Pseudostellaria Heterophylla Research Articles

Pseudosterins A-C, Three 1-Ethyl-3-formyl-β-carbolines from Pseudostellaria heterophylla and Their Cardioprotective Effects.

Pseudostellaria heterophylla is used in China not only as a functional food but also as an herb to tonify the spleen, enhance immunity, and treat palpitation. Our previous investigation showed that a fraction enriched in glycosides obtained from the roots of P. heterophylla possessed pronounced protective effects on H9c2 cells against CoCl2-induced hypoxic injury. However, the active compounds responsible for the observed effects were still unknown. In the current investigation, pseudosterins A–C (1–3), three new alkaloids with a 1-ethyl-3-formyl-β-carboline skeleton, together with polydatin, have been isolated from the active fraction. Their structures were elucidated on the basis of spectroscopic analysis and quantum chemical calculations. The four compounds showed cardioprotective effects against sodium hydrosulfite-induced hypoxia-reoxygenation injury in H9c2 cells, with the three alkaloids being more potent. This is also the first report of alkaloids with a β-carboline skeleton isolated from P. heterophylla as cardioprotective agents.

Heterophyllin B, a cyclopeptide from Pseudostellaria heterophylla, enhances cognitive function via neurite outgrowth and synaptic plasticity

Neurite outgrowth-induced construction of neural circuits and networks is responsible for memory generalization, consolidation, and retrieval. In this study, we found that the traditional Chinese medicine Pseudostellaria heterophylla promoted neurite regrowth and enhanced cognitive function in normal mice. Further, we orally administered Pseudostellaria heterophylla water extracts (PHE) to ICR mice, and detected heterophyllin B (HET-B), an important cyclopeptide, in the plasma and cerebral cortex. We demonstrated that neurites were significantly elongated after coculturing with HET-B for 4 days. Next, the intraperitoneal injection of HET-B on seven consecutive days in 3-month-old ICR mice significantly enhanced the object recognition memory and object location memory than that in control. Immunohistochemical analysis indicated significantly increased β3-tubulin-positive neurite density, synaptophysin, and postsynaptic density 95 in the perirhinal cortex and hippocampus after administering HET-B. Furthermore, the concentration of neurotransmitters was measured using HPLC analysis; HET-B significantly increased five-levels of HT in the hippocampus, and decreased metabolites of dopamine, dihydroxyphenylacetic acid, and homovanillic acid, in the prefrontal cortex and hippocampus. Taken together, HET-B induces neurite elongation and neurotransmitter regulation and possibly enhances cognitive memory.

Effects of combined application of boron-molybdenum-copper microfertilizers on photosynthetic characteristics of Pseudostellaria heterophylla

Effects of combined application of boron-molybdenum-copper microfertilizers on photosynthetic characteristics of Pseudostellaria heterophylla

Stimulatory Effect of Fe3O4 Nanoparticles on the Growth and Yield of Pseudostellaria heterophylla via Improved Photosynthetic Performance

Nanomaterials have recently been used as growth stimulants to promote the production of crops in saline-alkali through root application. However, if applied through leaves, little is known about the effect of Fe3O4 nanoparticles (NPs) on the root growth and yield, especially for medicinal crops. To fill this gap, a single factor experiment was conducted to explore the effects of Fe3O4 NPs on growth, yield, the dry matter distribution, chlorophyll content, photosynthetic characteristics, chlorophyll fluorescence parameters, and polysaccharide content of Pseudostellaria heterophylla by foliar spraying under field conditions. Fe3O4 NPs (20–50 mg·L–1) significantly promoted growth, the dry matter distribution of root and root tuber yield per unit area. Fe3O4 NPs enhanced net photosynthetic rate (Pn) by increasing chlorophyll content. And Fe3O4 NPs increased the daily mean and peak value of Pn, and alleviated the phenomenon of “midday depression” by improving nonstomatal limitation. Chlorophyll fluorescence parameters indicating that Fe3O4 NPs promoted the photochemical activity of PSII and alleviated photoinhibition by enhancing the photochemical use of excess excitation energy. Gray correlation analysis showed that Fe3O4 NPs enhanced the adaptability of P. heterophylla photosynthesis to high temperatures and strong light. Of note, Fe3O4 NPs enhanced the polysaccharide content of the root tuber. Phytotoxic effect was recorded at high NPs (100 mg·L–1) doses. Collectively, Fe3O4 NPs could promote performance of P. heterophylla by improving photosynthetic performance, enhancing its adaptability to the environment, and increasing the distribution ratio of photosynthates to the underground part.

