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Plant domestication often alters plant traits, including chemical and physical defenses against herbivores. In squash, domestication leads to reduced levels of cucurbitacins and leaf trichomes, influencing interactions with insects. However, the impact of domestication on inducible defenses in squash remains poorly understood. Here, we investigated the chemical and physical defensive traits of wild and domesticated squash (Cucurbita argyrosperma), and compared their responses to belowground and aboveground infestation by the root-feeding larvae and the leaf-chewing adults of the banded cucumber beetle Diabrotica balteata (Coleoptera: Chrysomelidae). Wild populations contained cucurbitacins in roots and cotyledons but not in leaves, whereas domesticated varieties lacked cucurbitacins in all tissues. Belowground infestation by D. balteata larvae did not increase cucurbitacin levels in the roots but triggered the expression of cucurbitacin biosynthetic genes, irrespective of domestication status, although the response varied among different varieties. Conversely, whereas wild squash had more leaf trichomes than domesticated varieties, the induction of leaf trichomes in response to herbivory was greater in domesticated plants. Leaf herbivory varied among varieties but there was a trend of higher leaf damage on wild squash than domesticated varieties. Overall, squash plants responded to both belowground and aboveground herbivory by activating chemical defense-associated gene expression in roots and upregulating their physical defense in leaves, respectively. While domestication suppressed both chemical and physical defenses, our findings suggest that it may enhance inducible defense mechanisms by increasing trichome induction in response to herbivory., (© 2024. The Author(s).)

To explore the influence of tea trichomes on the quality of white tea, liquid chromatography quadrupole time-of-flight mass spectrometry (LC-Q-TOF-MS), and headspace solid phase microextraction gas chromatography-mass spectrometry (HS-SPME-GC-MS) were used to identify non-volatile and volatile compounds white tea without trichomes (WTwt) and pure trichomes (PT). It was found that the bitter and astringent compounds, caffeine (CAF), epigallocatechin gallate (EGCG), epicatechin gallate (ECG) and flavonol glycosides, were mainly enriched in the WTwt, with 16.3-fold, 47.1-fold and 28.7-fold decrease in CAF and EGCG and ECG, respectively, and the content of these compounds in PT were lower than the taste thresholds. In PT, kaempferol-3-O-(p-coumaroyl)-glucoside and kaempferol-3-O-(di-p-coumaroyl)-glucoside were non-volatile marker compounds, and decanal was significant aroma contributor with rOAV = 250.86. Moreover, the compounds in trichomes mainly contributed to the fruity and floral aroma of white tea, among which benzyl alcohol, (E)-geranylacetone, decanal, dodecanal and 6-methyl-5-hepten-2-one were the crucial aroma components, which were 2.1, 1.7, 1.8, 1.4 and 2.2 times as much as the WTwt in the PT, respectively. In conclusion, trichomes can improve the quality of white tea by reducing the bitterness and astringency, increasing the umami, as well as enhancing the fruity and floral aromas., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Tea trichomes were regarded as an essential evaluation index for reflecting tea flavor quality in terms of aroma and influence on infusion color. This study reveals the impact of golden oxidized trichomes on the color, volatile and non-volatile metabolites of black teas through comparative metabolomics combined quantitative analysis on hongbiluo (trichomes-deficiency black teas), hongjinluo (trichomes-rich black teas), and trichomes (from hongjinluo). Forty-six volatile components were detected using headspace solid-phase microextraction gas chromatography-mass spectrometry, while the results suggested that the contribution of trichomes to black teas is limited. A total of 60 marker non-volatile compounds were identified, including catechins, catechin oxidation products, flavonoid glycosides, organic acids, hydrolysable tannins and amino acids. Notably, p-coumaroyl-kaempferol glucosides, and catechin dimers demonstrated high levels in independent trichomes and showed a positive correlation with the brightness and yellow hue of black tea infusions, specifically kaempferol 3-O-di-(p-coumaroyl)-hexoside. Furthermore, results from fractional extraction analysis of separated trichomes