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Apiaceae Medicinal Plants: A Review of Traditional Uses, Phytochemistry, Bolting and Flowering, and Controlling Approaches

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04 May 2023

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05 May 2023

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Abstract
Apiaceae plants have been widely used as traditional Chinese medicines (TCMs) for the treatment of removing dampness, relieving superficies, and dispelling cold, etc. In order to exploit the potential application and improve the yield and quality of Apiaceae medicinal plants (AMPs), The traditional use, phytochemistry, modern pharmacological use, effect of bolting and flowering (BF), and approaches for controlling the BF were summarized. Currently, about 228 AMPs have been recorded as TCMs with 6 medicinal parts, 72 traditional uses, 62 modern pharmacological uses, and 5 main kinds of metabolites. Three effect degrees (i.e., significantly affected, affected to some extent, and no significantly affected) could be classed based on the yield and quality. The BF of individual plants (e.g., Angelica sinensis) could be effectively controlled by the standard cultivation techniques, while the mechanism of BF has not been systemically revealed. This review will provide useful references for the reasonable exploration and high-quality production of AMPs.
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Subject: Biology and Life Sciences  -   Life Sciences

1. Introduction

Apiaceae (syn. Umbelliferae) is one of the largest angiosperm families including 300 genera (3000 species) in the world and 100 genera (614 species) in China [1]. Apiaceae plants have been widely used as medical healthcare, nutrition, food industry, and other fields [2]. Currently, 55 genera (230 species) of Apiaceae plants have been applied in medical healthcare, and over 20 species have been widely used as traditional Chinese medicines (TCMs) [3]. Extensive studies have demonstrated that Apiaceae medicinal plants (AMPs) present a variety of pharmacological properties for the treatment of central nervous system, cardiovascular, and respiratory system diseases, amongst others [1,4]. These pharmacological activities are largely associated with bioactive metabolites such as polysaccharides, alkaloids, phenylpropanoids (simple phenylpropanoids and coumarins), flavonoids, and polyene alkynes [1,5,6].
In China, Apiaceae plants have been primarily used as traditional medicines for main treatment of removing dampness, relaxing tendons, activating blood, relieving superficies, and dispelling cold, etc. [1,2]. For example, rhizomatous and whole plants are mainly used for the treatment of common cold due to wind-cold, cough, asthma, rheumatic arthralgia, as well as ulcerative carbuncle and pyogenes infections; fruits are mainly used for the treatment of expelling pathogenic wind and regulating vital energy, harmonizing the stomach and promoting digestion, as well as relaxing abdominal pain and expelling parasite [1,2].
As known, the occurrence of bolting and flowering (BF) plays a critical role in transiting from vegetative growth to reproductive development in plant life cycle [7]. However, the BF significantly reduces the accumulation of metabolites in vegetative organs, which ultimately leads to the lignification of rhizomes and/or roots such as sugar beet [8], lettuce [9], and Chinese cabbage [10]. Particularly, it is more common that the BF significantly reduces the yield and quality for the rhizomatous AMPs [11]. Extensive studies have demonstrated that the BF is regulated by both internal factors (e.g., germplasm resource, seedling size, and plant age) and external factors (e.g., vernalization, photoperiodism, and environmental stresses) [12]. To date, the BF, especially in the rhizomatous AMPs, has not been effectively controlled [11,13].
In order to comprehensively learn about the current status of AMPs, herein, the progress on traditional use, phytochemistry, bolting and flowering, and controlling approaches were summarized. These reviews will provide references for efficient cultivation and quality improvement of AMPs.

2. Materials and Methods

All information involved in AMPs was searched on scientific databases (i.e., PubMed, Web of science, Springer, and CNKI) using the keywords including: Apiaceae plant, traditional use, phytochemistry, bolting and flowering, and lignification. Additional information was collected from ethnobotanical literatures focusing on herb from Flora of China and local herbal classic literature, such as Divine Husbandman’s Classic of the Materia Medica (Shen Nong Ben Cao Jing), Compendium of Materia Medica, Illustrated Book on Plants, Collection of National Chinese Herbal Medicine, and Pharmacopoeia of the People’s Republic of China. The names of all the plants were corresponded to the Catalogue of Life China. Chemical structures were drawn using ChemDraw 2021 software.

3. A tour of Apiaceae Medicinal Plants (AMPs)

Apiaceae plants have been traditionally used as medicines in China for ca. 2400 years (Figure 1). In 390-278 BC, 3 Apiaceae plants including Angelica dahurica, Ligusticum chuanxiong, and Cnidium monnieri were firstly recorded as medicines in Sorrow after Departure [1,2]. With the progress of Chinese civilization, ca. 100 Apiaceae plants were historically recorded as medicines. Specifically, 12 AMPs such as Angelica decursiva, Bupleurum chinense, and Centella asiatica were recorded in the known herbal text of China, the Divine Husbandman’s Classic of the Materia Medica (Shen Nong Ben Cao Jing) in 1st-2nd century AD [14]; In 1578 and 1848, 24 and 31 AMPs were respectively recorded in the Compendium of Materia Medica and Illustrated Book on Plants [15]. In the 21st century, the number of AMPs has been increasing up to 93 species recorded in Flora of China in 2002 [16], and 96 species in Collection of National Chinese Herbal Medicine in 2014 [17]. In recent years, 22 species are recorded in Pharmacopoeia of the People’s Republic of China [18]; specifially, 18 species are used with rhizomes and/or roots (Table 1).

4. Classification of AMPs Species

To our best knowledge, a total of 228 AMPs used as TCMs are collected from previously published literatures and books (Table 1). Based on the traditionally used medicinal parts, the 228 AMPs are categorized into 6 classes including: 51 species (21 genera) used with the whole plants (i.e., rhizome and/or root, stem, and leaf), 184 species (44 genera) used with rhizomes and/or roots, 5 species (5 genera) used with stems, 9 species (8 genera) used with leaves, 17 species (14 genera) used with fruits, and single species (single genus) used with seeds.
Specifically, the 51 species (21 genera) used with whole plants include: Anethum, Anthriscus, Apium, Bupleurum, Centella, Conium, Coriandrum, Cryptotaenia, Eryngium, Ferula, Foeniculum, Hydrocotyle, Oenanthe, Peucedanum, Pimpinella, Pleurospermum, Pternopetalum, Sanicula, Sium, Spuriopimpinella, and Torilis genera. Specially, Sanicula (e.g., S. astrantiifolia, S. caerulescens, S. chinensis), Hydrocotyle (e.g., H. himalaica, H. hookeri, and H. nepalensis), and Pimpinella (e.g., P. candolleana, P. coriacea, and P. diversifolia) genera plants are usually used with whole plants.
The 184 species (44 genera) used with rhizomes and/or roots, which are mainly used in AMPs, include: Angelica, Anthriscus, Apium, Archangelica, Bupleurum, Carum, Changium, Chuanminshen, Cicuta, Cnidium, Conioselinum, Daucus, Eriocycla, Ferula, Foeniculum, Glehnia, Heracleum, Hymenidium, Kitagawia, Levisticum, Libanotis, Ligusticopsis, Ligusticum, Meeboldia, Nothosmyrnium, Oenanthe, Osmorhiza, Ostericum, Peucedanum, Phlojodicarpus, Physospermopsis, Pimpinella, Pleurospermum, Pternopetalum, Sanicula, Saposhnikovia, Selinum, Semenovia, Seseli, Seselopsis, Spuriopimpinella, Tongoloa, Torilis, and Vicatia genera. Specially, Angelica (e.g., A. biserrata, A. dahurica, and A. sinensis), Bupleurum (e.g., B. bicaule, B. chinense, and B. scorzonerifolium), and Ligusticum (L. chuanxiong, L. jeholense, and L. sinense) genera plants are usually used with whole plants.
The 5 species (5 genera) used with stems include: Aegopodium (A. alpestre), Coriandrum (C. sativum), Foeniculum (F. vulgare), Ligusticum (L. chuanxiong), and Oenanthe (O. javanica); the 9 species (8 genera) used with leaves include: Aegopodium (A. alpestre), Anethum (A. graveolens), Angelica (A. morii), Anthriscus (A. nemorosa and A. sylvestris), Carum (C. carvi), Daucus (D. carota), Foeniculum (F. vulgare), and Ligusticum (L. chuanxiong); the 17 species (14 genera) used with fruits include: Ammi (A. majus), Carum (C. buriaticum and C. carvi), Cnidium (C. monnieri), Coriandrum (C. sativum), Cuminum (C. cyminum), Cyclorhiza (C. peucedanifolia), Daucus (D. carota L. and D. carota var. Carota), Pimpinella (P. anisum), Trachyspermum (T. ammi), and Visnaga (V. daucoides) genera; as well as the single genera used with seeds is Ferula (F. bungeana) (Table 1).

5. Traditional Uses

As shown in Table 1, distinct traditional uses of the 228 AMPs were recorded. Based on their clinical agents, a total of 79 traditional uses are enriched, with 40 species (e.g., Angelica apaensis, Conium maculatum, and Hydrocotyle hookeri) contributing to the treatment of relieving pain, 36 species (e.g., Aegopodium alpestre, Apium graveolens, and Carum carvi) to the treatment of dispelling wind; and 21 species (e.g., Conioselinum vaginatum, Hydrocotyle sibthorpioides var. batrachium, and Ligusticum sinense) to the treatment of eliminating dampness (Figure 2).
What’s more, AMPs were also widely used as ethnodrug in ethnic minority of China. For example, Carum carvi was used as Tibetan medicine for the treatment of dispelling wind and eliminating dampness, treating cat fever and joint pain [86], Trachyspermum ammi [235] was used as Uygur medicine for the treatment of eliminating cold damp, dispelling coldness, and promoting digestion; Angelica acutiloba was used as Korean nationalities medicine for the treatment of strengthening spleen, enriching blood, stopping bleeding, and promoting coronary circulation [236]; Angelica sinensis was used as Tujia minority medicine for the treatment of enriching the blood, treating dysmenorrheal, and relaxing bowel [237]; and Chuanminshen violaceum was used as geo-authentic medicine of Sichuan province for the treatment of moistening lung melt phlegm, as well as nourishing spleen and stomach [89].
Meanwhile, AMPs combined with other herbs have also been applied in many prescriptions for thousands of years [238]. For example, Decoction of Notopterygium for Rheumatism, a famous Chinese prescription, composed of Notopterygium incisum, Angelica biserrata, Ligusticum sinense, Eryngium foetidum, and Ligusticum chuanxiong, etc., has been widely used for the treatment of exopathogenic wind-cold, rheumatism, headache, and pantalgia [94]. Xinyisan composed of Yulania liliiflora, Actaea cimicifuga, Angelica dahurica, Eryngium foetidum, and Ligusticum sinense, etc., has been widely used for the treatment of deficiency of pulmonary qi and nasal obstruction due to wind-cold pathogens and damp-heat in lung channel [94,167]. Shiquan Dabu Wan of Angelica sinensis, recorded in Pharmacopoeia of the People’s Republic of China, has been mainly used for treatment of pallor, fatigability, and palpitations [239]; and Juanbi Tang of Notopterygium incisum and Angelica biserrata, recorded in Medical Words (Qing dynasty), has been mainly used for treatment of arthralgia due to wind cold-dampness [120].