Quality and yield of Pseudostellaria heterophylla treated with GGBS as pH adjuster against the toxicity of Cd and Cu

The contamination of Cd and Cu in soil is a great threat to medicinal plant. Ground granulated blast furnace slag (GGBS) is a potential soil pH adjuster to reduce metal toxicity. However, how GGBS affects the quality and yield of herbal plants under the stress of Cd and Cu is not clear. This study aims to investigate the quality and yield of a medicinal plant (Pseudostellaria heterophylla) responding to GGBS treatment in Cd and Cu contaminated soil. GGBS with three mass percentages (0%, 3%, 5%) was added into contaminated lateritic soils for planting. Each condition had 21 replicated seedlings. The concentrations of Cd and Cu in plant, amounts of active ingredients (polysaccarides and saponins) in medicinal organ, and tuber properties were measured after harvest. The results showed that under 3% and 5% GGBS treatments, Cd and Cu accumulations in all plant organs (leaf, stem, root and tuber) were reduced by 69.4–86.0% and 10.3–30.1%, respectively. They were below the permissible limits (World Health Organization, WHO). Even though the concentrations of active ingredients in P. heterophylla tuber decreased by up to 35.8%, they still met Hong Kong Chinese Materia Medica standard. Besides, the biomass of root tuber increased by 9.8% and 46%, due to 3% and 5% GGBS treatments, respectively. The recommended 5% GGBS treatment in practice can balance the reduction of active ingredients and the increase of plant yield when minimizing Cd and Cu accumulation in tuber.

Identification of bioactive polysaccharide from Pseudostellaria heterophylla with its anti-inflammatory effects

Abstract Radix Pseudostellariae, the root of Pseudostellaria heterophylla (Miq.) Pax, has historically been used as medicine food homology plant for thousand years in China. Herein, PHP, a water soluble polysaccharides from Radix Pseudostellariae, was isolated and identified by HPGPC, GC–MS, FT-IR, and scanning electron microscopy. Results showed that PHP (Mw = 1.07 × 104 kDa) mainly consisted of rhamnose, galacturonic acid, glucose, galactose, arabinose, fucose in a molar ratio of 0.055:0.208:0.306:0.055:0.753:0.594:0.014. It contained nine types of linkages, including → 6)-Galp(A)-(1→, →6)-Galp-(1→, →3)-Arap-(1→, Galp-(1→, →3)-Rhap-(1→, Glup-(1→, →3)-Araf-(1→, →2,3,4)-Manp-(1 → and → 3,4)-Fucp-(1→. Morphology analysis showed that PHP took flakes with irregular size and properties. It significantly activated TLR4 in vivo. In DSS induced mouse-acute-colitis molds, PHP90-3a improved and the colon length was on ulcerative in mice. the production of IL-1β and TNF-α were decreased. The intestinal mucosa was more complete, there were much more goblet cells, and the infiltration of lymphocytes and neutrophils was significantly decreased. The content of SCFA increased, whereas acetic acid, propionic acid and butyric acid increased. 16S RNA sequencing and OTUs cluster analysis revealed that Bacteroides and Pseudomonas were the dominant intestinal flora. The analysis of community thermogram showed that the abundance ratio of Pseudomonas and Bacteroides (F/B) was significantly lower than the dominant group and significantly inhibited the levels of Verrucomicrobia and Akkermansia. Altogether, the data suggested that the anti-inflammatory activity might be implicated with the activation of TLR4 signal pathways, reducing IL-1β and TNF-α, and the regulation of mucosal immunity and the composition of intestinal flora.

A new cyclic peptide from the fibrous root of Pseudostellaria heterophylla

A new cyclic peptide, Pseudostellarin K (1), together with thirteen known compounds, including two cyclic peptides (2 and 3), one β-carboline alkaloid (4), two amides (5 and 6), three phenylpropanoids (7–9) and other compounds (10–14), were isolated from the fibrous root of Pseudostellaria heterophylla. Their structures were elucidated by extensive spectroscopic analysis. Compounds 1, 4–6, 10 were isolated from the genus pseudostellaria for the first time. All compounds were evaluated for cytotoxic activities against MCF-7, A549, HCT-116 and SGC-7901 cell lines by MTT assay. Unfortunately, all these compounds displayed weak cytotoxic activities.

