Received: 5 June 2023 Accepted: 5 June 2023
DOI: 10.1111/ppl.13947
EDITORIAL
Plant volatiles and color compounds: From biosynthesis to
function
Plants biosynthesize secondary metabolites that can be useful in various
applications, including health promotion, thanks to their organoleptic
properties and antioxidant activity. Moreover, pigments such as caroten-
oids, apocarotenoids, and polyphenols and volatiles such as fatty acid
derivatives, terpenoids, and phenylpropanoids contribute to attracting pol-
linators to flowers and to facilitating seed dispersal by animals (Abbas
et al., 2020, 2023; Dudareva et al., 2013). Floral volatiles are complex,
multifunctional signals that are often used by pollinators in combination
with other signals, such as color (Mostafa et al., 2022; Muhlemann
et al., 2014). Floral visitors use floral scent to estimate the amount of
reward present in flowers, to facilitate the identification of a specific host
flower or as signals that chemically resemble those important for pollina-
tor insects in other ecological contexts. Some plant volatiles also have
defensive roles, for example, as repellant for pest insects, which impacts
their interaction with their environment (Pichersky & Gershenzon, 2002).
There are good pieces of evidence that floral scent evolves under selec-
tion imposed by both mutualists and antagonists (Nadimpalli et al., 2000;
Ninkovic et al., 2019, 2021). Antagonists may often limit the amount of
scent emitted by flowers, thus contributing to spatial population variation.
Floral scent is also an important component of pollinator-mediated
reproductive isolation, as it often covaries with color and morphology in
sister species with different pollination systems (Ninkuu et al., 2021). The
molecular and biochemical mechanisms underlying the interactions and
cross talks between color and volatiles' biosynthesis pathways and their
regulation are not fully understood, although it has been discovered that
some transcription factors involved are common (Abbas, Ke, Zhou, Yu,
Waseem, Ashraf, Li, et al., 2021; Abbas, Ke, Zhou, Yu, Waseem, Ashraf,
Wang, et al., 2021; Mostafa et al., 2022; Qiao et al., 2021). The roles of
these metabolites in plant biotic or abiotic stress and plant communica-
tion are also still not fully understood. The special issue includes several
research and review articles highlighting the role of floral volatiles and
color compounds in various plant life span as well as in human life.
Herein, Sarkar and Sadhukhan (2023) addressed the modification
of plant volatiles in response to mycorrhizal fungi, which act as a
defensive shield toward plant pathogens. Arbuscular mycorrhizal fungi
(AMFs) not only induced the plan defense mechanism but also modi-
fied the emission pattern of volatile organic compounds (VOCs). The
authors also highlight up-to-date understanding of the cross talk of
AMF and VOC patterns in plants and the subsequent modulation of
resistance against microbial pathogens.
Abbas, O'Neill Rothenberg, et al. (2022) reviewed the diverse role
of plant VOCs, including plant defenses, attracting pollinators, and
mediating various interactions between plants and their environment.
Physiologia Plantarum. 2023;175:e13947. wileyonlinelibrary.com/journal/ppl
https://doi.org/10.1111/ppl.13947
Physiologia Plantarum
The authors described the recent updates on the ecological aspects
and context-dependence of plant–plant communication driven by vol-
atiles, crucial elements that influence plant performance in different
habitats. The authors described the biosynthesis pathways of volatile
terpenoids as being the most abundant class of VOCs. The interesting
aspect of the review was about the adaptability of crop plants and the
interplay of VOCs upon global climate change.
Karalija et al. (2023) reviewed how plants use volatile compounds
as a defense mechanism against aphids. The authors emphasized the
complexity and ecological significance of the tritrophic interplay
between host plants, aphids, and parasitic wasps, all of which are
directly influenced by climate change. Aphids use VOC emissions from
plants to locate their host plant, but plants can use the same signaling
mechanism to entice parasitic wasps as an indirect defense system.
The release of certain terpenoids in response to an aphid threat is
linked to the appeal of host-specific parasitic wasps.