6. Modern Pharmacological Uses

Modern pharmacological researches of the 228 AMPs were recorded (Table 1). Based on their pharmacological effects, a total of 62 modern uses are enriched (Figure 3), with 36 species (e.g., Angelica biserrata, Bupleurum. marginatum, and Foeniculum vulgare) showing anti-inflammatory activity, 20 species (e.g., Chuanminshen violaceum, Cryptotaenia japonica, and Ferula songarica) showing antioxidant activity, and 16 species (e.g., Anethum graveolens, Centella asiatica, and Changium smyrnioides) showing antitumor activity.
Specifically, the sesquiterpene-coumarin, such as (3’S, 5’S, 8’R, 9’S, 10’R)-kellerin, gummosin, galbanic acid, and methyl galbanate in Ferula sinkiangensis resin, showed the anti-neuroinflammatory effect and might be a potential natural therapeutic agent for Alzheimer’s disease [240]. The ferulin B and C in Ferula ferulaeoides rhizomes could restrain the multiplication of HepG2 stomach cancer cell lines, and 2,3-dihydro-7-hydroxyl-2R*, 3R*-dimethyl-2-[4,8-dimethyl-3(E),7-nonadienyl]-furo[3,2-c] coumarin could restrain the proliferation of HepG2, MCF-7, and C6 cancer cell lines [107,241]. The osthole in Angelica biserrata could restrain the multiplication of human gastric cancer cell lines MKN-45 and BGC-823, human lung adenocarcinoma cell line A549, human mammary carcinoma cell line MCF-7, and human colon carcinoma cell line LOVO [242]. The phthalides (i.e., sedanolide and 3-n-butylphthalide) in Apium graveolens showed the anticarcinogenic and neuroprotective properties [243,244].

7. Phytochemistry

As shown in Table 1, hundreds of bioactive metabolites have been identified from the 228 AMPs [1,245]. Based on their chemical structures, these metabolites can be categorized into 5 main classes including: (1) polysaccharides, (2) alkaloids, (3) phenylpropanoids, (4) flavonoids, and (5) terpenoids (Figure 4).
Among the 22 AMPs recorded in the Pharmacopoeia of the People’s Republic of China [18], 18 bioactive metabolites in the 17 AMPs (e.g., Angelica biserrata, Bupleurum chinense DC., and Centella asiatica) (Figure 5) were described as quality control indicators, which include: 10 phenylpropanoids (i.e., osthole, columbianadin, imperatorin, isoimperatorin, nodakenin, ferulic acid, ttrans-anethole, notopterol, praeruptorin A, and praeruptorin B), 4 terpenoids (i.e., saikosaponin a, saikosaponin d, asiaticoside, and madecassoside), 2 chromones (i.e., prim-O-glucosylcimifugin and 5-O-methylvisammioside), and 2 phthalides (i.e., ligustilide and levistilide A,); and there is no specific quality marker mentioned for the other 5 AMPs (e.g., Changium smyrnioides, Daucus carota L., and Glehnia littoralis) (Table 2).

7.1. Polysaccharides

Polysaccharides are the largest component of biomass and account for ca. 90% of the carbohydrates in plants [246]. Studies have demonstrated that polysaccharides in medicinal plants are the indispensable bioactive compounds presenting uniquely pharmacological effects such as immunomodulatory, hypoglycemic, antitumor, anti-diabetic, and antioxidant, amongst others, with almost no side effect or adverse drug reaction [247,248]. To date, polysaccharides in the 228 AMPs have also been identified to show multiple pharmacological effects. For example, polysaccharides in Angelica sinensis present the effect of hematopoietic, antitumor, and liver protection, etc., [238,249]; polysaccharides in Angelica dahurica protect the effect of spleen lymphocytes, natural killer cells, and procoagulant, etc., [250,251]; as well as polysaccharides in Bupleurum chinense and Bupleurum smithii present the effect of macrophage modulation, kidney protection, and inflammatory alleviation, etc., [252,253,254].

7.2. Alkaloids

About 27 000 alkaloids presenting as water-soluble salts of organic acids, esters, and combined with tannins or sugars have been found in plants [255]. Many alkaloids are valuable medicinal agents that can be utilized to treat various diseases including malaria, diabetes, cancer, cardiac dysfunction, and blood clotting related diseases, etc., [256,257,258]. While alkaloids in the 228 AMPs mainly exist in the Ligusticum, Apium, Conium, and Cuminum genera [245]. Pharmacological studies demonstrated that alkaloids in Ligusticum chuanxiong show the activity of inhibiting myocardial fibrosis, protecting ischemic myocardium, and relieving cerebral ischemia-reperfusion injury [150,259,260]; and a novel alkaloid 2-pentylpiperidine named as conmaculatin in Conium maculatum shows strong peripheral and central antinociceptive activity [261]. While some alkaloids have been verified to show antidepressant activity, such as berberine in Berberis aristata, strictosidine acid in Psychotria myriantha, and Anonaine in Annona cherimolia, which could be explored as an emerging therapeutic alternative for the treatment of depression of AMPs.

7.3. Phenylpropanoids

Phenylpropanoids are a large class of secondary metabolites biosynthesized from the amino acids, phenylalanine, and tyrosine [262]. Over 8000 aromatic metabolites of the phenylpropanoids that have been identified in plants include simple phenylpropanoids (propenyl benzene, phenylpropionic acid, and phenylpropyl alcohol), coumarins, lignins, lignans, and flavonoids [263].

7.3.1. Simple Phenylpropanoids

To date, limited simple phenylpropanoids have been identified from the AMPs, such as 3 phenylpropanoids (e.g., trans-isoelemicin, sarisan, and trans-isomyristicin) existed in roots of Ligusticum mutellina [264]; and ferulic acid, one of the phenylpropionic acids, as an important bioactive metabolite of AMPs had many activities, mainly existed in the Angelica, Ligusticum, Ferula, and Pleurospermum genera [238,265,266]. Pharmacological studies demonstrated that ferulic acid in Angelica sinensis shows strong properties in inhibiting platelet aggregation, increasing coronary blood flow, and stimulating smooth muscle [267,268]; ferulic acid in Angelica acutiloba shows antidiabetic effects, immunostimulant properties, antiinfammatory, antimicrobial, anti-arrhythmic, and antithrombotic activity [269]; and ferulic acid in Ligusticum tenuissimum shows anti-melanogenic and anti-oxidative effects [270].

7.3.2. Coumarins

Coumarins are the most widespread in 20 genera of AMPs (e.g., Angelica, Bupleurum, and Peucedanum) and mainly include simple coumarins, pyranocoumarins, and furocoumarins [56,271,272]. In recent years, distinct coumarins have been identified from the AMPs, such as 99 coumarins in Ferula [273], 116 coumarins in Angelica decursiva and Peucedanum praeruptorum [179], as well as 9 coumarins in Angelica dahurica [274]. Furthermore, 8 coumarins have been selected as quality markers including cnidiadin (1) in Angelica biserrata and Cnidium monnieri, zosimin (2) in Angelica biserrata, imperatorin (3) in Angelica dahurica and Angelica dahurica cv. Hangbaizhi, isoimperatorin (4) in Angelica dahurica, Angelica dahurica cv. Hangbaizhi, Notopterygium franchetii, and Notopterygium incisum, nodakenin (5) in Angelica decursiva, Notopterygium franchetii, and Notopterygium incisum, notopterol (14) in Notopterygium franchetii and Notopterygium incisum, as well as praeruptorin A (15) and praeruptorin B (16) in Peucedanum praeruptorum (Table 2 and Figure 5) [18].
To date, various biological activities of coumarins have been demonstrated including antifungal, antimicrobial, antiviral, anti-cancerous, antitumor, anti-inflammatory, anti-filarial, enzyme inhibitors, antiaflatoxigenic, analgesics, antioxidant, and oestrogenic [275,276,277,278]. For example, coumarins are recognized as the main bioactive constituents in Peucedani genus and play critical roles in relieving cough and asthma, strengthening heart function, as well as preventing and treating cardiovascular diseases such as nodakenin, (+)-praeruptorin B, and praeruptorin C [279]; imperatorin oxypeucedanin hydrate, xanthotoxol, bergaptol, 5-methoxy-8-hydroxypsoralen, isoimperatorin, phelloptorin, and pabularinone in Angelica dahurica exhibited moderate DPPH•scavenging activity, strong ABTS·+ scavenging activity, and significant inhibition on HepG2 cells, which could be explored as new and potential natural antioxidants and cancer prevention agents [30]; pabulenol and osthol extract in Angelica genuflexa show anti-platelet and anti-coagulant components [38]; decursinol angelate in Angelica gigas shows platelet aggregation and blood coagulation activity [38].

7.4. Flavonoids

Flavonoids are a group of the most abundant secondary metabolites in plants [262]. Generally, flavonoids can be further categorized into 8 subgroups including: flavones (e.g., apigenin, luteolin, and baicalein), flavonols (e.g., kaempferol, quercetin, and myricetin), flavanones (e.g., naringenin, hesperitin, and liquiritigenin), flavanonols (e.g., dihydrokaempferol, dihydromyricetin, and dihydroquercetin), isoflavones (e.g., daidzein, purerarin, and peterocarpin), aurones, anthocyanidins, and proanthocyanidins [280,281,282]. In recent years, flavonoids have been identified from the AMPs, such as 6 flavonoids (e.g., luteolin, isoquercitrin, and rutin) in Ferula [107], 12 flavonoids (e.g., quercetin-3-O-rutinoside, kaempferol-3,7-di-O-rhamnoside, quercetin-3-O-arabinoside) in Bupleurum [283], and 18 flavonoids (e.g., rutin, quercetin, and quercitrin) in Hydrocotyle [134].
To date, various biological activities of flavonoids have been demonstrated including antioxidant, antiinflammatory, antidiabetic, anticancer, antiobesity, and cardioprotective [280,284]. For example, apigenin in Apium graveolens shows anticancer property [21], and flavonoids in Pimpinella diversifolia DC., Anthriscus sylvestris, and Sanicula astrantiifolia show antioxidant effect [196,285]; as well as quercetin and its metabolites show vasodilator effects with selectivity toward the resistance vessels [286].

7.5. Terpenoids

About 25 000 terpenoids have been reported in plants and they are most diverse secondary metabolites containing three subgroups including: monoterpenoids, sesquiterpenes, and triterpenoids [287]. Actually, terpenoids have been also identified from the AMPs, such as 4 terpenoids (e.g., angelicoidenol, pregnenolone, and β-sitosterol) in Pleurospermum [141], 75 terpenoids (e.g., myrcene, farnesene, and xiongterpene) in Ligusticum [140], 109 terpenoids (e.g., nerolidol, guaiol, and ferulactone A) in Ferula [273], and 13 triterpenoids (e.g., ranuncoside, oleanane, and barrigenol) in Hydrocotyle sibthorpioides Lam. [135].
Studies have found that terpenoids possess various biological activities such as anti-inflammatory, anti-oxidation, and anti-fibrosis activities, antitumor, anti-Alzheimer’s disease, and anti-depression [288,289]. For example, xiongterpene in Ligusticum chuanxiong shows insecticide effect [150], asiaticoside in Centella asiatica shows antitumor property [290], as well as saikosaponin d in Bupleurum chinense DC. and Bupleurum scorzonerifolium show the effect of reducing blood glucose, inhibiting inflammation, and reducing insulin resistance [291].