Long-Read Genome Sequence Resource of Ascochyta versabilis Causing Leaf Spot Disease in Pseudostellaria heterophylla.

Ascochyta versabilis is the fungal pathogen that causes the severe leaf spot disease of Pseudostellaria heterophylla (Miq.) Pax, a vital Chinese herbal plant. Here, we deployed PacBio single-molecule real-time long-read sequencing technology to generate a near-complete genome assembly for the A. versabilis KC1 strain and obtained a total of 9.80 Gb raw reads. These reads were processed into a 41.05 Mb genome assembly containing 95 contigs with N50 of 1.70 Mb and a maximum length of 3.93 Mb. A total of 10,457 gene models, of which 1,004 encode putatively secreted proteins, were identified in the genome. This high-quality genome assembly and gene annotation resource will facilitate the institution of functional genetic studies aimed at providing a better insight into the infection mechanisms of A. versabilis to support the development of effective control strategies for leaf spot disease of P. heterophylla.

Isolation, Identification, and Immunomodulatory Effect of a Peptide from Pseudostellaria heterophylla Protein Hydrolysate.

In this study, a bioactive peptide YGPSSYGYG (YG-9) with immunomodulatory activity was isolated and identified from Pseudostellaria heterophylla protein hydrolysate. The highest proliferation index of mouse spleen lymphocytes reached 1.19 in the presence of 50 μg/mL YG-9. YG-9 could activate RAW264.7 cells by promoting the secretions of NO, the pinocytosis activity, and the productions of ROS and TNF-α. Moreover, YG-9 enhanced the expressions of TLR2 and TLR4 in RAW264.7 cells. TNF-α secretions induced by YG-9 were reduced in TLR2 and TLR4 siRNAs knockdown cells, and this suggested that macrophage activation of YG-9 was through TLR2 and TLR4. Furthermore, YG-9 promoted the translocation of NF-κB through the acceleration of IκB-α phosphorylation and degradation. Also, TNF-α secretions promoted by YG-9 were inhibited by NF-κB-specific inhibitors pyrrolidine dithiocarbamate and BAY11-7082. Altogether, these results suggested YG-9 activated RAW264.7 cells via the TLRs/NF-κB/TNF-α signaling pathway.

Gibberellin disturbs the balance of endogenesis hormones and inhibits adventitious root development of Pseudostellaria heterophylla through regulating gene expression related to hormone synthesis.

The adventitious roots of some plants will develop into tuberous roots which are widely used in many traditional Chinese medicines, including Pseudostellaria heterophylla. If adventitious root development is inhibited, the yield of Chinese medicinal materials will be reduced. Gibberellic acid is an important phytohormone that promotes plant growth and increases the resistance to drought, flood or disease. However, the effects of gibberellic acid on adventitious roots of Pseudostellaria heterophylla are not clear. Here, we reports GA3 suppressed adventitious root development of Pseudostellaria heterophylla by disturbing the balance of endogenesis hormones. By detecting the contents of various endogenous hormones, we found that the development of adventitious roots negatively correlated with the content of CA3 in tuberous roots. Exogenous GA3 treatment decreased the diameter of adventitious roots, but increased the length of adventitious roots of Pseudostellaria heterophylla. In contrast, blocking the biosynthesis of GA3 suppressed stem growth and promoted the xylem of tuberous roots development. Moreover, exogenous GA3 treatment resulted in imbalance of endogenesis hormones by regulating their synthesis-related genes expression in xylem of tuberous roots. These results suggest GA3 broke the established distribution of hormones by regulating synthesis, transport and biological activation of hormones to activate the apical meristem and suppress lateral meristem. Regulating GA3 signaling during adventitious roots development would be one of the possible ways to increase the yield of P. heterophylla.