Imran et al. (2023) investigated the significance of CRISPR-Cas9
and its function in aroma fragrance in rice using CRISPR-Cas9-mediated
editing of BETAINE ALDEHYDE DEHYDROGENASE 2 (BADH2) gene.
The researchers emphasize that the natural aroma compound 2-acetyl-
1-pyrroline (2-AP) is a pivotal fragrance substance among the over
200 volatiles outlined in fragrant rice. They stated that BADH2 is an
important gene for transcriptional regulation of 2-AP biosynthesis and
that CRISPR-Cas9 is a useful weapon for aroma enhancement. In addi-
tion, the author discusses the transcriptional cascades in the regulation
of 2-AP biosynthetic pathway under Zn availability in fragrant rice
(Imran et al., 2022). There were over 678 transcription factors involved
in the upregulation of aroma compound (2-AP), with the majority of
them belonging to the WRKY, bHLH, and MYB families. Likewise, Ash-
raf et al. (2022) addressed how alternate wetting and drying (AWD)
affected physio-biochemical attributes, grain yield, and aroma volatiles
in fragrant rice. They concluded that AWD is more effective during the
reproductive stage, promoting yield and quality attributes as well as
grain aroma volatiles in terms of 2-AP content. Furthermore, the puta-
tive role of type-B authentic response regulator genes in fragrant rice
under alkaline salt stress was highlighted (Rehman et al., 2022).
Wang et al. (2023) performed metabolome and transcriptome
analysis to uncover the framework of anthocyanin biosynthesis deficit
in litchi (Litchi chinensis Sonn.) aril. The authors show that anthocya-
nins are more abundant in mature litchi pericarp than in aril (flesh). In
addition to LcUFGT1 and LcMYB1, WGCNA identified 52 other can-
didate genes with a consistent expression profile. SGR, CYP75B1,
GST, and MATE were among the seven genes linked to anthocyanin
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Physiologia Plantarum
synthesis and transport, while another gene (LcMYB1) was found to
positively stimulate the promoters of LcUFGT1, LcGST4, and
LcSWEET10 through the prediction and identification of regulatory
interplay among structural candidate genes. A similar methodology
was used to unravel the role of putative structural terpene synthase
genes involved in aroma biosynthesis (Abbas, Guo, et al., 2022). First,
the terpene synthase gene family was thoroughly examined, followed
by a metabolomic approach to assess the volatile profile of litchi
flowers and fruits at various stages of development. After integrating
the data with multivariate analysis, key terpene synthase genes
involved in fruit and flower aroma biosynthesis were discovered.
Ke et al. (2023) used a similar strategy to identify putative structural
biosynthesis genes and key transcription factors associated with floral
aroma production in Hydrangea arborescens “Anabella.” The volatile pro-
file of ‘Annabelle’ was analyzed during flowering stages (F1–F3) using
HS-SPME-GC-MS and UPLC-MS/MS metabolome approach. Most
flower volatile components released during the F2 stage were found to
be benzenoids and phenylpropanoids. Multiple genes showed differential
expression, and overlapping benzenoid/phenylpropanoid pathways were
discovered in developing flowers. There was substantial multinode con-
nectivity between GRAS/bHLH/MYB/AP2/WRKY, making them promis-
ing candidates for regulating benzenoids/phenylpropanoids volatiles.
The ornamental trait diversity and classification of 144 caladium
accessions were investigated by Zhou et al. (2023), who used EST-
SSR-based association analysis. Based on the transcriptome data, the
authors isolated 7706 SSR markers and subsequently screened
26 SSR markers for polymorphism and reproducibility. SSR markers
associated with petiole color, main vein color, and blade upperside
glossiness were found ( p < 0.01).