7.6. Other Compounds

Chromones and phthalides also exist in the AMPs and show pharmacological properties. Specifically, phthalides (e.g., ligustilide, n-butylidenephthalide, and Z-ligustilide) in the Angelica sinensis show the effect of inhibiting vasodilation, decreasing platelet aggregation, as well as exerting analgesic, anti-inflammatory, and anti-proliferative [238]; butylphthalide in Ligusticum sinense shows the effect of anti-inflammatory, antithrombus, dilate blood vessels, improve brain microcirculation, and anti-myocardial ischemia [154].
For the chromones, 3 chromones [i.e., 5 thydroxy 2 [(angebyloxy) mehyI] fuan [3, 2’: 6, 7] chrmone, angeliticin A, and noreugenin)] in Angelica polymorpha [292], 10 chromones (e.g., cnidimoside A, cnidimol B, and peucenin) in Cnidiummonnieri (L.) Cuss. [93], and 22 chromones [e.g., edebouriellol, hamaudol, and 3ʹ(R)-(+)-hamaudol] in Saposhnikovia divaricate [217] have been identified. Studies have found that 2 chromones 3ʹS-(-)-O-acetylhamaudol and (±)-hamaudol in Angelica morii show the effect of inhibiting Ca2+ influx of vascular smooth muscle [293], prim-O-glucosylcimifugin and 5-O-methylvisammioside show the effect of antipyretic, analgesic, and anti-inflammatory [294], and chromones in Bupleurum multinerve shows the analgesic effect [295].

8. Effect of Bolting and Flowering (BF) on Yield and Quality

Previous literatures have repeatedly emphasized that the BF reduces the yield and quality of plants, especially in rhizomatous medicinal plants [11]. Here, a total of 38 rhizomatous plants reported in the 228 AMPs are associated with the BF (Table 3). Based on the effect degree of the BF on the yield and quality, the 38 rhizomatous AMPs belonging to 17 genera can be categorized into 3 classes including: (1) the BF significantly affects the yield and quality of 14 AMPs (i.e., Angelica acutiloba, Angelica biserrata, Angelica dahurica, Angelica dahurica cv. Hangbaizhi, Angelica decursiva, Angelica polymorpha, Angelica sinensis, Daucus carota, Heracleum hemsleyanum, Heracleum rapula, Libanotis iliensis, Libanotis seseloides, Peucedanum praeruptorum, and Saposhnikovia divaricata), and their rhizomes and/or roots are wholly lignified and absolutely useless for clinical effects; (2) the BF affects the yield of 11 AMPs (i.e., Angelica gigas, Bupleurum chinense, Bupleurum scorzonerifolium, Changium smyrnioides, Chuanminshen violaceum, Glehnia littoralis, Ligusticum chuanxiong, Ligusticum jeholense, Ligusticum sinense, Notopterygium franchetii, and Notopterygium incisum), while their rhizomes or roots can be used as medicine to some extent; as well as (3) the BF has no significant effect on the yield and quality of 13 AMPs (i.e., Angelica sylvestris, Cicuta virosa, Ferula ferulaeoides, Ferula fukanensis, Ferula lehmannii, Ferula olivacea, Ferula sinkiangensis, Ferula teterrima, Levisticum officinale, Libanotis buchtormensis, Libanotis lancifolia, Libanotis spodotrichoma, and Pimpinella candolleana), and their rhizomes or roots are still used as medicine (Figure 6).
For the class (1), bolting and flowering reduce the yield and contents of bioactive compounds of plants with none or almost no medicinal value, representatively, a 8.3- and 16.1-fold reduction of dry weight and quality marker ferulic acid content in Angelica sinensis [296]; and a 1.5- and 1.5-fold reduction of dry weight and quality marker isoimperatorin content in Angelica dahurica [297] have been observed after the BF. For the class (2), bolting and flowering reduce the yield and contents of bioactive compounds of plants with little medicinal value, representatively, a 1.34-fold reduction of saikosaponinsands, while no significant change of dry weight in Bupleurum chinense [298,299]; and a 2.0- and 1.7-fold reduction of dry weigh and polysaccharides content in Changium smyrnioides [300] have been observed after the BF. For the class (3), the yield and quality of the 13 AMPs are not affected after bolting and flowering by the harvest stages [19].

9. Approaches to Control the BF

Generally, Most Apiaceae plants are “low-temperature and long-day” perennial herbs, in other words, the plants must experience vernalization (i.e., an extended period of cool weather at 0 to 10℃) and long days (> 12 h daylight) to induce the BF, such as Angelica sinensis [320], Daucus carota [321], and Coriandrum sativum [322].
As shown in Table 4, approaches to inhibit the BF of 24 AMPs have been listed. For example, the bolting rate of Angelica sinensis can be significantly decreased through planting the green stem cultivar (Mingui 2) instead of the purple stem cultivar (Mingui 1) [323], selecting smaller seedlings (i.e., root-shoulder diameter <0.55 cm) instead of larger seedlings [324,325], storing the seedlings at freezing temperature (i.e., <0℃ ) during overwinter stage [320], shading the plants under sunshade (i.e., >40%) during growth stage [326], and providing the plants with good growth conditions (e.g., plant intensity, nutrient and water balance) [327]. The bolting rate of Angelica dahurica can be significantly decreased through planting the purebreeds [328], selecting the immature seeds for seeding [303], increasing the potassic fertilizer while decreasing the nitrogen and phosphorus fertilizers [329], and planting with standard techniques [330]. The bolting rate of Saposhnikovia divaricata can also be significantly decreased through controlling the sunshade [331], sowing date [332], planting density [333], and preventing the excessive growth [331].
To inhibit the occurrence of BF of AMPs, plenty of measures that can be used include: breeding new cultivars to reduce the BF, controlling the seedling age and size to delay the transition from vegetative growth to flowering, storing seedlings at freezing temperatures to avoid vernalization, growing the plants under sunshade to avoid long-day photoperiodism, and planting with standard techniques to reduce pests and diseases (Figure 7).

10. The Mechanism of BF Inducing the Rhizome Lignification

Extensive experiments have demonstrated that the BF induces the lignification of fleshy rhizomes, meanwhile, enhances the degradation of bioactive metabolites [11,13,323]. Studies on anatomical structures reveal that the ratio of secondary phloem to secondary xylem respectively changes from 2:1 to 1:10 and 2/5-1/2 to 1/2-3/4 for the rhizomes of Angelica sinensis and Angelica dahurica before and after BF, meanwhile, the number of secretory cells producing essential oils significantly decreased [363,364]. Studies have been found that EARLY BOLTING IN SHORT DAY (EBS) acts as a negative transcriptional regulator preventing premature flowering of Arabidopsis thaliana and have been observed as co-enrichment of a subset of EBS-associated genes with H3K4me3, H3K27me3, and Polycomb repressor complex 2 [365]; a potential genetic resource for radish late-bolting breeding with introgression of the RsVRN1In-536 insertion allele into early-bolting genotype could contribute to delay bolting time of Raphanus sativus [366]; and peroxidases (PRXs) involved in lignin monomers biosynthesis were downregulated in Peucedanum praeruptorum at the bolting stage [367].
As known, lignin biosynthesis belongs to the general phenylpropanoid pathway, which starts from phenylalanine and is catalyzed by a serial of enzymes [13,368]. Specifically, phenylalanine is catalyzed to form p-Coumaroyl CoA sequentially through the 3 enzymes phenylalanine ammonia lyase (PAL), cinnamate 4-hydroxylase (C4H), and 4-coumarate-CoA ligase (4CL); lignin biosynthesis is synthesized via 3 sub-pathways including: (1) lignins are catalyzed to from p-Coumaroyl CoA sequentially through the three enzymes cinnamoyl-CoA reductases (CCR), cinnamyl alcohol dehydrogenases (CAD), and laccases (LACs); and then coniferyl aldehyde is catalyzed to from p-Coumaroyl CoA sequentially through the 4 enzymes hydroxycinnamoyl shikimate/quinate transferase (HCT), p-coumarate 3-hydroxylase (C3H), caffeoyl-CoA 3-O-methyltransferase (CCOMT), and CCR; (2) lignins are catalyzed to from coniferyl aldehyde sequentially through the 2 enzymes CAD and LACs; as well as (3) lignins are catalyzed to from coniferyl aldehyde sequentially through the 3 enzymes ferulate 5-hydroxylase (F5H), caffeic acid 3-O-methyltransferase (COMT), and LACs (Figure 8).
Although lignin biosynthesis has been depicted, studies on the mechanism of BF inducing rhizome lignification are still limited. To our knowledge, only the mechanism of BF affecting Angelica sinensis has been conducted, with the expression level of genes (e.g., PAL1, 4CLs, HCT, CAD1, and LACs) significantly upregulated at the stem-node forming and elongating stage compared with stem-node pre-differentiation stage, leading to the reduction of accumulation of bioactive metabolites (i.e., ferulic acid and flavonoids) [13].

11. Conclusions and Future Aspect

In this review, we summarized the tour of AMPs, classification of AMPs species, traditional use, modern pharmacological use, phytochemistry, effect of BF on yield and quality, approach to control the BF, and the mechanism of BF inducing the rhizome lignification. Although ca. 228 AMPs, 72 traditional uses, 62 modern uses, and 5 main kinds of metabolites have been recorded, the potential properties still need to be exploited. Although the urgent problems in the BF significantly reducing the yield and quality of AMPs have been found and several approaches have been applied in controlling the BF, the effective measures to inhibit the BF and its mechanism have not been systemically revealed. Thus, in order to effectively control the BF of AMPs, on one hand, standard cultivation techniques of AMPs should be applied; on the other hand, new cultivars should be innovated by the modern biotechnology such as the CRISPR/Cas9 system.

Author Contributions

collected and analyzed the references, drew the chemical structures, and wrote the manuscript, M.L.L.; checked the classification and traditional use of Apiaceae medicinal plants, M.L.; checked the language and modern pharmacological use, L.W.; Conceptualization, Methodology, Supervision, Writing-review and editing, M.F.L.; Conceptualization and Project administration, J.W.

Funding

This research was funded by the National Natural Science Foundation of China (32160083), earmarked fund for CARS (CARS-21), and Gansu Education Science and Technology Innovation Project (2022CXZXS-022).