In Vitro Regeneration, Ex Vitro Rooting and Foliar Stoma Studies of Pseudostellaria heterophylla (Miq.) Pax

Pseudostellaria heterophylla, in the family Caryophyllaceae, is an important Chinese medicinal plant commonly used to treat various diseases in children and valued for its ornamental properties. In this study, nodal segments were obtained from wild plants and used as explants to develop an efficient micropropagation protocol for this species. Murashige and Skoog (MS) medium supplemented with 1.5 mg·L−1 6-benzyladenine (6-BA) was the most suitable medium for inducing axillary buds and enhancing their growth, and MS medium containing 0.1 mg·L−1 indole-3-butyric acid (IBA) was the most effective for inducing in vitro rooting. To reduce labor, time, and cost, microshoots were rooted under ex vitro conditions. Pretreatments of the shoots with 100 mg·L−1 naphthaleneacetic acid (NAA) for 1 min ensured successful rooting in 86.7% of shoots. Comparison of the leaf microstructure between in vitro- and ex vitro-rooted plantlets revealed abnormal stomatal apparatus in the former. The stomatal apparatus of ex vitro plantlets were normal, although the stomatal density was reduced, which indicated that these plantlets were more likely to be able to adapt to environmental conditions in the field. We identified the optimal medium for P. heterophylla multiplication with respect to increased rooting efficiency of micropropagated shoots under ex vitro conditions. This results presented here will be helpful for agricultural cultivation of P. heterophylla.

Industrial applications of selected JFS articles

Industrial applications of selected <i>JFS</i> articles

Detection, isolation, and characterization of the cucumber mosaic virus inPseudostellaria heterophyllafrom Korea

Detection, isolation, and characterization of the cucumber mosaic virus in<i>Pseudostellaria heterophylla</i>from Korea

Cyclic Peptide Extracts Derived From Pseudostellaria heterophylla Ameliorates COPD via Regulation of the TLR4/MyD88 Pathway Proteins.

We have explored the method of extraction and purification of cyclic-peptide extract (CPE) from Pseudostellaria heterophylla (Miq.) Pax. (Taizishen, TZS), characterized the structure about cyclic-peptide compounds and investigated the biological activity of CPE attenuating chronic obstructive pulmonary disease (COPD) in rats. The CPE from TZS was obtained by ethyl acetate, petroleum ether, hot water extraction, and alcohol-precipitation. Cyclic-peptide structures were distinguished using ultra-high performance liquid chromatography-quadrupole time-of-flight tandem mass spectrometry (UPLC-Q-TOF-MS/MS). Rats were induced by solid combustibles smoke (SCS) for the COPD model, and the anti-COPD activity of CPE was detected using lung airway resistance and dynamic lung compliance, as well as pulmonary tissue hematoxylin and eosin (HE) staining. The relevant inflammatory cytokines were assayed by enzyme-linked immunosorbent assay (ELISA). CPE obtained from TZS contained 12 cyclic-peptide constituents; the purity was up to 92.94%. CPE (200, 400, or 500 mg/kg/day) was given to SCS-induced COPD model rats orally for 15 days. The results showed that in rats given CPE (400 mg/kg/day) there was a sharp fall in lung airway resistance but a rise in dynamic lung compliance. The image analysis of lung tissue sections suggested that CPE could decrease the degree of alveolar destruction (p <0.05), alleviate lung inflammation, increase alveolar space, and improve the infiltration of inflammatory cells. CPE was found to reduce the levels of TNF-α, but increase IL-10, adjusting multiple cytokines in rat serum; the TLR4 mRNA, MyD88 mRNA and AP-1 mRNA levels, the expressing levels of p-JNK, p-p38 and p-TAK1 protein were significantly down regulated in rat alveolar macrophages. CPE intervention could improve the pulmonary ventilation function on COPD rats, which may be related to its effect in inhibiting the abnormal activation of the TLR4-MyD88-JNK/p38 pathway. This is the first report that the CPE of TZS lessens the severity of COPD episodes. The new preparation process of CPEs implements the anticipated goal, which is to refine CPE and actualize quality control.

