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Open Access https://doi.org/10.48130/opr-0024-0029
Ornamental Plant Research 2024, 4: e032
Identification and content analysis of volatile components in 100
cultivars of Chinese herbaceous peony
Aixin Wang1#, Yasang Luo1#, Tongfei Niu1, Kai Gao2, Sitong Wang1, Xiaoyang Zhao1, Xiaogai Hou1 and Lili
Guo1*
1College of Agriculture/Tree Peony, Henan University of Science and Technology, Luoyang 471000, China
2Luoyang Academy of Agriculture and Forestry Sciences, Luoyang 471000, China
# Authors contributed equally: AixinWang, YasangLuo
* Corresponding author, E-mail: guolili@haust.edu.cn
Abstract
Herbaceous peony (Paeonia lactiflora Pall.) is a well-known and traditional flower in China, occupying a significant position in Chinese traditional
culture. The floral scent of the herbaceous peony however remains relatively understudied. The objective of this study was to investigate the
floral composition of herbaceous peony by collecting and identifying floral volatiles from 100 cultivars, including P. lactiflora 'Hangbaishao', P.
lactiflora 'Hongrongqiu', P. lactiflora 'Biandihong', P. lactiflora 'Zijin Daipao', P. lactiflora 'Zixia Yingxue', and P. lactiflora 'Fenchi Dicui'. The volatile
compounds were collected using the dynamic headspace technique and identified through gas chromatography-mass spectrometry (GC-MS).
The results demonstrated qualitative and quantitative variations in the floral fragrances emitted by the 100 cultivars, with a total of 16 volatiles
belonging to six categories (six alkanes, three alcohols and esters, two terpenes, as well as one each of ether and phenol) being identified.
However, it is notable that not all volatile categories were emitted by every cultivar. Moreover, while some compounds were present in all 100
herbaceous peony cultivars, others were exclusive to specific cultivars. The screening revealed that ten of the 16 identified flower volatile
compounds exhibited unique floral components. It is noteworthy that benzene,1,4-dimethoxy-, was identified as the most prominent compound
in several cultivars, including P. lactiflora 'Taohua Huancai', P. lactiflora 'Xishifen', P. lactiflora 'Dabanhong', P. lactiflora 'Fumantang', and P. lactiflora
'Zhushapan'. Furthermore, the clustering classification results demonstrated that benzene,1,4-dimethoxy-, exhibited the highest variable
importance in projection (VIP) value of 3.153, as determined by partial least squares discriminant analysis (PLS-DA).
Citation: Wang A, Luo Y, Niu T, Gao K, Wang S, et al. 2024. Identification and content analysis of volatile components in 100 cultivars of Chinese
herbaceous peony. Ornamental Plant Research 4: e032 https://doi.org/10.48130/opr-0024-0029
Introduction
The herbaceous peony, a well-known traditional flower in
China[1], is characterized by its large and aesthetically pleasing
flowers. The herbaceous peony is a member of the family
Paeoniaceae[2], displaying notable adaptability, and significant
ornamental value[3]. Studies on aromatic ornamental plants
involve an examination of aromatic components, and genetic
mechanisms[4], including Rosa rugosa Thunb.[5], Lilium brownii
var. viridulum Baker[6], and Paeonia suffruticosa Andr.[7], Pyrus
communis L.[8], Dendrobium officinale [9], Nymphaea
tetragona[10], Rhododendron simsii[11], Jasminum sambac[12],
studies were conducted on Chrysanthemum morifolium[13],
Osmanthus fragrans[14], Camellia japonica[15], Malus[16], and Iris
tectorum Maxim.[17] Historically, research has focused on factors
such as flower shape, color, blooming season, and resilience,
with less attention given to the floral scent[18].
The floral scent has been identified as a significant orna-
mental attribute of herbaceous peony[3,19], and is also a promi-
nent feature in numerous plant species[20]. It is frequently
described as the 'essence of flowers'[21] and is derived from a
range of volatile compounds that are synthesized within the
plant and subsequently released into the atmosphere[22]. To
date, over 1,700 volatile compounds have been identified in a
variety of plants, with a multitude of applications in the manu-
facture of perfumes, cosmetics, culinary seasonings, and
pharmaceuticals[23,24]. The composition and concentration of
these volatile compounds exhibit considerable variation across
different species, genus, and cultivars. Nevertheless, there is a
paucity of research dedicated to the analysis of fragrance cons-
tituents and their respective concentrations in herbaceous
peony and tree peony[25]. Song et al.[4] identified a total of 130
volatile compounds across 30 cultivars of herbaceous peony,
encompassing 72 aromatic constituents. The 24 cultivars exhi-
biting heightened fragrance were categorized into five distinct
aroma profiles: woody scent, fruity scent, lily scent, rose scent,
and an orange blossom scent. Zhao et al.[26] conducted a study
in which 68 volatile compounds and 26 significant aroma cons-
tituents were identified from a sample of 87 herbaceous peony
cultivars. The researchers determined that herbaceous peony
contain characteristic aromatic substances, including linalool
(resembling lily of the valley), geraniol (exhibiting a pleasant
geranium-like scent), citronellol (evoking a fresh and light rose
and leaf fragrance), and phenylethyl alcohol (noted for its dis-
tinctive rose aroma), based on the content and odor threshold
of these main aroma components. In a separate study, Li et
al.[27] identified 128 volatile compounds from 24 tree peony
cultivars, with the predominant classes being terpenes, alco-
hols, and esters. The distribution pattern of these primary
fragrance constituents led to the categorization of 24 tree
peony cultivars into four types: grass scent (ocimene), woody
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scent (longifolene), lily of the valley scent (linalool), and fruity
scent (2-ethyl hexanol). It has been demonstrated that the
distinctive fragrances of different plant species are the result of
the presence of specific volatile compounds in varying quanti-
ties and ratios. Furthermore, the quantity of fragrance emitted
by flowers is contingent upon their developmental stage[28].