Chen et al. (2023) emphasized the significance of anthocyanin in
fruit and flower color formation. They stated that acylation modification
is essential for anthocyanin modification and that the acyltransferase
gene (PhAAT1) is an important factor in the anthocyanin mecha-
nism in petunia. The authors mentioned that PhAAT1 was strongly
regulated by MYB TFs (PhAN2) and that silencing PhAAT1 lowered
anthocyanin stability and modified color by reducing caffeoylated
and p-coumaroylated anthocyanins, and PhAAT1 protein was located in
the cytoplasm. Another study found that 5-O-glucosyltransferase (5GT)
has a major impact on anthocyanin contents in petunia flowers, and
that suppressing the aforementioned gene changed the color of the
petunia from purple to light purple (Li et al., 2022). Furthermore, meta-
bolic findings indicate that glycosylation significantly altered the 5-O-
position of petunidins and malvidins. Besides that, petunia rhamnose
synthase (PhRHMs) plays an integral role in leaf and flower develop-
ment as well as anthocyanin synthesis (Sang et al., 2022). The authors
discovered and characterized three PhRHM genes (PhRHM1/2/3) and
one PhUER1 gene. Silencing PhRHM1/2 resulted in abnormal plant
growth and decreased total anthocyanin, L-rhamnose, and photosyn-
thetic pigments, whereas PhUER1 did not. Furthermore, a multilateral
crotonylation proteome evaluation of PaACL-silenced corollas and con-
trol plants disclosed that crotonylation protein was strongly linked to
protein acetylation and that proteins with far more crotonylation sites
frequently had several acetylation sites (Yang et al., 2022).
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EDITORIAL
Zhang et al. (2023) highlighted the anthocyanin role in Chaeno-
meles speciosa. The authors discovered that, under drought conditions,
CsMYB6 interacts with CsbHLH11 to regulate anthocyanin formation.
Using suppression and transient expression assays, the authors dem-
onstrated that CsMYB6 is a key TF in anthocyanin biosynthesis. They
also found that CsMYB6 directly binds to the promoters of
CsbHLH11 and structural anthocyanin biosynthesis genes (CsUFGT,
CsDFR, and CsANS). Another research found that loss-of-function
mutations in ObANS1/2 are involved in the all-green phenotype of
Ocimum basilicum L. (Gonda et al., 2023). The researchers looked at
quantitative trait loci for sweet basil flowers and stem color in a bipa-
rental F2 population and revealed that both ANS genes had a loss-
of-function mutation. ObANS has a single base pair insertion in the
active site, whereas ObANS has a missense mutation. In the purple-
flower parent, ANS1 was functional, but ANS2 has a nonsense muta-
tion. Complementation assays in an Arabidopsis ANS mutant were
used to validate the role of ObANS1 active allele. Another finding
showed that exogenous potassium application improves drought tol-
erance in Camelina (Camelina sativa L.) (Bukhari et al., 2022).
Ali et al. (2023) report the role of synthetic volatiles derived
from Bacillus strains in nematode mortality and induced growth in
rice under extreme oxidative stress. These volatiles [benzaldehyde
(BDH), albuterol, dimethyl disulfide (DMDS), 2-undecanone (2-UD),
1,2-benzisothiazol-3(2H)-one (1,2-HIT), and 1,3-propanediole
(1,3-PD)] were used as biocontrol agents to control nematodes and
have shown higher nematicidal activity with mortality rate than con-
trol. The growth-promoting traits of the Bacillus strains GBSC56,
SYST2, and FZB42, as well as their synthetic VOCs, led to notable
improvements in the corresponding physiological parameters. Addi-
tionally, the expression of specific growth-promoting and defense-
related genes was significantly upregulated in rice seedlings exposed
to synthetic VOCs.
Likewise, whitefly contact toxicity, repellency, and anti-oviposition
were all tested using three different essential oils (EOs) extracted from
plants using ethanol (Wagan et al., 2022). Under controlled conditions,
Allium ascalonicum, Cinnamomum camphora, and Mentha haplocalyx
were protected from whitefly (Bemisia tabaci) for up to 72 h when trea-
ted with EOs. C. camphora EOs outperformed M. haplocalyx and A.
ascalonicum in repellency, contact toxicity, and oviposition deterrence.
The chemical components of these EOs were detected via gas
chromatography–mass spectrometry (GC–MS) after their antagonistic
impacts were established.
In conclusion, the current special issue was initiated to bring
together articles highlighting various roles of floral volatiles and color
pigments in plants' life span. Numerous key structural genes and tran-
scription factors involved in floral aroma and anthocyanin biosynthesis
mechanisms, as well as their roles under stress conditions, were revealed.