Distribution status, driving force, and protection suggestions of vegetation in Jiangsu Province, China

To provide support for the drawing of 1:500000 vegetation maps and vegetation protection in Jiangsu, the researchers performed a review on the history of research, distribution pattern, change and driving force of vegetation in Jiangsu Province, according to the research results and field investigation of Jiangsu vegetation in recent years. The series research work of the Jiangsu vegetation map began in 1964, and the final map was published in 2001. Vegetation in Jiangsu includes cultivited vegetation, planted and natural forests, herbaceous swamps, meadows, and sparse shrub grass. Cultivited vegetation is distributed in most flat areas, which mainly includes Zea mays , Triticum aestivum , and Oryza sativa . Natural forests are primarily distributed in Xuzhou, Lianyungang, Yili, and Ningzhen mountains. The natural forests include the coniferous forests such as Pinus massoniana , Pinus densiflora forests; the deciduous broad-leaved forests such as Quercus variabilis , Quercus acutissima , and Liquidambar formosana forests; the evergreen broad-leaved forests such as Quercus glauca and Schima superba forests; the bamboo forests such as Phyllostachys heterocycle forest; the coniferous and broad-leaved mixed forests such as Pinus massoniana and Quercus variabilis mixed forests; the evergreen and deciduous broad-leaved mixed forests such as Castanopsis sclerophylla and Quercus glauca mixed forests. Planted forests are mainly distributed in the northern Jiangsu, Lixiahe and coastal plains, including Populus spp, Platycladus orientalis , Cunninghamia lanceolate , and Pinus massoniana forests. The natural wetland vegetation is mainly distributed in the Yancheng coastal wetlands, with Phragmites communis , Spartina anglica , and Spartina alterniflora marshes. Compared with the vegetation in 1983, the area of forests represented by Pinus massoniana , Platycladus orientalis , and Robinia pseudoacacia forests decreased, and herbaceous grasslands and meadows were also drastically reduced. The salt meadows of the coastal wetlands were succeeded by Spartina anglica and Spartina alterniflora swamps, and large areas of agricultural vegetation and a part of the natural forest vegetation were transformed into towns. Human activities led by urbanization affected the distribution pattern of the vegetation in Jiangsu as the major driving force for vegetation change. According to the current situation of natural vegetation in Jiangsu, three suggestions were proposed to protect natural vegetation communities. First, ecological red line protection zones should be established in Nanjing Mufu Mountain, Zhenjiang Baohua Mountain, Lianyungang Shazhou Bay, etc., where the vegetation suffered grave human interference. Second, the development and utilization of herbs and coastal cofferdam, such as Mentha canadensis and Pseudostellaria heterophylla , should be more meticulous. Third, it is necessary to enhance the control of invasive plants, such as Spartina anglica , Solidago canadensis , and Alternanthera philoxeroides .

Pseudostellaria heterophylla Extract Polysaccharide H-1-2 Suppresses Pancreatic Cancer by Inhibiting Hypoxia-Induced AG2

We aimed to examine the therapeutic potential of polysaccharide H-1-2, a bioactive component of Pseudostellaria heterophylla, against pancreatic cancer, as well as to demonstrate the underlying molecular mechanisms. Invasion and migration of pancreatic cells treated with H-1-2 were evaluated. A xenograft tumor mouse model was established to assess the effect of H-1-2 on tumor growth. Expression levels of hypoxic inducible factor-1α (HIF1α) and anterior gradient 2 (AGR2) were measured in pancreatic cells after H-1-2 treatment. Luciferase report and chromatin immunoprecipitation assays were conducted to investigate HIF1α regulation on AGR2. AGR2 expression was re-introduced into pancreatic cells to assess the role of AGR2 as a downstream effector of hypoxia after H-1-2 treatment. H-1-2 inhibited invasion and migration of pancreatic cancer cells, repressed xenograft pancreatic tumor growth, and increased survival of mice. H-1-2 repressed AGR2 expression in pancreatic cancer cells through the hypoxia response element (HRE) in its promoter region. Ectopic AGR2 expression partially negated the H-1-2 inhibitory effect on invasion and migration of pancreatic cells and on xenograft pancreatic tumors growth, and it also compromised the H-1-2 promotional effect on survival of mice. We conclude that H-1-2 suppresses pancreatic cancer by inhibiting hypoxia-induced AGR2 expression, supporting further investigation into its efficacy against pancreatic cancer in clinical settings.