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. A tour of Apiaceae medicinal plants (AMPs).
Figure 1. A tour of Apiaceae medicinal plants (AMPs).
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Figure 2. Traditional use of the 228 AMPs.
Figure 2. Traditional use of the 228 AMPs.
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Figure 3. Modern pharmacological uses of the 228 AMPs.
Figure 3. Modern pharmacological uses of the 228 AMPs.
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Figure 4. Core structures of five different bioactive compounds identified from the 228 AMPs.
Figure 4. Core structures of five different bioactive compounds identified from the 228 AMPs.
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Figure 5. Structures of 18 quality markers from the 22 AMPs in Pharmacopoeia of the People’s Republic of China (2020).
Figure 5. Structures of 18 quality markers from the 22 AMPs in Pharmacopoeia of the People’s Republic of China (2020).
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Figure 6. Cluster of the 38 rhizomatous AMPs affected by the bolting and flowering. The red color shows that the BF significantly affects the yield and quality; the yellow color shows that the BF differently affects the yield while the rhizomes or roots can be used as medicine to some extent; and the green color shows that the BF has no significant effect on the yield and quality. Different lower case letters a: Angelica, b: Ferula, c: Libanotis, d: Ligusticum, e: Heracleum, f: Notopterygium, g: Bupleurum, h: Changium, i: Peucedanum, j: Saposhnikovia, k: Glehnia, l: Cicuta, m: Daucus, n: Levisticum, o: Anthriscus, p: Chuanminshen, and q: Pimpinella.
Figure 6. Cluster of the 38 rhizomatous AMPs affected by the bolting and flowering. The red color shows that the BF significantly affects the yield and quality; the yellow color shows that the BF differently affects the yield while the rhizomes or roots can be used as medicine to some extent; and the green color shows that the BF has no significant effect on the yield and quality. Different lower case letters a: Angelica, b: Ferula, c: Libanotis, d: Ligusticum, e: Heracleum, f: Notopterygium, g: Bupleurum, h: Changium, i: Peucedanum, j: Saposhnikovia, k: Glehnia, l: Cicuta, m: Daucus, n: Levisticum, o: Anthriscus, p: Chuanminshen, and q: Pimpinella.
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Figure 7. Approach to control the BF of AMPs.
Figure 7. Approach to control the BF of AMPs.
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Figure 8. Schematic representation of biosynthetic pathways of lignins. Abbreviations: PAL, phenylalanine ammonia lyase; C4H, cinnamate 4-hydroxylase; 4CL, 4-coumarate-CoA ligase; HCT, hydroxycinnamoyl shikimate/quinate transferase; C3H, p-coumarate 3-hydroxylase; CCOMT, caffeoyl-CoA 3-O-methyltransferase; CCR, cinnamoyl-CoA reductases; CAD, cinnamyl alcohol dehydrogenases; LACs, laccases; F5H, ferulate 5-hydroxylase; COMT, caffeic acid 3-O-methyltransferase. The green color shows the common phenylpropanoid pathway of phenylpropanoids, and the red color shows the lignin biosynthetic sub-pathway.
Figure 8. Schematic representation of biosynthetic pathways of lignins. Abbreviations: PAL, phenylalanine ammonia lyase; C4H, cinnamate 4-hydroxylase; 4CL, 4-coumarate-CoA ligase; HCT, hydroxycinnamoyl shikimate/quinate transferase; C3H, p-coumarate 3-hydroxylase; CCOMT, caffeoyl-CoA 3-O-methyltransferase; CCR, cinnamoyl-CoA reductases; CAD, cinnamyl alcohol dehydrogenases; LACs, laccases; F5H, ferulate 5-hydroxylase; COMT, caffeic acid 3-O-methyltransferase. The green color shows the common phenylpropanoid pathway of phenylpropanoids, and the red color shows the lignin biosynthetic sub-pathway.
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Table 1. The list of the 228 AMPs.
Table 1. The list of the 228 AMPs.
No. Plant species Parts of plant used Traditional use Modern pharmacological use Main chemical constituents References
1 Aegopodium alpestre Ledeb. Stems and leaves Dispelling wind, relieving pain, and treatment of influenza Treatment of rheumatic diseases, obesity and hypotensive Apiole, undecane, and limonene [19,20,21]
2 Ammi majus L. Fruits Treatment of vitiligo \ Furanocoumarins [16]
3 Anethum graveolens L. Fruits, leaves or whole plant Treatment of bladder inflammation, liver diseases, and insomnia Antibacterial, antifungal, antioxidant Alkaloid, terpenoids, and flavonoids [22]
4 Angelica acutiloba (Siebold & Zucc.) Kitag. Roots Treatment of menoxenia and anemia Hemogenic, analgesic, and sedative activities Ferulic acid, ligustilide, and angelicide [23]
5 Angelica amurensis Schischk. Roots \ \ α-pinene, limonene, and sabinene [1,24]
6 Angelica anomala Avé-Lall. Roots Dispelling wind, eliminating dampness, and relieving pain Antioxidant, anti-inflammatory, and antitumor Isoimperatorin, umbelliferone, and adenosine [16,25,26,27]
7 Angelica apaensis R. H. Shan & C. C. Yuan Roots Relieving pain, relieving cough and asthma Bacteriostat, anti-inflammatory Oxypeucedanin, isoimperatorin, and oxypeucedanin hydrate [19,28]
8 **Angelica biserrata (R. H. Shan & C. C. Yuan) C. C. Yuan & R. H. Shan Roots Dispelling wind, eliminating dampness, and relieving pain Antitumor, anti-inflammatory, and antioxidant Coumarins osthole, columbianadin, and volatile oils [29]
9 Angelica cartilaginomarginata var. Foliosa C. C. Yuan & R. H. Shan Roots \ \ \ [17]
10 **Angelica dahurica (Fisch. Ex Hoffm.) Benth. & Hook. F. Ex Franch. & Sav. Roots Treatment of acne, erythema, and headache Antiinflammatory, anti-mutagenic, and antitumor Scopoletin, and psoralen [18,30,31,32,33]
11 **Angelica dahurica cv. Hangbaizhi Roots Treatment of headache, toothache, abscess, and furunculosis Estrogenic, cytotoxic, and anti-inflammatory isoimperatorin, imperatorin, and phellopterin [18,34,35]
12 Angelica dahurica var. Formosana (H. Boissieu) Yen Roots \ Anti-staphylococca Falcarindiol [33,34]
13 **Angelica decursiva (Miq.) Franch. & Sav. Roots A remedy for thick phlegm, asthma, and upper respiratory tract infections Antioxidant and anti-inflammatory potential Decursin, decursidin, and nodakenetin [36]
14 Angelica gigas Nakai Roots Treatment of dysmenorrhea, amenorrhea, and menopausal Anti-platelet effects Decursin, and decursinol angelate [37,38]
15 Angelica laxifoliata Diels Roots Dispelling wind, Dispelling wind, and relieving pain Treatment of wind-damp pain, aching lumbus and knees Angelicin, β-sitosterol, and laxifolin [16,26,39]
16 Angelica megaphylla Diels Roots Same as Angelica sinensis Same as A. Sinensis Ferulic acid, ligustilide, and angelol [40,41]
17 Angelica morii Hayata Roots and leaves Treatment of deficiency-cold in spleen and stomach, cold cough, and toothache Used for diarrhea caused by deficiency of spleen and for cough caused by weak-ness and chill Imperatorin, isoimperatorin, and phellopterin [42,43,44]
18 Angelica nitida H. Wolff Roots Nourishing the blood, regulating menstrual disorder, and relieving pain \ Isoimperatorin, imperatorin, and cnidilin [45]
19 Angelica polymorpha Maxim. Roots Dispelling wind and relieving pain Treatment of stomachache Coumarins, sesquiterpenoids, and alkaloid [19,46,47]
20 **Angelica sinensis (Oliv.) Diels Roots Nourishing the blood, regulating menstrual disorder, and relieving pain Cardio-cerebrovascular, anti-inflammatory, and antioxidant Ferulic acid, alkylphthalides, and polysaccharides [18,48,49]
21 Angelica sinensis var. Wilsonii Roots Same as Angelica sinensis, relieving pain Same as Angelica sinensis Isoimperatorin, coumarin, and oxypeucedanin [50]
22 Angelica sylvestris L. Roots Relieves rheumatism, sweating, and detoxification \ Cnidilide, sedanenolide, and ligustilide [19]
23 Angelica tsinlingensis K. T. Fu Roots \ \ \ [1]
24 Angelica valida Diels Roots \ \ \ [1]
25 Anthriscus nemorosa (M. Bieb.) Spreng. Roots, whole plant, and leaves Same as Peucedanum praeruptorum Same as Peucedanum praeruptorum \ [51]
26 Anthriscus sylvestris (L.) Hoffm. Roots and leaves Invigorating spleen and replenishing qi and expelling phlegm Antitumor, antioxidation, and antisenity Phenylpropanoids, flavonoids, and steroidal [19,52]
27 Apium graveolens L. Whole plant, roots, and rhizome Dispelling wind, eliminating dampness, and detoxification Hypertension, hyperlipidemia, and dysuria Organic acids, apigenin, and volatile oils [19,53,54]
28 Archangelica brevicaulisf Roots Same as Angelica biserrata Same as Angelica biserrata Osthol, imperatorin, and archangelicin [16,55]
29 Bupleurum angustissimum (Franch.) Kitag. Roots \ \ Saikosaponins (a, c, and d), β -terpinene, and β -thujene [56]
30 Bupleurum aureum Fisch. Roots \ \ Saikosaponins (a, c, and d) [1,57]
31 Bupleurum bicaule Helm Roots Same as Bupleurum scorzonerifolium Same as Bupleurum scorzonerifolium Saikosaponin d, prosaikogenin G, and prosaikogenin F [16,58,59]
32 Bupleurum candollei Wall. Ex DC. Whole plant Diminish inflammati and detoxify, dispelling wind, and relieving convulsion \ Saikosaponin and flavonoids [16,56]
33 Bupleurum chaishoui R. H. Shan & M. L. Sheh Roots and rhizome Same as Bupleurum Same as Bupleurum Saikosaponins (a, c, and d) [60]
34 **Bupleurum chinense DC. Roots Treatment of chronic hepatitis, kidney syndrome, and inflammatory diseases Anti-allergic, analgesic, and anti-inflammation Saikosaponins (a, c, and d) [18,61,62]
35 Bupleurum chinense DC. F. Octoradiatum (Bunge) Shan et Sheh Roots Same as Bupleurum Anti-allergic, analgesic, and anti-inflammation Saikosaponins (a, c, and d) [63,64]
36 Bupleurum chinense DC. F. Vanheurckii (Muell. -Arg.) Shan et Y. Li Roots Same as Bupleurum Anti-allergic, analgesic, and anti-inflammation Saikosaponins (a, c, and d) [63,64]