provided that N-ethyl-2-pyrrolidinone-substituted epicatechin gallates, acylated kaempferol glycosides, and chromogenic catechins dimers, such as theaflavins, were primary color contributors in oxidized trichomes. Especially, we found that epicatechin gallate (ECG) and its derivates, 3'-O-methyl-ECG and N-ethyl-2-pyrrolidinone-substituted ECG, highly accumulated in trichomes, which may be associated with the varieties of hongbiluo and hongjinluo black teas. Eventually, addition tests were applied to verify the color contribution of trichome mixtures. Our findings employed comprehensive information revealing that golden oxidized trichomes contributed significantly to the brightness and yellow hue of black tea infusion, but their contribution to the aroma and metabolic profile is limited. These findings may contribute to the effective modulation of the infusion color during black tea production by regulating the proportion of tea trichomes or screening trichomes-rich or deficiency varieties., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Cembranoids and labdanes are two important types of diterpenes in tobacco (Nicotiana genus) that are predominantly found in the leaf and flower glandular trichome secretions. This is the first systematic review of the biosynthesis, chemical structures, bioactivities, and utilisation values of cembranoid and labdane diterpenes in tobacco. A total of 131 natural cembranoid diterpenes have been reported in tobacco since 1962; these were summarised and classified according to their chemical structure characteristics as isopropyl cembranoids (1-88), seco-cembranoids (89-103), chain cembranoids (104-123), and polycyclic cembranoids (124-131). Forty natural labdane diterpenes reported since 1961 were also summarised and divided into epoxy side chain labdanes (132-150) and epoxy-free side chain labdanes (151-171). Tobacco cembranoid and labdane diterpenes are both formed via the methylerythritol 4-phosphate pathway and are synthesised from geranylgeranyl diphosphate. Their biosynthetic pathways and the four key enzymes (cembratrienol synthase, cytochrome P450 hydroxylase, copalyl diphosphate synthase, and Z-abienol cyclase) that affect their biosynthesis have been described in detail. A systematic summary of the bioactivity and utilisation values of the cembranoid and labdane diterpenes is also provided. The agricultural bioactivities associated with cembranoid and labdane diterpenes include antimicrobial and insecticidal activities as well as induced resistance, while the medical bioactivities include cytotoxic and neuroprotective activities. Further research into the cembranoid and labdane diterpenes will help to promote their development and utilisation as plant-derived pesticides and medicines., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Cannabis glandular trichomes produce and store an abundance of lipidic specialised metabolites (e.g. cannabinoids and terpenes) that are consumed by humans for medicinal and recreational purposes. Due to a lack of genetic resources and inherent autofluorescence of cannabis glandular trichomes, our knowledge of cannabinoid trafficking and secretion is limited to transmission electron microscopy (TEM). Advances in cryofixation methods has resulted in ultrastructural observations closer to the 'natural state' of the living cell, and recent reports of cryofixed cannabis trichome ultrastructure challenge the long-standing model of cannabinoid trafficking proposed by ultrastructural reports using chemically fixed samples. Here, we compare the ultrastructural morphology of cannabis glandular trichomes preserved using conventional chemical fixation and ultrarapid cryofixation. We show that chemical fixation results in amorphous metabolite inclusions surrounding the organelles of glandular trichomes that were not present in cryofixed samples. Vacuolar morphology in cryofixed samples exhibited homogenous electron density, while chemically fixed samples contained a flocculent electron dense periphery and electron lucent lumen. In contrast to the apparent advantages of cryopreservation, fine details of cell wall fibre orientation could be observed in chemically fixed glandular trichomes that were not seen in cryofixed samples. Our data suggest that chemical fixation results in intracellular artefacts that impact the interpretation of lipid production and trafficking, while enabling greater detail of extracellular polysaccharide organisation., (© 2022 Royal Microscopical Society.)