Floral substances derived from plants are classified as
secondary metabolites, which are released by flowering plants
and predominantly comprise a range of volatile compounds
characterized by relatively low molecular weights. In a compre-
hensive analysis of the aromatic compounds present in P. rockii
and P. ostii 'Fengdan', Wu et al.[29] employed two-dimensional
gas chromatography coupled with time-of-flight mass spectro-
metry (GC × GC-TOF/MS). The results indicated that the aroma
profile of P. rockii was primarily characterized by the presence
of alcohols, alkanes, and acids, while the aroma profile of P. ostii
'Fengdan' was predominantly defined by aldehydes, alcohols,
and terpenes. In a separate investigation, Li et al.[30] sought to
identify and analyze the volatile compounds present in the
flowers of seven pear cultivars (Anli, Bayuesu, Golden, Brown
Peel, KorlaXiangli, Lyubaoshi, and Xizilü). The findings indi-
cated that certain aldehydes constitute significant characteris-
tics of these cultivars and are recognized as essential active
odorants, which emit pronounced citrus and floral fragrances.
Yang et al.[31] successfully identified and characterized 34
volatile compounds in the Dendrobium officinale flowers. Of
these, 18 compounds were identified as principal odorants,
including 1-octen-3-ol, hexanal, nonanal, phenylacetaldehyde,
linalool, 4-oxoisophorone, theaspirane, and methyl salicylate.
Furthermore, Kimani et al.[32] identified geraniol, β-caryophyl-
lene, 2-phenylethanol, citronellol, and 1,8-cineole as the pri-
mary aromatic constituents in 24 cultivars of herbaceous
peony, including P. lactiflora 'LianTai' and P. lactiflora 'Hongyan
Feishuang'. Aromatic compounds are recognized as the
primary chemical constituents of aromatic plants, playing a
crucial role in the synthesis of secondary metabolites[33], and
fragrance development. These compounds exhibit a diverse
range of forms. For example, phenethyl alcohol is found in rose,
mint contains menthol, and lemon includes citric acid[22,34,35].
This study employed a combination of dynamic headspace
sampling technology[36] and GC-MS to analyze the volatile com-
ponents and concentrations in 100 international herbaceous
peony cultivars during the half-opening stage. The objective
was to elucidate the aromatic profile of the herbaceous peony.
The findings of this study establish a fundamental framework
for further investigation and exploitation of the fragrances of
herbaceous peony flowers and provide a valuable resource for
enhancing the economic value of herbaceous peony.
Materials and methods
Plants
The experimental materials used in this study were obtained
from the Luoyang Academy of Agriculture and Forestry
Sciences (Luoyang City, Henan Province, China) between 20
April and 8 May 2022. The majority of the materials were
collected between 10 and 12 am. The subjects of the experi-
ment were herbaceous peony plants sourced from the her-
baceous peony resource garden affiliated with the Henan
University of Science and Technology. As outlined in Table 1
and Fig. 1, herbaceous peony cultivars demonstrating consis-
tent growth patterns and flowering stages were identified, and
the methodology entailed the repetition of each sample on
three occasions.
Experimental equipment and materials
Equipment
The Gas Chromatography-Mass Spectrometry System
(GC8890-MS5977B) from Agilent Technologies, USA, and the
Atmospheric Sampler QC-1S from the Beijing Institute of Labor
Protection were utilized in the study.
Chemicals
The reagents used included Tenax TA as the adsorbent, ethyl
caprate, dichloromethane, pentane, n-hexane of chromatogra-
phy grade, ethyl decanoate, ethyl acetate, and a standard solu-
tion of n-alkane mixture (ranging from C8 to C40) obtained from
Sigma-Aldrich, USA.
Collection methods of volatile components
The dynamic headspace adsorption technique employed in
this study was a sampling bag (355 mm × 508 mm, Reynolds,
USA) hermetically sealed at one end with an activated carbon
filter tube. The bag was meticulously wrapped around a live
peony flower to minimize contact and prevent damage to the
bag. The bag's opposite end was connected to a Tenax TA
adsorption tube (6 mm outer diameter, 100 mm length, filled
with adsorbent) and an atmospheric sampler via tasteless
Table 1. Names and numbers of 100 herbaceous peony cultivars.
