransferasechr03_46291224_46293449
chr09_37050943_37053194		Locus_15867chr01_27228085_27230144AT3G15820.1Acyl-CoA independent pathwayGenome
Assembly, Transcriptomics		[3,25]
OLEO2Oleosin2chr15_52133834_52134256Locus_31790 AT5G40420.1TAG assemblyTranscriptomics[3,25]
chr17_50355018_50355440 chr09_02008310_02008732
OLEOOleosinchr14_08347244_08347714Locus_31788chr10_44101965_44102435AT3G18570.1TAG assemblyTranscriptomics[3,25]
chr18_08871500_08871970
OLEO1Oleosin1chr05_05196095_05196523Locus_29266chr02_64426568_64426996AT4G25140.1TAG assemblyTranscriptomics[3,25]
chr01_30156121_30156549
OLEO5Oleosin5chr05_59989345_59989911Locus_29276 AT3G01570.1TAG assemblyTranscriptomics[3,25]
chr05_59997449_59997976 chr02_07157257_07157823
chr02_69562819_69563393 chr02_07149192_07149719
chr02_69577662_69578195
* Perilla frutescens var. frutescens cv. PF40; ** Perilla frutescens var. frutescens cv. Dayudeulkkae; *** Perilla citriodora. The mentioned genes have been identified through de novo transcriptome mining coupled with Arabidopsis homologous sequences prediction.
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Bae, S.-H.; Zoclanclounon, Y.A.B.; Kumar, T.S.; Oh, J.-H.; Lee, J.; Kim, T.-H.; Park, K.Y. Advances in Understanding the Genetic Basis of Fatty Acids Biosynthesis in Perilla: An Update. Plants 2022, 11, 1207. https://doi.org/10.3390/plants11091207

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Bae S-H, Zoclanclounon YAB, Kumar TS, Oh J-H, Lee J, Kim T-H, Park KY. Advances in Understanding the Genetic Basis of Fatty Acids Biosynthesis in Perilla: An Update. Plants. 2022; 11(9):1207. https://doi.org/10.3390/plants11091207

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Bae, Seon-Hwa, Yedomon Ange Bovys Zoclanclounon, Thamilarasan Senthil Kumar, Jae-Hyeon Oh, Jundae Lee, Tae-Ho Kim, and Ki Young Park. 2022. "Advances in Understanding the Genetic Basis of Fatty Acids Biosynthesis in Perilla: An Update" Plants 11, no. 9: 1207. https://doi.org/10.3390/plants11091207

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