Background: Lavender (genus Lavandula, family Lamiaceae) is an aromatic plant widely grown as an ornamental plant. The chemical composition of lavender is characterized by monoterpenoids, sesquiterpenoids, and other compounds, which are primarily synthesized and stored in epidermal secretory structures called glandular trichomes (GTs). Volatile organic compounds (VOCs) are responsible for the aroma characteristics of plant oil that drive consumer preference. Aroma is usually regarded as a characteristic trait for the classification of aromatic plants. Interestingly, VOCs are synthesized and stored in GTs. Lamiaceae species such as purple perilla, peppermint, basil, thyme, and oregano usually possess two types of GTs: peltate glandular trichomes (PGTs) and capitate glandular trichomes (CGTs). But the development process of PGTs in lavender has been reported in only a few studies to date., Results: In this study, we identified and quantified the VOCs in four lavender cultivars by headspace-solid phase micro extraction-gas chromatography mass spectrometry (HS-SPME-GC-MS). A total of 66 VOCs were identified in these four cultivars, the most prominent of which were linalyl acetate and linalool, and flowers were the main site of accumulation of these VOCs. Here, we examined the developmental process of PGTs, including the formation of their base, body, and apex. The apex cells contained secretory cavities, which produced VOCs. Based on the reference genome sequence of the lavender cultivar 'Jingxun 2', several R2R3-MYB subfamily genes related to GT formation were identified. These results will guide the engineering of GTs and molecular breeding of lavender for improving the VOC content., Conclusions: In this study, we identified the VOCs in four lavender cultivars. We analyzed the formation of GTs, and compared the number and diameter size of PGTs among four lavender cultivars. Additionally, we identified four candidate genes belonging to the R2R3-MYB family., (© 2023. The Author(s).)

Introduction: In recent years, industrial production of Cannabis sativa has increased due to increased demand of medicinal products based on the plant. In these medicinal products, it is mainly the contents of cannabinoids like THCA and CBDA which are of interest, but also the flavonoids of C. sativa have pharmaceutical interest., Objectives: The primary aim is to study the distribution of the different cannabinoids in leaves of C. sativa and specifically to which extent they are located on the trichomes found on the surface of C. sativa leaves. Desorption electrospray ionization (DESI) and matrix assisted laser desorption ionization (MALDI) mass spectrometry imaging (MSI) provide non-targeted imaging of numerous compounds in the same experiment. Therefore, the distribution of flavonoids is also mapped in the same experiments., Material and Methods: Fan leaves from C. sativa were imaged in the lateral dimension using direct DESI-MSI as well as indirect DESI-MSI via a porous PTFE surface using pixel sizes of 150-200 μm. For cross sections of sugar leaves, MALDI-MSI was performed at 20 μm pixel size., Results: From indirect DESI-MSI experiments, a connection was made between the cannabinoid CBGA and capitate-stalked trichomes. Other cannabinoids like THCA/CBDA (isomers, which are not resolved in an MSI experiment) were also detected in the capitate-stalked trichomes, but in addition to this also in the small glandular trichomes. MALDI-MSI experiments on cross sections of sugar leaves confirmed that the cannabinoids were not an integral part of the leaf tissue itself, but originated from the trichomes on the surface of the leaf., Conclusion: The study provides visual evidence that the cannabinoids are produced and accumulated in the trichomes of C. sativa leaves., (© 2023 The Authors. Phytochemical Analysis published by John Wiley & Sons Ltd.)

This work represents the first multi-scale study on Teucrium fruticans L. cultivated at the Ghirardi Botanic Garden (Lombardy, Northern Italy), combining a micromorphological and a phytochemical survey on the plant's aerial parts. Micromorphological investigations, performed by Light Microscopy, Fluorescence Microscopy and Scanning Electron Microscopy, highlighted the presence of five trichomes morphotypes, distinguished by a different distribution pattern: peltates, short-stalked and ball-like medium-stalked capitates, ubiquitous on the whole plant, medium-stalked and long-stalked capitates, exclusive to the floral whorls. Both peltates and medium-stalked capitates were recognized as the main terpene production sites. Phytochemical characterization focused on the essential oils (EOs), obtained by Clevenger-type hydrodistillation in February and April 2022 and characterized by Gas Chromatography-Mass Spectrometry (GC/MS), which resulted mainly formed by sesquiterpene hydrocarbons. The February EO profile was characterized by β-caryophyllene (28.30 %) and germacrene D (19.16 %) as main compounds, while in April β-myrcene was detected at high percentage (13.77 %), in addition to the previous two components (15.72 % and 11.55 %, respectively). Literature data, dealing with the biological activities of the main oil constituents, highlighted an anti-microbial, anti-inflammatory, and anti-tumor potential, due to the high content in sesquiterpenes and, particularly, of β-caryophyllene and germacrene D., (© 2023 The Authors. Chemistry & Biodiversity published by Wiley-VHCA AG, Zurich, Switzerland.)