100 herbaceous peony cultivars
'Hangbaishao' 'Hongrongqiu' 'Biandihong' 'Zijin Daipao' 'Zixia Yingxue' 'Fenchi Dicui' 'Xishifen' 'Yinlong
Hanzhu'
'Yinxian
Xiuhongpao'
'Jindaiwei'
'Luhong' 'Xueyuan
Hongxing'
'Mozijin' 'Yahong' 'Wulong
Tanhai'
'Hongyan
Zhengshuang'
'Xingguang
Chanlan'
'Yanlihong' 'Hongling
Chijin'
'Fenzhuangyuan'
'Taohua
Huancai'
'Zhongshenghong' 'Ziling' 'Luxihong' 'Zifurong' 'Hongling Chijin' 'Huguang
Shise'
'Hongyuqiu' 'Yanzhi
Dianyu'
'Lantian
Piaoxiang'
'Zhushapan' 'Hongyun Yingri' 'Yanzi Xiangyang' 'Yanzhihong' 'Zaoyuanhong' 'ChilongCaifeng' 'Chaoshihong' 'Qingwen' 'Shaifugui' 'Ziyanshuang'
'Gaoganfen' 'Qundiehui' 'Meirenmian' 'Meiju' 'Dafugui' 'Zhifeng
Zhaoyang'
'Xueyuan
Hongxing'
'Dahongpao' 'Zixiuqiu' 'Canglong'
'Gaoganhong' 'Hongyan
Feishuang'
'Dabanhong' 'Zifengyu' 'Hongpan
Jinqiu'
'Hushui Dangxia' 'Yinlong
Huihai'
'Baihuazi' 'Taohuafen' 'Wawamian'
'Fenpanjinxing' 'Heixiuqiu' 'Shuanghonglou' 'Changshouhong' 'Hongyan
Lushuang'
'Tuopan Jinhua' 'Hongling
Chijin'
'Linglongyu' 'Jinzan Ciyu' 'Xiangyang
Qihua'
'Jinbian Hongge' 'Duoyezi' 'Fenzilou' 'Furong Jinhua' 'Fenkui' 'Guifei Chacui' 'Huolian
Jingang'
'Hongguanfang' 'Fenmian
Taohua'
'Taoranzui'
'Zhaoyanghong' 'Hongfengyu' 'Fumantang' 'Shaonvfen' 'Danfeng' 'Liantaizi' 'Meiguihong' 'Fenfurong' 'Fenling
Hongzhu'
'Fenqiu'
'Fencuiqiu' 'FengChao Chuyu' 'Lanju' 'Jinsanhong' 'Zhaoyuanfen' 'Hongfeng' 'Qiaoling' 'Tuanye Jinqiu' 'Guohong' 'Tongquechun'
The numbers 1−100 are listed from top to bottom, left to right respectively.
Identification and analysis of volatiles in herbaceous peony
Page 2 of 10 Wang et al. Ornamental Plant Research 2024, 4: e032
silicone tubing. The flow rate of the atmospheric sampler was
set at 400 mL·min−1 and the sampling duration was 3 h. Follo-
wing the sampling period, the adsorption tube was sealed with
cling film and aluminum foil, then placed in a self-sealing bag
and stored in an ultra-low temperature cooler for transport to
the laboratory. The sample was then eluted with n-hexane
during sample processing, and the eluate was transferred to a
brown sample bottle for subsequent analysis.
The following conditions were observed in the gas chroma-
tography (GC) analysis: the chromatographic column employed
is a flexible quartz capillary column, with a length of 30 mm, an
internal diameter of 0.25 mm, and a pore size of 0.25 μm. The
flow rate of the column is set at 1.2 mL·min−1. The temperature
of the column is maintained according to a specific protocol.
It is initially set at 70 °C and held for 1 min, then increased to
136 °C at a rate of 6 °C·min−1, followed by further increases
to 138 °C at a rate of 1 °C·min−1, then to 142 °C at a rate of
2 °C·min−1, and finally to 143 °C at a rate of 0.5 °C∙min−1. The
temperature is increased by 5 °C·min−1 and subsequently to
160 °C at a rate of 2 °C·min−1, before reaching 250 °C at a rate of
10 °C·min−1. The injector temperature is set at 250 °C, with a
carrier gas of high-purity helium at a flow rate of 1 mL·min−1.
The injection mode is a split injection, with a split ratio of 9:1,
and the injection volume is 2 μL.
The following conditions were employed for the mass spec-
trometry (MS) analysis: The electron impact (EI) source is
operated at 70 eV, with the interface temperature set to 250 °C
and the ion source temperature maintained at 230 °C. The
quadrupole temperature is controlled at 150 °C, and the scan
range is from 25 to 400 amu.
