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130 A. Gupta et al.
8.3.3 HLH TFs
This class of potential regulatory TFs act by formation of homo/heterodimeric
complexes with the MYB TFs via their two constituent alpha helices that tend to
form a helix-loop-helix association with the proteins (Frerigmann et al.
2014). These
TFs are composed of a conserved bipartite domain made up of 60-am ino-acids. They
generally bind to the promoter sequence of the target DNA by means of N-terminals
of their key amino acid residues (Yamamura et al.
2015). Typically, the bHLH TFs
are involved in the upregulation of specific biosynthetic genes of anthocyanin,
alkaloids, glucosinolate, diterpenoid phytoalexin and saponins (Nemesio-Gorriz
et al.
2017), (Table 8.3).
8.3.4 bZIP TFs
Similar to bHLH TF family, the bZIP TFs also occur as dimeric proteins typically
involved in imparting tolerance to ROS and osmotic imbalance in plant species
(Shaaban et al.
2010). Structurally, these consist of a conserved leucine zipper with a
DNA binding domain that contains positively charged amino acid residues specific
to the promoter region of the corresponding biosynthetic genes (Hong et al.
2013)
(Table 8.4)
8.3.5 AP2/ERF TFs
This family of TFs have been the most studied class of TFs in plants and have been
reported with the occurrence of a unique and conserved DNA binding domain
known as AP2. This domain is ~60-amino acid long and has been named after a
floral homeotic gene APETALA2 (AP2) of A. thaliana in which it was first
described (Schnall and Quatrano
1992). In addition, it also has other conserved
domains, such as ERF, RAV and DREB, which differ in terms of the number of AP2
repeats or their sub-families (Zhou and Memelink
2016). A study on different
members of AP2 ERF family suggested the role of two such proteins, ORCA and
ORCA2 in regulating the synthesis of terpenoid indole alkaloids by binding to the
respective promoter regions of their genes. Another similar protein ORCA3 was
reported to govern the synthesis of enzymes of the jasmonic acid pathway namely
strictosidine synthase and tryptophan deca rboxylase by binding to the JERE element
of the promoter regions in their respective genes (Wasternack and So ng
2017). A list
of AP2 ERF TFs studied as regulators of different classes of SMs in plants has been
given in Table
8.5.
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8 Transcriptional Regulation in Biosynthesis of Phytochemicals in... 131
Table 8.3 Different members of bHLH family responsible for regulation of secondary metabolite biosynthesis in plants
TF family Plant species Pathway Responsible gene
Class of SM
involved Associated property Reference
bHLH04,
bHLH05 and
bHLH06 with
MYB51
A. thaliana Phenylpropanoid
pathway
CYP79B2, CYP79B3 and
CYP83B1, CYP79F2
CYP83A1, SOT16,
PAPST1
Glucosinolates Wound/insect responses,
flavonoid biosynthesis
and oxidative stress
tolerance
Frerigmann
et al. (2014)
CrMYC2 Catharanthusroseus Jasmonic acid ORCA2 and ORCA3 Alkaloid Defence against insects
and certain pathogens
Zhang et al.
(2011)
DPF Rice (Oryza sativa) Methylerythritol
phosphate pathway,
salicylic acid
pathway
CPS2, CYP99A2 Diterpenoid
phytoalexin
Abiotic stress tolerance Yamamura
et al. (2015)
NbbHLH1,
NbbHLH2 and
NbbHLH3
Nicotiana
benthamiana
Jasmonic acid
pathway
PMT, MPO Alkaloid
nicotine
Protects against biotic
stress
Todd et al.
(
2010)
TSAR1 and
TSAR2
Medicago
truncatula
Mevalonate
pathway
HMGR Triterpene
saponin
Defense against insect
feeding and microbial
pathogens
Mertens
et al. (
2016)
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132 A. Gupta et al.
Table 8.4 Different members of bZIP family responsible for regulation of secondary metabolite biosynthesis in plants
TF family
Plant
species Pathway Responsible gene
Class of SM
involved Associated property Reference
SmbZIP20,
SmbZIP7 and
AabZIP1
Salvia
miltiorrhiza
Tanshinone
biosynthetic
pathway
HMGR2, CPS1,
KSL1 and
CYP76AH1
Tanshinone Anti-atherosclerosis, antioxidant, anti-
cancer and anti-inflammatory activities
Zhang
et al.
(2018)
MdHY5 Apple Phenylpropanoid
pathway
MdMYB10 Anthocyanin Nitrate assimilation An et al.
(2017)
OsTGAP1 Rice MEP pathway OsDXS3, OsKSL4
and OsKSL7
Terpenoid
phytoalexins
Blast pathogen-resistance Yoshida
et al.
(2017)
AtfB and Apyap1 Aspergillus
parasiticus
cAMP/PKA/PI3K
signaling pathway
Nor-1, ver-1 and
aflR
Aflatoxin Promotes resistance of conidiospores to heat
shock, osmotic stress and oxidative stress
Roze
et al.
(2011)
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8 Transcriptional Regulation in Biosynthesis of Phytochemicals in... 133
Table 8.5 Different members of AP2 ERF family responsible for regulation of secondary metabolite biosynthesis in plants
TF family Plant species Binding site Responsible gene
Class of SM
involved Pathway Reference
AaERF1 and AaERF2 Artemisia
annua
CBF2 and
RAA
CYP71AV1 and ADS Artemisinin and
artemisinic acid
Jasmonic acid Yu et al.