37 Bupleurum commelynoideum var. Flaviflorum R. H. Shan & Yin Li Roots, rhizome and whole plant Antipyretic-analgesic effect, choleretic, and hepatoprotection Treating or relieving inflammatory bowel disease Saikosaponins (a, c, and d), β-pinene, and perillen [65,66]
38 Bupleurum densiflorum Rupr. Roots \ \ \ [63]
39 Bupleurum dielsianum H. Wolff Roots \ \ \ [63]
40 Bupleurum euphorbioides Nakai Roots \ \ Saikosaponins, perillen, and undecanal [56]
41 Bupleurum exaltatum M. Bieb. Roots \ \ \ [64]
42 Bupleurum falcatum L. Roots \ Treatment of colds and upper respiratory tract infections Saikosaponins (a, c, and d) [64,67,68]
43 Bupleurum gansuense S. L. Pan et Hsu Roots \ \ \ [56]
44 Bupleurum hamiltonii N. P. Balakr. Roots or whole plant Antipyretic-analgesic effect, treatment of chill, and fever alternation Treatment of stomach pain, dysuria, and cough Kaerophyllin, isokaerophyllin, and ethyl caffeic acid [69]
45 Bupleurum hamiltonii var. Hamiltonii / Bupleurum tenue Roots or whole plant Same as Bupleurum hamiltonii N. P. Balakr. Same as Bupleurum hamiltonii N. P. Balakr. Same as Bupleurum hamiltonii N. P. Balakr. [70]
46 Bupleurum hamiltonii var. Humile (Franch.) R. H. Shan & M. L. Sheh Roots \ \ \ [64]
47 Bupleurum huizei S. L. Pan sp. Nov. Roots \ \ \ [64]
48 Bupleurum kaoi T. S. Liu, C. Y. Chao & T. I. Chuang Roots \ Treatment of influenza and fever Saikosaponin a and saikosaponin c [64]
49 Bupleurum komarovianum Lincz. Roots Same as Bupleurum chinense Same as Bupleurum chinense Saikosaponins (a, c, and d) and volatile oils (1-caprylene, limonene, and thymol) [71,72]
50 Bupleurum krylovianum Schischk. Ex Krylov Roots \ \ Saikosaponins (a, c, and d) [56,57]
51 Bupleurum kunmingense Yin Li & S. L. Pan Roots \ Immunomodulatory Saikosaponins (a, c, and d), cyclohexanone, and 2- methyldodecane [56]
52 Bupleurum longicaule var. Amplexicaule C. Y. Wu Roots \ \ Saikosaponins (a, c, and d) [64]
53 Bupleurum longicaule var. Franchetii H. Boissieu Roots or whole plant \ \ Saikosaponins (a, c, and d), cyclohexanone, and myrcene [56]
54 Bupleurum longicaule var. Giraldii H. Wolff Roots \ \ Saikosaponins (a, c, and d), narcissin, and rutin [56]
55 Bupleurum longiradiatum Turcz. Roots Treatment of gout and inflammatory illness Anti-inflammatory and/or antimicrobial Thymol, butylidene phthalide and 5-indolol [73]
56 Bupleurum luxiense Yin Li & S. L. Pan Roots \ \ Saikosaponins (a, c, and d), n-heptaldehyde, and octanal [56]
57 Bupleurum malconense R. H. Shan & Yin Li Whole plant Hepatoprotection and antipyretic effect Acute toxicity Saikosaponins (a, c, and d), rutin, and quercetin [74,75,76]
58 Bupleurum marginatum var. Marginatum Whole plant Hepatoprotection and antipyretic effect Anti-allergic, analgesic, and anti-inflammatory Saikosaponins (a, c, and d), rutin, and quercetin [74,75,77]
59 Bupleurum marginatum var. Stenophyllum (H. Wolff) R. H. Shan & Yin Li Whole plant \ \ Saikosaponins (a, c, and d), chikusaikoside I, II, and 2- methylcyclopentanone [56]
60 Bupleurum marginatum Wall. Ex DC. Whole plant and roots Hepatoprotection and antipyretic effect Anti-allergic, analgesic, and anti-inflammatory Saikosaponins (a, c, and d), rutin, and quercetin [74,75,77]
61 Bupleurum microcephalum Diels Whole plant and roots Hepatoprotection and antipyretic effect Anti-allergic, analgesic, and anti-inflammatory Saikosaponins (a, c, and d), rutin, and quercetin [74,75]
62 Bupleurum petiolulatum var. tenerum R. H. Shan & Yin Li Whole plant Antipyretic-analgesic effect Anti-inflammatory \ [63,78]
63 Bupleurum polyclonum Yin Li & S. L. Pan Roots \ Anticancer Saikosaponins (a, c, and d), 4’-O-saikosaponin-a, and fenchane [56]
64 Bupleurum rockii H. Wolff Roots \ \ Saikosaponins (a, c, and d), thymol, and β-guaiene [56]
65 Bupleurum scorzonerifolium f. Longiradiatum Roots Same as Bupleurum Same as Bupleurum Same as Bupleurum [19]
66 Bupleurum scorzonerifolium f. Pauciflorum Roots Same as Bupleurum Same as Bupleurum Same as Bupleurum [19]
67 **Bupleurum scorzonerifolium Willd. Roots Antipyresis, relieve liver depression and menstrual disorder Same as Bupleurum chinense Rutin, quercetin, and kaempferol [18,19]
68 Bupleurum sibiricum var. Jeholense (Nakai) Y. C. Chu ex R. H. Shan & Yin Li Roots \ \ \ [1]
69 Bupleurum sibiricum Vest Roots Same as Bupleurum Same as Bupleurum Saikosaponin a, rutin, and quercetin [16,79,80]
70 Bupleurum sichuanense S. L. Pan et Hsu. Roots \ \ Saikosaponins (a, c, and d) [56]
71 Bupleurum smithii H. Wolff Roots Antipyretic-analgesic effect Anti-inflammatory, immunomodulatory, and anti-hepatic injury Saponins, volatile oils, and lignans [81]
72 Bupleurum smithii var. Parvifolium R. H. Shan & Yin Li Roots Relieve liver depression and activate the yang-energy Anti-inflammatory, immunomodulatory, and antitumor falcarinol, saponins, and flavonoids [82]
73 Bupleurum thianschanicum Freyn Roots \ \ Saikosaponins (a, c, and d) [57]
74 Bupleurum triradiatum Adams ex Hoffm. Roots \ \ \ [1]
75 Bupleurum wenchuanense R. H. Shan & Yin Li Roots Same as Bupleurum Same as Bupleurum Quercetin-3-O-α-L-rhamnoside, quercetin, and rutin [16,75]
76 Bupleurum yinchowense R. H. Shan & Yin Li Roots Antipyresis, relieve liver depression, and activate the yang-energy Same as Bupleurum Saikosaponins (a, c, and d) [16,65,83,84]
77 Carum buriaticum Turcz. Roots and fruits \ \ \ [5]
78 Carum carvi L. Roots, fruits, and leaves Dispelling wind and eliminating dampness, invigorate the stomach, and treatment of heart disease Anti-bacterial, antioxidant, and antitumor Carvone, limonene, and dihydrocarvone [19,85,86]
79 *Centella asiatica (L.) Urb. Whole plant Clearing heat, promoting diuresis, and toxicity Anti-bacterial, anti-depression and neuroprotection Asiaticoside, madecassoside, and elemene [18,87]
80 **Changium smyrnioides H. Wolff Roots Strengthening with tonics, moistening lung melt phlegm, and calm the liver Immunomodulatory, relieve fatigue, and enhance adaptability Cetylic acid, succinic acid, and imperatorin [18,88]
81 Chuanminshen violaceum M. L. Sheh & R. H. Shan Roots Moistening lung melt phlegm, harmonize the stomach, and engender liquid Antioxidant, enhancing immunity, and antimutation Polysaccharides, coumarins, and flavonoids [89,90,91]
82 Cicuta virosa L. Roots and rhizome Expelling phlegm and detoxification Treatment of osteomyelitis, gout, and rheumatism P-cymene, cicutoxine, and L-limonene [17,92]
83 *Cnidium monnieri (L.) Spreng. Fruits Dispelling wind, relieving convulsion, and Impotence Antibacterial, antiviral, and antimutagenesis Osthole, limonene, and cnidimoside A [18,93]
84 Cnidium officinale Roots Same as Cnidium monnieri Same as Cnidium monnieri \ [1]
85 Conioselinum acuminatum (Franch.) Lavrova Roots \ \ Sabinene, α-pinene, and aromadendrene [11]
86 Conioselinum anthriscoides ’Fuxiong’ Roots \ \ β-bergamotene [11]
87 Conioselinum tenuisectum (H. Boissieu) Pimenov & Kljuykov Roots \ \ \ [94]
88 Conioselinum vaginatum (Spreng.) Thell. Roots Dispelling wind, eliminating dampness, and relieving pain Treatment of common cold due to wind-cold and gastro spasm Diligustilide, daucosterol, and palmitic acid [19,95]
89 Conium maculatum L. Whole plant Relieving pain and relieving muscular spasm Treatment of cancer Coniine, N-methyl-coniine, conhydrine 2-(1-hydroxypropyl)-piperidine [16,96,97]
90 Coriandrum sativum L. Whole plant, fruits, and stems Invigorate the stomach and promoting eruption Antibacterial, antifungal, and antioxidant Petroselinic acid, linoleic acid, and oleic acid [19,98]
91 Cryptotaenia japonica Hassk. Whole plant Treatment of weakness, urinary closure, and swelling Antioxidant, protect liver, and anticancer Friedelin, stigmasterol, and apigenin [19,99,100]
92 Cuminum cyminum L. Fruits Treatment of indigestion and stomach cold abdominal pain Antibacterial, antioxidant, and radical-scavenging properties α-pinene, 1,8-cineole, and linalool [19,101]
93 Cyclorhiza peucedanifolia (Franch.) Constance Fruits Enriching the blood, activating blood, and regulating menstrual disorder \ \ [102]
94 Daucus carota L. Fruits Treatment of ascariasis, enterobiasis, and tapeworm disease Insecticide, anti-bacterial, and anticancer α-pinene, isophorone oxide, and and quercetrin [18,103]
95 Daucus carotavar. Carota Fruits Treatment of ascariasis, enterobiasis, and tapeworm disease Insecticide, anti-bacterial, and anticancer α-pinene, β-bisabolene , and luteolin [18,103]
96 Daucus carota var. Sativus Hoffm. Roots and basal leaves Strengthening spleen, treatment of dyspepsia, and chronic dysentery Enhancing immunity, anticancer, and prevents aging Carotene, (1R)-α-pinene, and β-carotene [19,104]
97 Eriocycla albescens (Franch.) H. Wolff Roots \ \ \ [1]
98 Eryngium foetidum L. Whole plant Diuresis, treatment of dropsy, and snakebite Bacteriostat, diminish inflammati, and detumescence Lanolin alcohol, carotene, and n-nonyl aldehyde [19,105]
99 Ferula bungeana Kitag. Whole plant and seeds Heat-clearing and detoxifying, relieving pain and expelling phlegm, and arresting coughing Treatment of cold, bronchopneumonia, and pulmonary tuberculosis Anisole, d-fenchone, and limonen [19,106]
100 Ferula caspica M. Bieb. Roots and resin Eliminating stagnated food, relieving dyspepsia, and insecticide Toxicity effect Umbelliprenin, farnesyl alcohol, and umbelliferone [107]
101 Ferula conocaula Korovin Resin, roots, and rhizome Eliminating stagnated food, insecticide, treatment of abdominal mass, and a lump in the abdomen Anticancer and treatment of influenza Umbelliprenin, fezelol, and feterin [107]
102 Ferula feruloides (Steud.) Korovin Roots and resin Treatment of chilliness, and pain of the heart and abdomen Insecticidal, bacteriostat and antitumor α-pinene, farnesene and toluene [108,109]
103 **Ferula fukanensis K. M. Shen Resin Eliminating stagnated food, relieving dyspepsia and insecticide Treatment of stomach disease, rheumatism and joint pain Ferulic acid, guaiol and ethyl-p-hydroxybenzoate [18,19,110,111,112]