Analysis methods of volatile components
Qualitative analysis
Before the analysis of the sample using gas chromatography,
the 500 mg∙L−1 n-alkane mixed standard solution should be
diluted with n-hexane at a ratio of 1:50, in accordance with the
specified conditions for the chromatography. It is essential to
record the retention time for each n-alkane and to compare the
resulting Retention Index (RI) values with those documented in
the literature to facilitate the identification of the compounds
in question. The following formula is used to calculate the RI:
RI =100 ×n+100 ×(tx−tn)/(tn+1−tn)
The location of the aforementioned item is as follows: The
retention index (RI) represents the retention time of the volatile
Fig. 1 Morphological characteristics of 100 herbaceous peony cultivars at the half-opening stage. The order of the above pictures is relative
to the order of cultivars in Table 1.
Identification and analysis of volatiles in herbaceous peony
Wang et al. Ornamental Plant Research 2024, 4: e032 Page 3 of 10
substances under examination. The number of carbon atoms in
the straight-chain alkane preceding the analyte is represented
by n. The retention time of the analyte is represented by tx. The
retention time of the straight-chain alkane before the analyte is
represented by tn. The retention time of the straight-chain
alkane following the analyte is represented by tn+1. The reten-
tion time of the analyte falls between the retention times of
tn and tn+1. Qualitative analysis of volatile components is con-
ducted through consultation with the NIST 17 spectral library,
with cross-referencing of RI values, literature sources, and other
pertinent resources, including books.
Quantitative analysis
An internal standard solution, comprising 69.32 mg∙L−1 of
ethyl decanoate in ethyl acetate, is employed. A volume of
0.4 μL of the internal standard solution is added to each 80 μL
sample. Subsequently, quantitative calculations are performed
in accordance with the following formula:
Content of each aroma substance (μg·g−1)=
Peak area of each aroma substance
Peak area of the internal standard ×
Concentration of the internal standard (mg ·L−1)×
Volume of the internal standard (μL)
Volume of the sample (g) ×f
where, f is the correction factor of each component to the internal
standard, f = 1.
Data cleaning and analysis
The analysis of variance can be conducted using the statis-
tical software package SPSS, while graph plotting can be
accomplished with the Origin 2022 software. The software
Metaboanalyst and the Microbiome Analysis Platform are capa-
ble of performing data normalization, partial least squares
discriminant analysis (PLS-DA), and cluster analysis.
Results
Examination of the primary volatile compounds in
various cultivars of herbaceous peony during the
half opening stage
An analysis was conducted to determine the main volatile
compounds present in 100 herbaceous peony cultivars during
the half-opening stage. This was achieved through the utiliza-
tion of database retrieval and manual identification methods.
The results are outlined in Table 2. A total of 16 volatile compo-
nents were identified and classified into six distinct groups. The
data revealed that alkanes constituted six types, representing
37.5% of the total volatile components. This was followed by
four types of esters at 25%, three types of alcohols at 18.75%,
and one type each of terpenes, ethers, and phenols, each
accounting for 6.25% of the total volatile components. The
results of this analysis indicate that the predominant cate-
gories of volatile compounds found in herbaceous peony culti-
vars are alkanes, esters, and alcohols.
Comparison of volatile compounds content in
different cultivars of herbaceous peony during
the half opening stage
Alkane compounds
As illustrated in Fig. 2, alkane compounds were undetected
in 30 cultivars, including P. lactiflora 'Hushui Dangxia', P. lacti-
flora 'Tuopan Jinhua', P. lactiflora 'Qiaoling', P. lactiflora 'Yinlong
Hanzhu', and P. lactiflora 'Yanlihong'. Among the 100 herba-
ceous peony cultivars, the highest concentration of alkane
compounds was observed in P. lactiflora 'Heizijin' (10.66 ±
2.01 μg·g−1), with the range of alkane compounds concentra-
tion spanning from 0.00 to 10.66 μg·g−1.
Ester compounds
As shown in Fig. 2, ester compounds were discernible in all
44 cultivars of herbaceous peony at the half-opening stage.
However, the content of ester compounds was generally not
notably high in most cultivars. The highest ester compounds
content was observed in P. lactiflora 'Changshouhong' (9.15 ±
0.03 μg·g−1), followed by P. lactiflora 'Zaoyuanhong' (3.55 ±
0.40 μg·g−1), P. lactiflora 'Hongyun Yingri' (3.37 ± 0.11 μg·g−1),
and P. lactiflora 'Saifugui' (3.25 ± 0.67 μg·g−1). The ester
compounds content among these three cultivars was found to
be similar, with a range of 0.00 to 9.15 μg·g−1.
Alcohol compounds
As depicted in Fig. 2, the majority of the 100 cultivars of
herbaceous peony at the half-opening stage exhibited the
presence of alcohol compounds. Only 23 cultivars, including
Table 2. The volatile components of 100 herbaceous peony cultivars.