Furthermore, the role of EOs in the defense mechanism was determined.
The application of modern tools and techniques, both analytical and com-
putational, useful for identifying, characterizing, and functionally validating
posttranscriptional regulators were key in the research cited above. This
special issue brings new insights on the physiological, molecular, and bio-
chemical cues to plant secondary metabolites.

EDITORIAL
ACKNOWLEDGMENTS
This work was supported by fundings from the Guangzhou Science
and Technology Project (202103000057 and 202206010023), and
China Litchi and Longan Industry Technology Research System
(CARS-32-08).
FUND ING INFORMATION
Guangzhou Science and Technology Project, Grant/Award Numbers:
202206010023, 202103000057; China Litchi and Longan Industry
Technology Research System, Grant/Award Number: CARS-32-08
Farhat Abbas 1
Yixun Yu 1
Mohammed Bendahmane 2
Hui-Cong Wang 1
1
Key Laboratory of Biology and Genetic Improvement of Horticultural
Crops-South China/Guangdong Litchi Engineering Research Center,
College of Horticulture, South China Agricultural University, Guangzhou,
China
2
Laboratoire Reproduction et Development des Plantes, INRA-CNRS-
Lyon1-ENS, Ecole Normale Supérieure de Lyon, Lyon, France
Correspondence
Hui-Cong Wang, Key Laboratory of Biology and Genetic
Improvement of Horticultural Crops-South China/Guangdong Litchi
Engineering Research Center, College of Horticulture, South China
Agricultural University, Guangzhou, China.
Email: wanghc1972@263.net
ORCID
Farhat Abbas https://orcid.org/0000-0003-3704-5213
Mohammed Bendahmane https://orcid.org/0000-0003-1661-1060
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Physiologia Plantarum
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Rehman, O.U., Uzair, M., Chao, H., Fiaz, S., Khan, M.R. & Chen, M. (2022)
Role of the type-B authentic response regulator gene family in fragrant
rice under alkaline salt stress. Physiologia Plantarum, 174(3), e13696.
Sang, L., Chen, G., Cao, J., Liu, J. & Yu, Y. (2022) PhRHMs play important
roles in leaf and flower development and anthocyanin synthesis in
petunia. Physiologia Plantarum, 174(5), e13773.
Sarkar, A.K. & Sadhukhan, S. (2023) Unearthing the alteration in plant vola-
tiles induced by mycorrhizal fungi: a shield against plant pathogens.
Physiologia Plantarum, 175(1), e13845.
Wagan, T.A., Hua, H., Bozdog an, H. & Khan, M.M. (2022) Repellency,
contact toxicity, and anti-oviposition effects of three ethanol-
extracted plant essential oils on Bemisia tabaci (Hemiptera: Aleyrodidae).
Physiologia Plantarum, 174(6), e13799.
13993054, 2023, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/ppl.13947 by Cochrane Mexico, Wiley Online Library on [29/06/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
EDITORIAL
Wang, D., Chen, L., Yang, Y., Abbas, F., Qin, Y., Lu, H. et al. (2023) Inte-
grated metabolome and transcriptome analysis reveals the cause of
anthocyanin biosynthesis deficiency in litchi aril. Physiologia Plantarum,
175(1), e13860.
Yang, W., Li, X., Jiang, G., Long, Y., Li, H., Yu, S. et al. (2022) Crotonylation
versus acetylation in petunia corollas with reduced acetyl-CoA due to
PaACL silencing. Physiologia Plantarum, 174(5), e13794.
Zhang, S., Ren, Y., Zhao, Q., Wu, Y., Zhuo, Y. & Li, H. (2023) Drought-induced
CsMYB6 interacts with CsbHLH111 to regulate anthocyanin biosynthe-
sis in Chaenomeles speciosa. Physiologia Plantarum, 175(1), e13859.
Zhou, Y., Ye, Y., Zhu, G., Xu, Y., Tan, J. & Liu, J. (2023) Diversity, classifica-
tion, and EST-SSR-based association analysis of caladium ornamental
traits. Physiologia Plantarum, 175(1), e13841.