(2012)
NtORC1/ERF221 and
NtJAP1/ERF10
Tobacco PMT NtORC1 and NtJAP1 Nicotine and
pyridine alkaloids
Jasmonic acid De Sutter
et al. (2005)
GAME9 Tomato CaMV 35S JRE1, JRE2, JRE3, JRE4,
JRE5 and JRE6
Steroidal
glycoalkaloids
Mevalonate
pathway
Thagun et al.
(2016)
PnERF1 Panax
notoginseng
CYP71AV1 HMGR, EPS, DS and SS Saponins PNS biosynthetic
pathway
Deng et al.
(
2017)
GbERF1 Gossypium
barbadense
GhHCT1,
AtPAL3
PAL, C4H, C3H, CCR, HCT,
CoMT and F5H
Lignin Jasmonic acid,
ethylene pathway
Guo et al.
(
2016)
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134 A. Gupta et al.
8.3.6 NAC TFs
NAC TFs have been named after their characteristic NAC domain whose C-terminus
is diverse while the N-terminus remains highly conserved and acts as a transcrip-
tional regulator (Olsen et al.
2005). The NAC TFs have been predominantly found in
plants namely rice and Arabidopsis with nearly 100 members of this family (Olsen
et al.
2005), (Table 8.6).
8.4 Techniques Intricated in Delineating the Regulatory
Pathways
In addition to the biosynthesis of the different classes of SMs and their TFs, the
co-synthesis of the metabolite as well as the transcription factors is also regulated by
key regulatory mechanisms involved in the prediction of gene-to-metabolite
correlations (Saito
2013). Apparently, this takes place via an integrated network
analysis of transcriptomes and metabolomes (Fukushima et al. 2009). Towards this,
the advent of high-throughput DNA sequencing techniques and the current trends
and advances in ‘omics’ technology has readily assisted in identification of the
candidate genes (Rai et al.
2017). These techniques delineate the correlation between
the expression pattern of genes and metabolite accumulation or identify
co-expressed clusters of genes within a pathway (Winzer et al.
2012). More often,
these strategies are concerned with the application of elicitors or exogenous
hormones in culture systems in order to study their stimulating effects on the
accumulation of SM (Villarreal-García et al.
2016).
These strategies basically involve the over-expression of yield-boosting biosyn-
thetic enzymes or inactivation of endogenous genes by virus-induced gene silencing
or RNA interference (Gupta et al.
2017). Moreover, in order to enhance the produc-
tion of suitable phytochemicals under stress conditions, the target peptides of
heterologous enzymes are either added, removed or modified to alter their subcellu-
lar location. In addition, phylogenetics and genome mining techniques are also
employed as strategies to optimize the production of phytochemicals during stress
(Vashisht et al.
2015).
8.5 Conclusion
SMs are protective components of the plant system that are usually produced in
small quantities. Stress stimuli in the environment, however, induce their increased
secretion by effecting characteristic metabolomic changes in the host plant that affect
their synthesis. This environment induced biosynthesis of SMs is regulated at the
molecular level by the expression o f individual or multiple marker genes
participating in the biosynthetic pathways. Moreover, these genes further are
regulated by the associated TFs, which work in synchrony as a network to mediate
the regulation involved in environmental stress tolerance. Both biotic and abiotic
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8 Transcriptional Regulation in Biosynthesis of Phytochemicals in... 135
Table 8.6 Different members of NAC family responsible for regulation of secondary metabolite biosynthesis in plants
TF family
Plant
species Pathway Responsible gene
Class of SM
involved Property Reference
ANAC042 A. thaliana Ethylene signaling
pathway
CYP71A12, CYP71A13 and
CYP71B15
Camalexin Tolerance to Alternaria
brassicicola infection
Saga et al.
(2012)
PtrNAC72 Poncirus
trifoliata
ABA signaling
pathway
Arginine decarboxylase (ADC) Putrescine ROS homeostasis Wu et al.
(2016)
MfNAC Medicago
falcate
KEGG pathway Glyoxalase-1 gene (GLO1) Glutathione ROS homeostasis Duan et al.
(2017)
HbNAC1 Hevea
brasiliensis
Jasmonic acid Small rubber particle protein Latex Drought tolerance Cao et al.
(
2017)
ANAC032 A. thaliana Phenylpropanoid
pathway
CHS, F30H and LAR3, PAL1,
DFR and ANS
Flavonoid Tolerance to salinity stress Mahmood et al.
(
2016)
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stress tolerance is governed by the specific TFs, which either inhibit pest and
pathogen infection, deter herbivore attack or maintain redox balance and confer
resistance to oxidative stress.
136 A. Gupta et al.
8.6 Future Directions
Overall, this chapter directs further research towards enhancing bioactive accumula-
tion of SMs in stress conditions. Besides, it also provides scope to additional
experiments for the creation of stress tolerant plants. These experiments would be
based on the exploration, identification and functional characterization of candidate
and marker genes, metabolites and TFs as well as the use of advanced techniques for
the validation of stress tolerance property in plants on exogenous application
of SMs.
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