Compound
number RT (min) CAS number Compounds Compound
classification
Chemical
formula
Retention index
Calculated value Reference value
1 3.273 111-84-2 Nonane Alkanes C9H20 900 900
2 4.805 124-18-5 Decane Alkanes C10H22 1,000 1,000
3 5.727 13877-91-3 (Z)-β-ocimene Terpenes C10H16 1,038 1,037
4 7.063 60-12-8 Phenylethyl alcohol Alcohols C8H10O 1,115 1,109
5 8.133 150-78-7 Benzene,1,4-dimethoxy- Ethers C8H10O21,165 1,168
6 9.502 106-22-9 Citronellol Alcohols C10H20O 1,228 1,228
7 10.084 106-25-2 Nerol Alcohols C10H18O 1,220 1,219
8 10.187 103-45-7 Methyl cinnamate Esters C10H12O21,260 1,258
9 12.976 103-26-4 2-Propenoic acid,3-phenyl-,methyl ester Esters C10H10O21,389 1,380
10 14.88 131-11-3 Dimethyl phthalate Esters C10H10O41,456 1,466
11 16.174 629-62-9 Pentadecane Alkanes C15H32 1,500 1,500
12 16.664 128-37-0 Butylated hydroxytoluene Phenols C15H24O 1,513 1,513
13 19.877 544-76-3 Hexadecane Alkanes C16H34 1,600 1,601
14 24.137 629-78-7 Heptadecane Alkanes C17H36 1,699 1,700
15 31.517 84-74-2 Dibutyl phthalate Esters C16H22O41,964 1,907
16 33.398 646-31-1 Tetracosane Alkanes C24H50 2,400 2,400
Identification and analysis of volatiles in herbaceous peony
Page 4 of 10 Wang et al. Ornamental Plant Research 2024, 4: e032
P. lactiflora 'Taohua Huancai', P. lactiflora 'Zhushapan', and P.
lactiflora 'Gaoganhong' exhibited no detection. The highest
alcohol compounds content was observed in P. lactiflora
'Hongfeng' (22.98 ± 3.86 μg·g−1), which was significantly higher
than that of other herbaceous peony cultivars. Subsequently,
P. lactiflora 'Wandai Shengse' (16.23 ± 2.28 μg·g−1) exhibited
the second-highest alcohol compounds content, with a range
of 0.00 to 22.98 μg·g−1.
Terpene compounds
As illustrated in Fig. 2, only 19 of the herbaceous peony culti-
vars exhibited detectable levels of terpene compounds, with
significant differences in content (p < 0.05). The highest
content was observed in P. lactiflora 'Hongfengyu' (8.19 ±
1.02 μg·g−1), followed by P. lactiflora 'Wandai Shengse' (4.93 ±
0.09 μg·g−1), P. lactiflora 'Jinzan Ciyu' (2.92 ± 1.75 μg·g−1), P.
lactiflora 'Dabanhong' (0.07 ± 0.13 μg·g−1), P. lactiflora 'Jinbian
Hongge' (0.14 ± 0.23 μg·g−1), and P. lactiflora 'Mozi Hanjin' (0.16
± 0.28 μg·g−1), among others. The range of terpene compounds
content was found to vary from 0.00 to 8.19 μg·g−1.
Ether compounds
The analysis of 50 herbaceous peony cultivars revealed the
presence of ether compounds in all samples, with notable varia-
tions in their content (p < 0.05). The highest content of ether
compounds was observed in P. lactiflora 'Dabanhong' (22.84 ±
2.15 μg·g−1), followed by P. lactiflora 'Taohua Yingcai' (19.53 ±
2.44 μg·g−1). The lowest levels were observed in P. lactiflora
Alkanes
Esters
Alcohols
Terpenes
Ethers
Phenols
8
6
4
2
0
100
80
94
93
83
87
74
92
56
99
73
31
27
82
89
61
63
62
40
66
35
39
13
42
36
44
2
30
28
20
32
5
22
46
14
19
37
75
21
78
95
91
96
90
76
88
86
98
49
48
50
1
10
69
68
55
71
45
70
16
51
12
65
97
79
81
72
77
85
52
38
58
57
67
8
7
47
53
6
34
24
25
18
11
23
64
54
41
33
15
9
43
17
60
59
84
29
4
3
26
Fig. 2 Comparative heat map depicting the release of six types of volatile compounds from various herbaceous peony cultivars.
Identification and analysis of volatiles in herbaceous peony
Wang et al. Ornamental Plant Research 2024, 4: e032 Page 5 of 10
'Danfeng' (0.06 ± 0.11μg·g−1), P. lactiflora 'Ziling' (0.15 ± 0.26
μg·g−1), and P. lactiflora 'Huolian Jingang' (0.12 ± 0.21 μg·g−1).
The range of ether compounds content was observed to vary
from 0.00 to 22.84 μg·g−1.
Phenol compounds
The analysis revealed that only five herbaceous peony culti-
vars exhibited discernible levels of phenol compounds, namely
P. lactiflora 'Jinbian Hongge' (0.15 ± 0.05 μg·g−1), P. lactiflora
'Zhaoyanghong' (0.34 ± 0.02 μg·g−1). The remaining cultivars
exhibited lower levels of phenol compounds, with the lowest
concentration observed in P. lactiflora 'Hongrongqiu' (0.17 ±
0.03 μg·g−1), followed by P. lactiflora 'Xueyuan Honghua' (0.01 ±
0.02 μg·g−1), and P. lactiflora 'Ziling' (0.27 ± 0.05 μg·g−1). The five
cultivars exhibited notably lower levels of phenol compounds,
with values consistently below 1 μg·g−1. The remaining culti-
vars were found to be devoid of phenol compounds.