104 Ferula jaeschkeana Vatke Resin of overground part Eliminating stagnated food, insecticide, treatment of tumour, wound, and peptic ulcer Antifertility Jaeschkeanadiol, α-pinene and β-pinene [107]
105 Ferula krylovii Korovin Resin Eliminating stagnated food and insecticide \ fekrynol, ferukrin and fekrynol acetate [107]
106 Ferula lehmannii Boiss. Resin Detoxification, deodorize, and insecticide Treatment of gastropathy, rheumatism and arthralgia Lehmannolone, sinkianone, and lehmannolone A [16,113]
107 Ferula moschata (Reinsch) Koso-Pol. Roots Sedative, spasmolysis, and treatment of hysteria Suppress the replication of human immunodeficiency virus in H9 lymphocytes and suppress the production of cytokine fezelol, fesumtuorin A and fesumtuorin B [107]
108 Ferula olivacea (Diels) H. Wolff ex Hand.-Mazz. Resin Wind-heat dispersing, expelling phlegm, and arresting coughing \ \ [16]
109 **Ferula sinkiangensis K. M. Shen Resin Eliminating stagnated food, detoxification, and insecticide Antioxidant, antitumor, and antiviral Ferulic acid, fekrynol, and lehmannolone [16,18,114,115]
110 Ferula songarica Pall. Ex Schult. Resin and whole plant Eliminating stagnated food and insecticide \ 2,4-dihydroxylacetophenone, 3,3′, 4,4′-biphenyltetracarboxylic acid, and Δ3-carene [116]
111 Ferula teterrima Kar. & Kir. Resin Eliminating stagnated food, and insecticide Treatment of malaria and dysentery Feterin, badrakemin, and badrakemin acetate [116]
112 *Foeniculum vulgare Mill. Fruits, roots, stems, leaves, and whole plant Dispelling wind, relieving pain, and harmonize the stomach Bacteriostat, anti-inflammatory, and antianxiety Trans-anethole, estragole, and anisaldehyde [18,19,117]
113 **Glehnia littoralis F. Schmidt ex Miq. Roots Heat-clearing and detoxifying, diminish inflammation, and expelling phlegm and arresting coughing Anti-inflammatory, bacteriostat, and antitumor Phenyllactic acid, catechol, and quercetin [18,118]
114 Hansenia oviformis (R. H. Shan) Pimenov & Kljuykov Rhizome, roots, and leaves Treatment of rheumatic arthralgia, cold due to wind-cold, and headache \ \ [16,102]
115 Heracleum barmanicum Kurz Roots Treatment of cold abdominalgia \ \ [16]
116 Heracleum candicans Wall. Ex DC. Roots Dispelling wind, eliminating dampness, and relieving pain Treatment of cold headache Bergapten, heraclenin, and imperatorin [19,119]
117 Heracleum dissectifolium K. T. Fu Roots Dispelling wind, eliminating dampness, and relieving pain \ \ [16]
118 Heracleum fargesii H. Boissieu Roots \ \ \ [17]
119 Heracleum franchetii M. Hiroe Roots and rhizome \ \ \ [120,121]
120 Heracleum hemsleyanum Roots and rhizome Dispelling wind, eliminating dampness, and relieving pain Antioxidant, anti-inflammatory, and antitumor β-pinene, α-pinene, and (1S)-6,6-dimethyl-2-methylene-bicyclo[3.1.1] heptane [26,27,122,123]
121 Heracleum hemsleyanum Diels Roots and rhizome Dispelling wind, eliminating dampness, and relieving pain Antioxidant, anti-inflammatory, and antitumor Osthole, columbianadin, and columbianetin [26,27]
122 Heracleum henryi H. Wolff Roots Clearing and activating the channels and collaterals, relieving pain, and scattered stasis \ Turgeniifolin B, turgeniifolin C, and bergapten [124]
123 Heracleum millefolium var. Millefolium Roots and rhizome Detumescence, disintegrate masse, and treatment of leprosy \ \ [102,120,121]
124 Heracleum moellendorffii Hance Roots and rhizome Clearing and activating the channels and collaterals, relieving pain, and scattered stasis Bacteriostat β-pinene, α-pinene, and pentadecane [122,124,125,126]
125 Heracleum oreocharis H. Wolff Roots \ \ \ [121]
126 Heracleum rapula Franch. Roots Clearing and activating the channels and collaterals, relieving pain, and scattered stasis Bacteriostat, treatment of asthma, and chronic bronchitis Ostholce, marmesin, and imperatorin [19,124,127]
127 Heracleum scabridum Franch. Roots, rhizome, and fruits Treatment of common cold due to wind-cold, headache, and cough asthma \ Heraclenol, oxypeucedanin-hydrate, and byakangelicin [128,129,130]
128 Heracleum souliei H. Boissieu Roots \ \ Bergapten [119,121]
129 Heracleum stenopterum Diels Roots Treatment of cold and rheumatism \ Bergapten, isopimpinellin, and sphondin [16,131]
130 Heracleum tiliifolium H. Wolff Roots Dispelling wind, eliminating dampness, and relieving pain \ \ [16]
131 Heracleum vicinum H. Boissieu Roots Same as Notopterygium incisum \ \ [120,121]
132 Heracleum wenchuanense F. T. Pu & X. J. He Roots \ \ \ [121]
133 Heracleum wolongense F. T. Pu & X. J. He Roots \ \ \ [1,121]
134 Heracleum yungningense Hand.-Mazz. Roots and rhizome Treatment of waist and knee pain, limb spasm, and leucoderma \ Pimpinellin, angelicin, and isobergapten [26,132]
135 Hydrocotyle himalaica P. K. Mukh. Whole plant Heat-clearing, detoxifying, and eliminating dampness \ Asiaticoside, madecassoside, and quercetin [133,134]
136 Hydrocotyle hookeri subsp. Chinensis (Dunn ex R. H. Shan & S. L. Liou) M. F. Watson & M. L. Sheh Whole plant Relieving pain, diuresis, and removing dampness Antiviral, antitumor, and anti-bacterial Flavonoids, triterpenes, and volatile oils [16,128,134]
137 Hydrocotyle nepalensis Hook. Whole plant Clearing heat and promoting diuresis, dissolving stasis, and hemostasis and detoxicate Antiviral, antitumor, and anti-bacterial Flavonoids, triterpenes, and volatile oils [16,134]
138 Hydrocotyle sibthorpioides Lam. Whole plant Heat-clearing, diuresis, and detumescence Anti-ulcer, antilipemic, and antiviral Quercetin, isorhamnetin, and asiaticoside [134,135]
139 Hydrocotyle sibthorpioides var. batrachium (Hance) Hand.-Mazz. Ex R. H. Shan Whole plant Heat-clearing and detoxifying, eliminating dampness, and diuresis Anti-ulcer, spasmolysis, and anti-inflammatory Benzene propane nitrile, phytol, and caryophyllene oxide [16,136,137]
140 Hydrocotyle wilfordii Maxim. Whole plant As Hydrocotyle nepalensis Hook. As Hydrocotyle nepalensis Hook. Asiaticoside, madecassoside, and quercetin [133,134]
141 Hymenidium chloroleucum (Diels) Pimenov & Kljuykov Roots or whole plant Regulating flow of qi, invigorating stomach, and activating blood Anti-inflammatory, analgesia, and nutritious function Nobiletin, falcarindiol, and isoliquiritingenin [19,138,139]
142 Hymenidium davidii (Franch.) Pimenov & Kljuykov Roots \ \ \ [140]
143 Hymenidium delavayi (Franch.) Pimenov & Kljuykov Roots \ \ \ [1,6]
144 Hymenidium lindleyanum (Klotzsch) Pimenov & Kljuykov Roots Treatment of hypertensive, coronary heart disease, and altitude stress \ Bergapten, isoimperatorin, and oxypeucedanin [141]
145 Kitagawia formosana (Hayata) Pimenov Roots \ \ \ [1]
146 Kitagawia macilenta (Franch.) Pimenov Roots Expelling phlegm \ \ [142]
147 Kitagawia terebinthacea (Fisch. Ex Trevir.) Pimenov Roots Cleaning heat and dispelling wind, alm the adverse-rising energy, and expelling phlegm Treatment of cold and cough, bronchitis, and cough during pregnancy Isoepoxybuterixin [19]
148 Levisticum officinale W. D. J. Koch Roots Diuresis, invigorate the stomach, and expelling phlegm Inhibition of rhythmic uterine contractions, Scavenging oxygen free radicals, and anti- lipid peroxidation Ligustilide, α-phellandrene, and β-phellandrene [19,143]
149 Libanotis buchtormensis (Fisch.) DC. Roots Divergent wind chill, dispel wind-damp, and relieving pain Bacteriostat, treatment of common cold due to wind-cold, generalized pain, and cough Falcarinone, isoimperatorin, and xanthotoxin [19,144]
150 Libanotis iliensis (Lipsky) Korovin Roots Expel wind-cold pathogens, thermolysis, and relieving pain Treatment of common cold due to wind-cold and rheumatic arthritis Archangelin and iliensin [19]
151 Libanotis lancifolia K. T. Fu Roots Divergent wind chill, dispel wind-damp, and relieving pain Bacteriostat, treatment of common cold due to wind-cold, generalized pain, and cough Falcarinone, isoimperatorin, and xanthotoxin [19,144]
152 Libanotis laticalycina R. H. Shan & M. L. Sheh Roots Dispelling wind, antispasmodic, and relieving pain Analgesia, sedation, and anti-inflammatory Octanal, hexanal, and 2-pentylfuran [16,145,146]
153 Libanotis seseloides (Fisch. & C. A. Mey. Ex Turcz.) Turcz. Roots Eliminating dampness, activating spleen, and promote blood circulation Treatment of damobstruction, dysentery, and sore Edultin [19]
154 Libanotis sibirica (L.) C. A. Mey. Roots \ \ \ [1]
155 Libanotis spodotrichoma K. T. Fu Roots Divergent wind chill, dispel wind-damp, and relieving pain Bacteriostat, treatment of common cold due to wind-cold, generalized pain, and cough Falcarinone, isoimperatorin, and xanthotoxin [19,144]
156 Ligusticopsis brachyloba (Franch.) Leute Roots Sudation, relieving pain, and dispelling wind Treatment of headache dizziness, arthralgia, and tetanus α-pinene, β-pinene, and sabinene [147,148,149]
157 Ligusticopsis daucoides (Franch.) Lavrova & Kljuykov Roots \ \ \ [1,94]
158 Ligusticopsis likiangensis (H. Wolff) Lavrova & Kljuykov Roots \ \ \ [1,94]
159 **Ligusticum chuanxiong Hort. Roots, rhizome, stems, and leaves Activating blood, relieving pain, and Dispelling wind Anti-inflammatory, antioxidant, and antitumor Abietene, tetramethylpyrazine, and glucose [18,19,150]
160 **Ligusticum jeholense Nakai et Kitag. Roots and rhizome Dispelling wind, dispersing cold, and eliminating dampness Anti-inflammatory, sedation, and anti-ulcer Ferulic acid, isoferulic acid, and daucosterol [18,19,151,152]
161 Ligusticum pteridophyllum Franch. Roots Dispelling wind, relieving pain, and eliminating dampness Treatment of cold due to wind-cold and migraine Asaricin, β-sitosterol, and daucosterol [26,153]
162 **Ligusticum sinense Oliv. Roots, rhizome, and tuber Expel wind-cold pathogens, eliminating dampness, and relieving pain Anti-inflammatory, central inhibitory, and anti-thrombotic effect 3-butylphthalide, opthalonide, and neopthalonide [18,154]