A specific peptide with immunomodulatory activity from Pseudostellaria heterophylla and the action mechanism

Pseudostellaria heterophylla protein hydrolysate has been reported to be a potential nutraceutical supplement to strengthen the immune system. A specific peptide from P. heterophylla protein hydrolysate with proliferation promotion effect on spleen lymphocytes was purified by RP-HPLC. The amino acid sequence of the peptide was determined to be RGPPP, with molecular weight of 522.29 Da. 100 μg/mL of RP-5 achieved the highest stimulation index of 1.27 on spleen lymphocytes and activated RAW264.7 cells resulting in up-regulating tumor necrosis factor-α (TNF-α) secretion to 827.24 pg/mL and elevating pinocytosis rate to 123.81%. Results of western-blot and siRNA experiments indicated that RP-5 activated nuclear factor-κB (NF-κB) signaling pathway through combining to TLR2 expressed on RAW264.7 cells surface. Furthermore, TNF-α secretion increased by RP-5 could be decreased to 401.68 pg/mL by pyrrolidine dithiocarbamate or BAY11-7082. These findings further the research progress of peptide with immunomodulatory activity and its action mechanism.

Modification of Rhizosphere Bacterial Community Structure and Functional Potentials to Control Pseudostellaria heterophylla Replant Disease.

Replant disease caused by negative plant-soil feedback commonly occurs in a Pseudostellaria heterophylla monoculture regime. Here, barcoded pyrosequencing of 16S ribosomal DNA amplicons combined with phylogenetic investigation of communities by reconstruction of unobserved states (PICRUSt) analysis was applied to study the shifts in soil bacterial community structure and functional potentials in the rhizosphere of P. heterophylla under consecutive monoculture and different soil amendments (i.e., bio-organic fertilizer application [MF] and paddy-upland rotation [PR]). The results showed that the yield of tuberous roots decreased under P. heterophylla consecutive monoculture and then increased after MF and PR treatments, which was consistent with the changes in soil bacterial diversity. Both principal coordinate analysis and the unweighted pair-group method with arithmetic means cluster analysis showed the distinct difference in bacterial community structure between the consecutively monocultured soil (relatively unhealthy soil) and other relatively healthy soils (i.e., newly planted soil, MF, and PR). Furthermore, taxonomic analysis showed that consecutive monoculture of P. heterophylla significantly decreased the relative abundances of the families Burkholderiaceae and Acidobacteriaceae (subgroup 1), whereas it increased the population density of families Xanthomonadaceae, Phyllobacteriaceae, Sphingobacteriaceae, and Alcaligenaceae, and Fusarium oxysporum. In contrast, the MF and PR treatments recovered the soil microbiome and decreased F. oxysporum abundance through the different ways; for example, the introduction of beneficial microorganisms (in MF) or the switching between anaerobic and aerobic conditions (in PR). In addition, PICRUSt analysis revealed the higher abundances of membrane transport, cell motility, and DNA repair in the consecutively monocultured soil, which might contribute to the root colonization and survival for certain bacterial pathogens under monoculture. These findings highlight the close association between replant disease of P. heterophylla and the variations in structure and potential functions of rhizosphere bacterial community.

First report of cucumber mosaic virus infecting Pseudostellaria heterophylla in China

First report of cucumber mosaic virus infecting Pseudostellaria heterophylla in China

The Biosynthesis of Heterophyllin B in Pseudostellaria heterophylla From prePhHB-Encoded Precursor.

Plant cyclic peptides (CPs) are a large group of small molecule metabolites found in a wide variety of plants, including traditional Chinese medicinal plants. However, the majority of plant CPs have not been studied for their biosynthetic mechanisms, including heterophyllin B (HB), which is one of the characteristic chemical components of Pseudostellaria heterophylla. Here, we screened the precursor gene (prePhHB) of HB in P. heterophylla and functionally identified its correctness in vivo and in vitro. First, we developed a new method to screen the precursors of HB from 16 candidate linear peptides. According to transcriptome sequencing data, we cloned the genes that encoded the HB precursor peptides and confirmed that the prePhHB-encoded precursor peptide could enzymatically synthesize HB. Next, we generated the transgenic tobacco that expressed prePhHB, and the results showed that HB was detected in transgenic tobacco. Moreover, we revealed that prePhHB gene expression is positively correlated with HB accumulation in P. heterophylla. Mutations in the prePhHB gene may influence the accumulation of HB in P. heterophylla. These results suggest that HB is ribosomally synthesized and posttranslationally modified peptide (RiPP) derived from the precursor gene prePhHB-encoded precursor peptide, and the core peptide sequence of HB is IFGGLPPP in P. heterophylla. This study developed a new idea for the rapid identification of Caryophyllaceae-type CP precursor peptides via RNA-sequencing data mining.