An analysis was conducted on the total release of
aroma components in different herbaceous peony
cultivars
The analysis of the 16 volatile compounds detected revealed
that, aside from alkanes such as nonane, the remaining 10 com-
pounds from five classes all exhibited characteristic aromas, as
detailed in Table 3. These aromatic compounds were present
in the majority of samples, with concentrations exceeding
0.01 μg·g−1. Of particular note is the detection of benzene,1,4-
dimethoxy-, in the majority of samples, with relatively high
concentrations observed (Fig. 3).
Cluster analysis of aromatic components in
various herbaceous peony cultivars
A data matrix of dimensions 100 × 10 was constructed, repre-
senting the content of 10 aromatic compounds in 100 herba-
ceous peony cultivars as variables. A cluster heatmap was
generated using the microbiome analysis platform, as illus-
trated in Fig. 4. In light of the clustering results and a compre-
hensive consideration of the major aromatic components, the
100 herbaceous peony cultivars are ultimately classified into
two groups (Table 4). The first group of herbaceous peony culti-
vars is distinguished by a marked prevalence of benzene,1,4-
dimethoxy-, with markedly elevated levels in comparison to
other cultivars. This gives rise to a pronounced clove scent,
indicative of a clove floral type. This initial classification is based
on the presence of specific compounds and is therefore
Table 3. Characteristics of aroma compounds.
No. Compound
name
Odor characteristics
1 (Z)-β-ocimene The scent of grass and flowers is accompanied
by the aroma of orange blossom oil[37]
2 Phenylethyl
alcohol
Sweet rose-like fragrance[38]
3 Benzene,1,4-
dimethoxy-
The fragrance of cloves[39]
4 Citronellol Has a sweet rose aroma[40]
5 Nerol There is a sweet rose fragrance[41]
6 Acetic acid, 2-
phenylethyl ester
There is a reminiscent of honey-like floral
fragrance[42]
7 Methyl
cinnamate
Sweet smelling fragrance[43]
8Dimethyl
phthalate The substance emits a delicate fragrance[44]
9 Butylated
hydroxytoluene
The presence of a carbonic acid taste can
influence the aroma of wine[45]
10 Dibutyl
phthalate
The substance emits a delicate fragrance[46]
Comparison of the contents of ten characteristic aroma components
of different herbaceous peony cultivars (μg·g−1)
Fig. 3 Content of characteristic aroma compounds in herbaceous peony cultivars.
Identification and analysis of volatiles in herbaceous peony
Page 6 of 10 Wang et al. Ornamental Plant Research 2024, 4: e032
applicable to only five cultivars. The cultivars in question are P.
lactiflora 'Taohua Huancai', P. lactiflora 'Xishifen', P. lactiflora
'Dabanhong', P. lactiflora 'Fumantang', and P. lactiflora
'Zhushapan'. The second group generally exhibits lower levels
of aromatic compounds, resulting in milder scents that may be
characterized as a light floral type. The second group comprises
95 cultivars, including representative cultivars such as P. lacti-
flora 'Meiju', P. lactiflora 'Shaonvfen', P. lactiflora 'Fenmian
Taohua', P. lactiflora 'Fenling Hongzhu', and P. lactiflora
'Guohuo', among others.
Partial Least Squares Discriminant Analysis (PLS-
DA)
Following the clustering of 100 cultivars into two groups,
a partial least squares discriminant analysis (PLS-DA) was
conducted on the content of 10 aroma compounds in the 100
cultivars using Metaboanalyst software. The results of the
analysis are presented in Fig. 5. The PLS model for aroma com-
pounds demonstrated satisfactory reliability, as evidenced by
R2 and Q2 values of 0.702 and 0.598, respectively. Moreover, the
PLS-DA results demonstrated variations in the profile of aroma
compounds between the two groups of cultivars (Fig. 5a).
The application of a VIP criterion greater than 1 identified a
differentiating component (Fig. 5b). The VIP values in the PLS-
DA model provided further insight into the contribution of
each component to the model, with components having a
value of VIP > 1 being considered significant. For instance,
benzene,1,4-dimethoxy-, exhibited a VIP value of 3.153 and was
identified as a principal component accountable for the dis-
crepancies among herbaceous peony cultivars (Fig. 5b),
corroborating the findings of the clustering analysis. It can
therefore be posited that benzene,1,4-dimethoxy- is a charac-
teristic aroma component of these herbaceous peony cultivars.
Variables A and B represent the first and second categories,
respectively. The specific variables include A-Benzene,1,4-
dimethoxy-, B-Citronellol, C-Nerol, D-Acetic acid, 2-phenylethyl
ester, E-Methyl cinnamate, F-Dimethyl phthalate, G-(Z)-β-
ocimene, H-Phenylethyl alcohol, I-Butylated hydroxytoluene,
J-Dibutyl phthalate.