163 Ligusticum tenuissimum (Nakai) Kitagawa Roots and rhizome Same as ligusticum sinense Oliv. Divergent wind chill, treatment of wind-cold headache, and diarrhoea. Analgesia and sedation Ferulic acid [19,94,155]
164 Meeboldia delavayi (Franch.) W. Gou & X. J. He Roots Treatment of cold, fever, headache \ \ [16]
165 Nothosmyrnium japonicum var. Japonicum Roots \ Sedation and analgesia \ [16]
166 Nothosmyrnium japonicum var. Sutchuensis H. Boissieu Roots \ Sedation and analgesia \ [16]
167 **Notopterygium franchetii H. De Boiss. Roots and rhizome Divergent wind chill, dispelling wind, and eliminating dampness Anti-inflammatory, analgesia, and antiviral Nodakenin, ferulic acid, and bergamot lactone [18,156,157]
168 **Notopterygium incisum Ting ex H. T. Chang Roots and rhizome Divergent wind chill, dispelling wind, and eliminating dampness Anti-inflammatory, analgesia, and antiviral Nodakenin, notopterol, and isoimperatorin [18,157]
169 Oenanthe benghalensis Benth. & Hook. Roots and whole plant Same as Oenanthe javanica (Blume) DC. Same as Oenanthe javanica (Blume) DC. \ [17,158]
170 Oenanthe javanica (Blume) DC. Roots, stems and whole plant Heat-clearing, detoxification, and removing liver-fire Enhancing immunity, antiarrhythmic, and hypoglycemic Phytic acid, γ-terpinene, and caryophyllene [19,159]
171 Oenanthe linearis subsp. Rivularis (Dunn) C. Y. Wu & F. T. Pu Roots and whole plant Same as Oenanthe javanica (Blume) DC. Same as Oenanthe javanica (Blume) DC. \ [17]
172 Osmorhiza aristata var. Laxa (Royle) Constance & R. H. Shan Roots Divergent wind chill, sudation, and relieving pain \ \ [16]
173 Ostericum citriodorum (Hance) C. C. Yuan & R. H. Shan Roots and whole plant Activating blood, dissolving stasis, and dispelling wind Expectorant, anti-inflammatory, and bacteriostat Isoapiole, panaxynol, and myristicin [19,160,161,162]
174 Ostericum grosseserratum (Maxim.) Kitag. Roots Activating spleen, dispersing cold, and invigorating spleen and replenishing qi \ Octanal, β-pinene, and myristic acid [16,163,164]
175 Ostericum sieboldii (Miq.) Nakai Roots \ \ \ [165,166,167]
176 Peucedanum dielsianum Fedde ex H. Wolff Roots and rhizome Relieving pain, dispelling wind, and eliminating dampness \ Isoimperatorin, Phellopterin, and 9-octadecenoic acid [19,168,169]
177 Peucedanum dissolutum (Diels) H. Wolff Roots \ \ \ [1]
178 Peucedanum harry-smithii var. Subglabrum Roots Same as Peucedanum praeruptorum, alleviate asthma, reducing phlegm, and heatelimination Treatment of bronchitis, hypertensive, and coronary heart disease Psoralen, bergapten, and xanthotoxin [170,171,172,173]
179 Peucedanum japonicum Thunb. Roots Clearing heat, relieving cough, and diuresis Antipyresis, analgesia, and anti-inflammatory Peucedanol, umbelliferone, and β-pinene [19,174,175]
180 Peucedanum ledebourielloides K. T. Fu Roots \ \ \ [1,167]
181 Peucedanum longshengense R. H. Shan & M. L. Sheh Roots \ \ \ [1]
182 Peucedanum mashanense R. H. Shan & M. L. Sheh Roots Expelling phlegm \ \ [142]
183 Peucedanum medicum Dunn Roots Expelling phlegm, alleviating asthma and cough, and arresting convulsion Anticoagulation, antioxidant, and antibacterial 2-methoxy-4-vinylphenol, p-menthan-1-ol, and cis-α-bisabolene [19,176,177]
184 Peucedanum medicum var. Gracile Dunn ex R. H. Shan & M. L. Sheh Roots and rhizome Expelling phlegm, alleviating asthma and cough, and arresting convulsion Anticoagulation, antioxidant, and antibacterial Isoimperatorin, phellorerin, and bergapten [19,176,178]
185 Peucedanum medicum var. Medicum Roots and rhizome Expelling phlegm, alleviating asthma and cough, and arresting convulsion Anticoagulation, antioxidant, and antibacterial 2-methoxy-4-vinylphenol, p-menthan-1-ol, and cis-α-bisabolene [19,176,177]
186 **Peucedanum praeruptorum Dunn Roots Divergent wind, clearing heat, and reducing phlegm Anticoagulation, antioxidant, and anticancer Praeruptorin A, praeruptorin B, and scopoletin [18,179]
187 Peucedanum shanianum F. L. Chen & Y. F. Deng Roots Relieving asthma, expelling phlegm, and spasmolysis Anti-inflammatory, antiallergic, and anti-ulcer Sinodielides A, deltoin, and (+)-pareruptorin A [180,181,182,183]
188 Peucedanum turgeniifolium H. Wolff / Peucedanum pulchrum Roots and whole plant Expelling phlegm, antibechic, and dispersing wind-heat Smooth muscle spasmolysis Turgenifolin A, turgenifolin B, and bergapten [19,183,184]
189 Peucedanum wawrae (H. Wolff) S. W. Su ex M. L. Sheh Roots Antibechic and expelling phlegm Analgesia, sedation, and anti-inflammatory Peucedanocoumarin, d-laserpitin, and bergapten [16,167,185]
190 Peucedanum wulongense R. H. Shan & M. L. Sheh Roots \ \ \ [1]
191 Phlojodicarpus sibiricus (Steph. Ex Spreng.) Koso-Pol. Roots \ \ \ [1]
192 Physospermopsis alepidioides (H. Wolff & Hand.-Mazz.) R. H. Shan Roots \ \ \ [1]
193 Physospermopsis delavayi (Franch.) H. Wolff Roots \ \ \ [1]
194 Pimpinella anisum L. Fruits Warming meridian and diuresis Treatment of paralysis, facial paralysis, and migraine Anisaldehyde, anisole, and (E)-anethole [186,187,188,189,190]
195 Pimpinella candolleana Wight & Arn. Roots or whole plant Warming spleen and stomach for dispelling cold, relieving pain, and dispelling wind Relieving muscular spasm, antiviral, and antibacterial α-zingiberene, pregeijerene, and β-elemene [19,191,192,193]
196 Pimpinella coriacea (Franch.) H. Boissieu Whole plant Warming spleen and stomach for dispelling cold, dispelling wind and eliminating dampness, and activating blood \ \ [194]
197 Pimpinella diversifolia DC. Whole plant Expelling phlegm, activating blood, relieving pain, and removing toxicity for detumescence Anti-inflammatory, antitumor, and antituberculous 1H-benzocycloheptene, sesquiphellandrene, and β-chamigrene [195,196,197]
198 Pimpinella diversifolia var. Diversifolia Roots or whole plant Invigorating stomach, dispersing accumulations, and antidiarrheic Anti-inflammatory, antitumor, and antituberculous 1H-benzocycloheptene, sesquiphellandrene, and β-chamigrene [19,195,196,197]
199 Pimpinella thellungiana H. Wolff Roots or whole plant Warming spleen and stomach for dispelling cold, benefiting qi and nourishing blood, and coordinating yin and yang Hypotensive, hypolipidemic, and modulates, and improves cellular immunity Protocatechuic acid, gallic acid and neochlorogenic acid [198,199,200,201,202]
200 Pleurospermopsis bicolor (Franch.) Jing Zhou & J. Wei Whole plant Warming spleen and stomach for dispelling cold, benefiting qi and nourishing blood, and coordinating yin and yang Hypotensive, antilipemic, and modulates and improves cellular immun antimicrobial ity Chlorogenic acid, isochlorogenic acid A, and apigenin-7-O-β-D-glucuronopyranoside [198,200,201]
201 Pleurospermum aromaticum W. W. Sm. Whole plant \ \ \ [1]
202 Pleurospermum giraldii Diels Whole plant and seeds Warming spleen, digesting food, and checking vaginal discharge Inhibition of smooth muscle contraction and release intestinal smooth muscle spasm Carvone, n-triactanol, and γ-sitosterol [19,203,204,205]
203 Pleurospermum rivulorum (Diels) K. T. Fu & Y. C. Ho Roots or whole plant Tonifying the kidney \ \ [1,102]
204 Pternopetalum leptophyllum (Dunn) Hand.-Mazz. Whole plant \ \ \ [16]
205 Pternopetalum vulgare var. Vulgare Roots or whole plant Treatment of lumbago \ \ [19]
206 Sanicula astrantiifolia H. Wolff ex Kretschmer Whole plant Tonifying the kidney and lung, treating tuberculosis, and kidney vacuity lumbar pain Antioxidant, antibacterial, and bacteriostat Total flavonoids, rutin, and polysaccharides [206,207,208]
207 Sanicula caerulescens Franch. Whole plant Dispelling wind, melting phlegm, and promoting blood circulation for regulating menstruation Expectorant, antibechic, and anti-inflammatory Angelicin, isoferulaldehyde, and 12-hydroxybakuchiol [19,209,210]
208 Sanicula chinensis Bunge Whole plant Detoxification, hemostasis, and treatment of throat pain Antiviral \ [128,211,212,213]
209 Sanicula elata Buch.-Ham. Ex D. Don Whole plant Same as Sanicula lamelligera Antiviral Oleanane saponins, saponins, and microelement [211,212,213,214,215,216]
210 Sanicula lamelligera Hance Whole plant Dispelling wind, melting phlegm, and promoting blood circulation for regulating menstruation Expectorant, antibechic, and anti-inflammatory Angelicin, isoferulaldehyde, and 12-hydroxybakuchiol [19,209,210]
211 Sanicula orthacantha S. Moore Roots or whole plant Heat-clearing and detoxifying, treatment of traumatic injury \ \ [16]
212 Sanicula orthacantha var. Brevispina H. Boissieu Whole plant Heat-clearing and detoxifying, treatment of traumatic injury \ \ [16]
213 **Saposhnikovia divaricata (Turcz.) Schischk. Roots Dispelling wind to relieve superficies, removing dampness to relieve pain, and arrest convulsio Analgesia, sedation, and anti-inflammatory Prim-o-glucosylcimifugin, 5-O-methylvisamitol glycoside, and cimifugin [18,217,218]
214 Selinum cryptotaenium H. Boissieu Roots \ \ \ [1]
215 Semenovia montana Kamelin & V. M. Vinogr. Roots \ \ \ [121]
216 Seseli delavayi Franch. Roots Dispelling wind, removing dampness, and relieving pain \ \ [19]
217 Seseli mairei var. Mairei Roots and rhizome Dispelling wind, removing dampness, and relieving pain Antipyretic, analgesia, and anti-inflammatory Sphondin, bergapten and isopimpinellin [19,219,220,221]
218 Seseli yunnanense Franch. Roots and rhizome Dispelling wind, removing dampness, and relieving pain Antipyretic, analgesia, and anti-inflammatory Falcarindiol, falcarinol, and glycerol monolinoleate [19,219,220,222]
219 Seselopsis tianschanica Schischk. Roots Treatment of fall injury, anemia, and other diseases Treatment of nasopharynx cancer \ [16]
220 Sium suave Walter Whole plant Dispersing cold, relieving headache, and decreasing blood pressure \ \ [16,223]