Discussion
The present study comprises a comprehensive identification
and analysis of the volatile constituents present in 100 her-
baceous peony cultivars during the half-opening stage. The
findings indicated that alkanes, alcohols, and ethers were
the most prevalent volatile compounds, with benzene,1,4-
dimethoxy- was identified as the distinctive aromatic
components.
One such molecule is benzene,1,4-dimethoxy-, a methoxy-
lated aromatic volatile compound that is known to elicit phy-
siological and behavioral responses in a diverse range of insect
pollinators. It serves as a principal floral volatile in a number of
plant species belonging to diverse genera, including Salix,
Lithophragma, Nelumbo, Catasetum, Allium, and Fragaria[47].
Wang et al.[40] identified the common floral component,
5
4
3
2
1
0
J
I
H
G
F
E
D
C
B
A
3
26
4
29
61
59
19
37
31
46
10
74
67
14
53
6
17
13
8
42
25
36
2
24
44
43
52
38
32
33
15
16
23
41
40
85
87
47
78
57
76
91
95
80
83
86
100
12
70
94
45
98
34
75
11
63
58
18
22
77
65
79
62
97
81
71
72
51
54
55
68
69
9
90
89
88
66
64
35
39
20
73
5
7
28
49
48
93
82
60
84
27
56
96
50
92
30
21
99
1
Fig. 4 Heat map showing the clustering analysis of 100 herbaceous peony cultivars. A-Benzene,1,4-dimethoxy-, B-Citronellol, C-Nerol, D-
Acetic acid, 2-phenylethyl ester, E-Methyl cinnamate, F-Dimethyl phthalate, G-(Z)-β-ocimene, H-Phenylethyl alcohol, I-Butylated hydroxy-
toluene, J-Dibutyl phthalate. The numbers 1−100 correspond to the cultivar names listed in Table 1.
Identification and analysis of volatiles in herbaceous peony
Wang et al. Ornamental Plant Research 2024, 4: e032 Page 7 of 10
benzene,1,4-dimethoxy-, in all eight herbaceous peony
cultivars. Furthermore, Kimani et al.[32] identified 95 volatile
organic compounds in 24 herbaceous peony cultivars, inclu-
ding benzene,1,4-dimethoxy-, which is a phenolic methyl ether
containing a benzene skeleton but not derived from aromatic
amino acids. Rather, it is a member of a particular chemical
class that is responsible for the olfactory characteristics of spe-
cific plant varieties. The types and contents of volatile compo-
nents of herbaceous peonies may be associated with the
sampling method, sampling location and time. Additionally,
the types and contents of volatile compounds in plants may be
influenced by different planting environmental conditions[48].
In recent years, there has been a growing emphasis on the
natural floral volatiles present in herbaceous peonies, with the
fragrance components demonstrating a diverse range of appli-
cations in the fields of healthcare, perfumes, and cosmetics[49].
Floral scent represents a significant component of plant vola-
tiles, which are primarily composed of terpenes, aromatic
hydrocarbons, fatty acids, and their derivatives, as well as sulfur
and nitrogen-containing compounds[27,50]. These compounds
are taxonomically categorized into three primary classes, con-
tingent upon their biogenic origins, namely fatty acid deriva-
tives, phenylpropanoids/benzenoids, and terpenoids[51]. In the
present study, the volatile components of the 100 cultivars of
herbaceous peony were predominantly identified as alkanes,
esters, and alcohols. The most abundant type of compound
was identified as alkanes. The available evidence suggests that
straight-chain alkanes represent the primary constituents of
plant leaf wax[52]. These waxes are not exclusive to leaves but
may also be found on other plant organs, including flowers and
fruit surfaces[53]. This indicates that wax layers may cover the
surfaces of the majority of herbaceous peony cultivars. Alco-
hols play a significant role in the fragrance industry, serving as
essential raw materials for synthetic fragrances and as an indis-
pensable component in perfumery[54]. The presence of abun-
dant ether compounds results in the production of pleasant
floral and fruity aromas, while simultaneously enhancing the
richness, typicality, and complexity of plant fragrances[55].
However, due to the constraints of the existing literature,
some volatile components, such as specific alkanes, have not
yet been conclusively identified as fragrance components.
Further research is required to ascertain whether these com-
ponents contribute to the fragrance of herbaceous species.
Alkane compounds have relatively high thresholds[56] and
make minimal contributions to the overall scent[28]. Accor-
dingly, the analysis of fragrance compounds excludes the
contributions made by nonane, decane, pentadecane, hexade-
cane, heptadecane, and tetracosane.
Conclusions
This study employed dynamic headspace bag adsorption of
live plant materials and gas chromatography-mass spectrome-
try (GC-MS) analysis techniques to identify a total of 16 volatile
Table 4. Cluster analysis of characteristic aroma components in different herbaceous peony cultivars.
Groups Herbaceous peony cultivars
1 'Taohua Huancai', 'Xishifen', 'Dabanhong', 'Fumantang', and 'Zhushapan'
2 'Liantaizi', 'Hushui Dangxia', 'Shaifugui', 'Hongfeng', 'Wandai Shengse', 'Zhaoyuanfen', 'Wawamian', 'Lanju', 'Shuanghonglou', 'Fenling
Hongzhu', 'Guohuo', 'Fenmian Taohua', 'Yinlong Tanhai', 'Chaoshihong', 'Shaonvfen', 'Meiju', 'Huolian Jingang', 'Meiguihong', 'Chilong
Huancai', 'Yinlong Hanzhu', 'Yanlihong', 'Zhaoyanghong', 'Yinxian Xiuhongpao', 'Fenchi Dicui', 'Xueyuan Hongxing', 'Fenfurong',
'Linglongyu', 'Xiangyang Qihua', 'Hongrongqiu', 'Huguang Shise', 'Yanzhihong', 'Duoyezi', 'Mozijin', 'Guifei Chacui', 'Ziling', 'Zixia
Yingxue', 'Zixiuqiu', 'Jinzan Ciyu', 'Meirenmian', 'Zifengyu', 'Jinshanhong', 'Hongyan Lushuang', 'Hongguanfang', 'Jindaiwei', 'Canglong',
'Tuopan Jinhua', 'Huolian Chijin', 'Fengchao Chuyu', 'Hongyuqiu', 'Xueyuan Hongxing', 'Qiaoling', 'Dahongpao', 'Qundiehui', 'Tuanye
Jinqiu', 'Dafugui', 'Taoranzui', 'Yanzhi Dianyu', 'Tongquechun', 'Ziyanshuang', 'Gaoganfen', 'Fenpan Jinxing', 'Fenkui', 'Lantian Piaoxiang',
'Zifeng Zhaoyang', 'Xingguang Canlan', 'Hongyan Feishuang', 'Biahuazi', 'Taohuafen', 'Danfeng', 'Hongfengyu', 'Fenzilou', 'Yanzi
Xiangyang', 'Zaoyuanhong', 'Luhong', 'Yahong', 'Luxihong', 'Furong Jinhua', 'Jinbian Hongge', 'Wulong Tanhai', 'Zhongshenghua',
'Zifurong', 'Hongyan Zhengshuang', 'Gaoganhong', 'Heixiuqiu', 'Hongling Chijin', 'Hongyun Yingri', 'Changshouhong', 'Fencuiqiu',
'Qingwen', 'Hongpan Jinqiu', 'Zijin Daipao', 'Biandihong', 'Fenqiu', 'Hangbaishao' and 'Fenzhuangyuan'
Scores plot
a b
4
2
0
−2
−4
A
H
G
E
J
B
C
F
I
D
−40 −30 −20
Component 1 (65.6%) VIP scores
Component 2 (9.3%)
−10 0 10 0.0 0.5 1.0 1.5 2.0 2.5 3.0
a
b
a b
High
Low
Fig. 5 PLS-DA scores of 100 herbaceous peony cultivars under two cluster groups.
Identification and analysis of volatiles in herbaceous peony
Page 8 of 10 Wang et al. Ornamental Plant Research 2024, 4: e032
components in 100 herbaceous peony cultivars at the half-
opening stage[57]. The components were primarily categorized
into six major groups: alkanes, esters, alcohols, terpenes, ethers,
and phenols. The predominant volatile compounds were alka-
nes, alcohols, and ethers, while benzene,1,4-dimethoxy- was
identified as the main aromatic component. Significant varia-
tions in the total content of the main aromatic components
were observed among the different herbaceous peony culti-
vars at the half-opening stage. In particular, P. lactiflora 'Taohua
Huancai', P. lactiflora 'Xishifen', P. lactiflora 'Dabanhong', P. lacti-
flora 'Fumantang', and P. lactiflora 'Zhushapan' exhibited the
highest content of aromatic components, resulting in a more
intense floral fragrance. The intensity and characteristics of the
aroma exhibited notable variation among different herbaceous
peony cultivars, attributable to differences in the quantity and
composition of the aromatic components. This is a crucial indi-
cator for evaluating the quality of herbaceous peony. This study
provides a theoretical foundation for understanding the forma-
tion and regulation mechanisms of herbaceous peony aroma
characteristics, while also offering technical support for accele-
rating industrial development and utilization of herbaceous
peony aromas.
Author contributions
The authors confirm contribution to the paper as follows:
study conceptualization, reviewing, editing and funding acqui-
sition: Guo L; material preparation: Wang A, Luo Y, Niu T, Zhao
X, Gao K; data curation: Wang A, Luo Y, Niu T, Wang S; draft
manuscript preparation: Wang A, Luo Y; manuscript reviewing
and editing: Hou X. All authors reviewed the results and
approved the final version of the manuscript.
Data availability
All data generated or analyzed during this study are included
in this published article.
Acknowledgments
This research was funded by the Science and Technology
Innovation Talents in Universities of Henan Province (Grant No.
22HASTIT036) and the Project of Henan Province Traditional
Chinese Medicine Industry Technology System (Grant No. 2024-
24).
Conflict of interest
The authors declare that they have no conflict of interest.
Dates
Received 2 July 2024; Revised 11 September 2024; Accepted
12 October 2024; Published online 11 December 2024
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