221 Spuriopimpinella arguta (Diels) X. J. He & Z. X. Wang Roots and whole plant \ \ \ [194]
222 Tongoloa silaifolia (H. Boissieu) H. Wolff Roots Stopping bleeding, relieving pain, and activating blood Treatment of traumatic injury, trauma bleeding, and rheumatic pain Suberosin, crenulatin, and isoimperatorin [19,224,225]
223 Tongoloa stewardii H. Wolff Roots \ \ \ [1]
224 Torilis japonica (Houtt.) DC. Fruits and roots Lumbricide ascaricide, and external antiphlogistic agent \ Essential oil [19]
225 Torilis scabra (Thunb.) DC. Fruits or whole plant Activating blood, insecticide, and antidiarrheal Bacteriostat Cyclohexene, 6,6-dimethyl-bicyclo [3.1.1] heptane-2-carboxaldehyde, and endo-borneol [19,194,226]
226 Trachyspermum ammi (L.) Sprague. Fruits Dispersing cold, relieving pain, and treatment of indigestion Antibacterial, antimicrobial, and antifungal thymol, ρ-cymene, and β-pinene [19,187,227] [228,229,230]
227 Vicatia thibetica H. Boissieu Roots Dispelling wind, eliminating dampness, and dispelling cold Anti-fatigue, antioxidant, and enhancing immunity Umbelliferone, bergapten, and ferulic acid [231,232,233]
228 Visnaga daucoides Gaertn. Fruits Treatment of coronary artery disease, such as panhandling coronary thrombosis Treatment of renal colic, angina pectoris, and urinary calculi Khellin, visnagin, and khellol glycoside [16,234]
Note: “*” means the plant reported in Pharmacopoeia of the People’s Republic of China (2020), “**” means the plant roots used as medicine reported in Pharmacopoeia of the People’s Republic of China (2020), the same below.
Table 2. Quality markers in the 22 AMPs recorded in the Pharmacopoeia of the People’s Republic of China (2020).
Table 2. Quality markers in the 22 AMPs recorded in the Pharmacopoeia of the People’s Republic of China (2020).
No. /No. in Table 1 Plant species Quality markers Classification Biosynthetic pathway
1/8 Angelica biserrata Osthole (1) and columbianadin (2) Coumarins Phenylpropanoids
2/10 Angelica dahurica Imperatorin (3) and isoimperatorin (4) Coumarins Phenylpropanoids
3/11 Angelica dahurica cv. Hangbaizhi (3) and (4) Coumarins Phenylpropanoids
4/13 Angelica decursiva Nodakenin (5) Coumarins Phenylpropanoids
5/20 Angelica sinensis Ferulic acid (6) and ligustilide (15) Propenyl benzenes and phthalides Phenylpropanoids and phthalides
6/34 Bupleurum chinense Saikosaponin a (11) and saikosaponin d (12) Triterpenes Terpenes
7/67 Bupleurum scorzonerifolium (11) and (12) Triterpenes Terpenes
8/79 Changium asiatica Asiaticoside (13) and madecassoside (14) Triterpenes Terpenes
9/80 Changium smyrnioides – – –
10/83 Changium monnieri (1) Coumarins Phenylpropanoids
11/94 Daucus carota – – –
12/102 Ferula fukanensis – – –
13/109 Ferula sinkiangensis – – –
14/112 Foeniculum vulgare Trans-anethole (7) Phenylpropene Phenylpropanoids
15/113 Glehnia littoralis – – –
16/159 Ligusticum chuanxiong (6) and levistilide A (16) Phenylpropanoids and phthalide Phenylpropanoids and phthalides
17/160 Ligusticum jeholense (6) Phenylpropanoids Phenylpropanoids
18/162 Ligusticum sinense (6) Phenylpropanoids Phenylpropanoids
19/167 Notopterygium franchetii (4), (5), and notopterol (8) Coumarins Phenylpropanoids
20/168 Notopterygium incisum (4), (5), and (8) Coumarins Phenylpropanoids
21/186 Peucedanum praeruptorum Praeruptorin A (9) and praeruptorin B (10) Coumarins Phenylpropanoids
22/213 Saposhnikovia divaricata Prim-O-glucosylcimifugin (17) and 5-O-methylvisammioside (18) Chromones Chromones
Note: The “–” indicates there are no specific quality markers recorded in the Pharmacopoeia of the People’s Republic of China (2020).
Table 3. Classification of the 38 rhizomatousAMPs affected by the BF.
Table 3. Classification of the 38 rhizomatousAMPs affected by the BF.
No. /No. in Table 1 Plant species Classes References No. /No. in Table 1 Plant species Classes References
1/4 Angelica acutiloba (Siebold & Zucc.) Kitag. (1) [301] 20/109 *Ferula sinkiangensis K. M. Shen (3) [19]
2/8 **Angelica biserrata (R. H. Shan & C. C. Yuan) C. C. Yuan & R. H. Shan (1) [302] 21/111 Ferula teterrima Kar. & Kir. (3) [19]
3/10 **Angelica dahurica (Fisch. ex Hoffm.) Benth. & Hook. f. ex Franch. & Sav. (1) [303] 22/113 **Glehnia littoralis F. Schmidt ex Miq. (2) [304]
4/11 **Angelica dahurica cv. Hangbaizhi (1) [303] 23/121 Heracleum hemsleyanum Diels (1) [302]
5/13 **Angelica decursiva (Miq.) Franch. & Sav. (1) [305] 24/126 Heracleum rapula Franch. (1) [19]
6/14 Angelica gigas Nakai (2) [306] 25/148 Levisticum officinale W. D. J. Koch (3) [19]
7/19 Angelica polymorpha Maxim. (1) [19] 26/149 Libanotis buchtormensis (Fisch.) DC (3) [307]
8/20 **Angelica sinensis (Oliv.) Diels (1) [308] 27/150 Libanotis iliensis (Lipsky) Korovin (1) [19]
9/26 Anthriscus sylvestris (L.) Hoffm. (3) [309] 28/151 Libanotis lancifolia K. T. Fu (3) [19,307]
10/34 **Bupleurum chinense DC. (2) [310] 29/153 Libanotis seseloides (Fisch. & C. A. Mey. ex Turcz.) Turcz. (1) [19]
11/67 **Bupleurum scorzonerifolium Willd. (2) [310] 30/155 Libanotis spodotrichoma K. T. Fu (3) [19,307]
12/80 **Changium smyrnioides H. Wolff (2) [311] 31/159 **Ligusticum chuanxiong Hort. (2) [312]
13/81 Chuanminshen violaceum M. L. Sheh & R. H. Shan (2) [313] 32/160 **Ligusticum jeholense Nakai et Kitag. (2) [314]
14/82 Cicuta virosa L. (3) [19] 33/162 **Ligusticum sinense Oliv. (2) [314]
15/96 Daucus carota var. sativus Hoffm. (1) [315] 34/167 **Notopterygium franchetii H. de Boiss. (2) [316]
16/102 Ferula feruloides (Steud.) Korovin (3) [19] 35/168 **Notopterygium incisum Ting ex H. T. Chang (2) [316]
17/103 Ferula fukanensis K. M. Shen (3) [19] 36/186 **Peucedanum praeruptorum Dunn (1) [317]
18/106 Ferula lehmannii Boiss. (3) [19] 37/195 Pimpinella candolleana Wight & Arn. (3) [19]
19/108 Ferula olivacea (Diels) H. Wolff ex Hand.-Mazz. (3) [19] 38/213 **Saposhnikovia divaricata (Turcz.) Schischk. (1) [318,319]
Note: (1) the BF significantly affects the yield and quality, and the rhizomes or roots are absolutely useless for clinical effects; (2) the BF differently affects the yield, while the rhizomes or roots can be used as medicine to some extent; and (3) the BF has no significant effect on the yield and quality, and their rhizomes or roots are used as medicine without doubtedly, the same below.
Table 4. Approach to inhibit BF of 25 AMPs have been reported.
Table 4. Approach to inhibit BF of 25 AMPs have been reported.
Classes No. /No. in Table 1 Plant species Measure Ⅰ
(Seeding)		 Measure Ⅱ
(Cultivation)		 Measure Ⅲ
(Abiotic)		 Measure Ⅳ
(Molecular biology)
(1) 1/4 Angelica acutiloba (Siebold & Zucc.) Kitag. Seedling diameter [334] Density of planted seedlings [334] Paclobutrazol concentration [334] \
(1) 2/8 **Angelica biserrate (R. H. Shan & C. C. Yuan) C. C. Yuan & R. H. Shan Seedling size and root length [302] \ \ \
(1) 3/10 **Angelica dahurica (Fisch. ex Hoffm.) Benth. & Hook. f. ex Franch. & Sav. Seed quality and seed maturity degree [303,328] Soil selection should avoid continuous cropping and fertile sticky soil, density of planted seedlings, and seeding time [328,330,335] Rational application of fertilizer, and appropriate N, P, and K fertilizer [303,328,336] Seven types of reproductive conversion genes, and adconstans-like gene [337,338]
(1) 4/11 **Angelica dahurica cv. Hangbaizhi Seed quality and seed maturity degree [303,328] Soil selection should avoid continuous cropping and fertile sticky soil, density of planted seedlings, and seeding time [328,330,335] Rational application of fertilizer, and appropriate N, P, and K fertilizer [303,328,336] Seven types of reproductive conversion gene, and adconstans-like gene [337,338]
(1) 5/13 **Angelica decursiva (Miq.) Franch. & Sav. \ \ \ \
(1) 6/19 Angelica polymorpha Maxim. \ \ \ \
(1) 7/20 **Angelica sinensis (Oliv.) Diels Seed maturity degree, seeding age, seeding weight, root diameter, and excellent variety [323,325,339,340,341] Short-day, storage temperature, and reasonable planting and cultivation [308,339,342] Plant growth retardant [343] Four pathways of genes for regulating early bolting and flowering [344,345]
(1) 8/96 Daucus carota var. Sativus Hoffm. Endogenous hormone content and different cultivars [346,347] Temperature, short-day, and seeding time [346,348,349,350] \ Two major genes: Bol1-1 and Bol1-2 [351]
(1) 9/121 Heracleum hemsleyanum Diels \ \ \ \
(1) 10/126 Heracleum rapula Franch. \ \ \ \
(1) 11/150 Libanotis iliensis (Lipsky) Korovin \ \ \ \
(1) 12/153 Libanotis seseloides (Fisch. & C. A. Mey. ex Turcz.) Turcz. \ \ \ \
(1) 13/186 **Peucedanum praeruptorum Dunn \ Compact planting and seeding time [352,353] \ \
(1) 14/213 **Saposhnikovia divaricata (Turcz.) Schischk. \ Density of planted seedlings [333] \ Differentially expressed genes associated with bolting and flowering during early flowering, flower bud differentiation, and late flowering [354]
(2) 15/14 Angelica gigas Nakai \ \ \ \
(2) 16/34 **Bupleurum chinense DC. \ Cut the flowers [310] Temperature [355] Flowering gene (bcsvp, bcpaf1, bcco and bcft) [356]
(2) 17/67 **Bupleurum scorzonerifolium Willd. \ \ \ \
(2) 18/80 Changium smyrnioides H. Wolff \ Cut the flowers [311] \ \
(2) 19/81 Chuanminshen violaceum M. L. Sheh & R. H. Shan \ \ \ \
(2) 20/113 **Glehnia littoralis F. Schmidt ex Miq. \ Cut the flowers [304] \ \
(2) 21/159 **Ligusticum chuanxiong Hort. Asexual reproduction and tissue cultur [312,357] Cut the bolted stem [358] \ Transcriptome original data by Illumina sequencing technology [359]
(2) 22/160 **Ligusticum jeholense Nakai et Kitag. \ Cut the flower [360,361] \ \
(2) 23/162 **Ligusticum sinense Oliv. \ Cut the flower [360,361] \ \
(2) 24/167 **Notopterygium franchetii H. de Boiss. \ Cut the flower [362] \ \
(2) 25/168 **Notopterygium incisum Ting ex H. T. Chang \ Cut the flower [316] \ \
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