In the present scenario, remediation of heavy metals (HMs) contaminated soil has become an important work to be done for the well-being of human and their environment. Phytoremediation can be regarded as an excellent method in environmental technologies. The present contemporary research explores the Solanum viarum Dunal function as a potential accumulator of hazardous HMs viz. lead (Pb), cadmium (Cd), zinc (Zn), and their combination (CHM). On toxic concentrations of Pb, Cd, Zn, and their synergistic exposure, seeds had better germination percentage and their 90d old aerial tissues accumulated Pb, Cd, and Zn concentrations ranging from 44.53, 84.06, and 147.29 mg kg -1 DW, respectively. Pattern of accumulation in roots was as Zn 70.08 > Pb 48.55 > Cd 42.21 mg kg -1 DW. Under HMs treatment, positive modulation in physiological performances, antioxidant activities suggested an enhanced tolerance along with higher membrane stability due to increased levels of lignin, proline, and sugar. Phenotypic variations were recorded in prickles and roots of 120 d old HM stressed plants, which are directly correlated with better acclimation. Interestingly, trichomes of the plant also showed HM accumulation. Later, SEM-EDX microanalysis suggested involvement of S. viarum capitate glandular trichomes as excretory organs for Cd and Zn. Thus, the present study provides an understanding of the mechanism that makes S. viarum to function as potent accumulator and provides information to generate plants to be used for phytoremediation., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Mentha haplocalyx Briq (M. haplocalyx) is a herbaceous plant that has long been used as a food, medicinal spice, and flavoring agent in traditional Chinese medicine. Its secondary metabolites, having high commercial values, are mainly produced in tiny specialized structures called glandular trichomes (GTs). The primary purpose of this study was to examine the morphology and metabolites of peltate GTs in M. haplocalyx.Peltate GTs possessed globular dome shapes and intense auto-fluorescence on the surfaces of M. haplocalyx leaves. Structure subsidence and cuticle rupture were found throughout the aging stage of peltate GTs. According to histochemical staining results, the secretion of peltate GTs contained anthraquinone, flavonoids, phenolic acid and terpenoids. In M. haplocalyx peltate GTs and leaf tissues without peltate glandular trichomes, ten and two volatile compounds were identified respectively. Peltate GTs contained 42 non-volatile chemicals with a variety of structural types, including 20 flavonoids, 17 phenolic acids,1 diterpene, 3 anthraquinone and 1 alkane. Meanwhile, 15 non-volatile compounds were discovered in leaf tissues without peltate glandular trichomes, and they were all included in the list of peltate GTs' 41 components. Therefore, Peltate GTs were shown to be the primary site of not just volatile compounds but also non-volatile chemicals in M. haplocalyx. This study provides an important theoretical basis and technical approach for clarifying the bio-active metabolite biosynthesis in M. haplocalyx., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Buddleja officinalis is a traditional Chinese medicinal plant covered with glandular and non-glandular trichomes on leaves. Phytochemical investigation of its leaves led to the identification of one undescribed tetranorcycloartane 3-oxo-25,26,27,29-tetranorcycloartan-24-oic acid (1) and one first identified natural product tetranorcycloartane 3-oxo-25,26,27,29-tetranorcycloartan-24-oic methyl ester (2), along with an undescribed megastigmane glucoside (3) and 14 known constituents (4-17). Structures of undescribed chemicals were elucidated by comprehensive 1D and 2D NMR, MS and CD analysis. Further chemical investigation resulted in six triterpenoids (4-9) being localized to the trichomes of B. officinalis. The major trichome components cycloeucalenone (4) and 24-oxo-29-norcycloartan-3-one (5) showed potent antifeedant activity against a generalist insect cotton bollworm (Helicoverpa armigera), but no obvious activity against the specialist herbivore Hyphasis inconstans. Compounds 4 and 7 also displayed inhibitory effects on seed germination of Arabidopsis thaliana. In addition, 1 and 4 exhibited moderate antibacterial activity toward three gram-positive bacteria., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Despite its use by humans for thousands of years, the technology of cannabis usage and extraction is still evolving. Given that the primary pharmacological compounds of interest are cannabinoid and terpenoids found in greatest abundance in capitate glandular trichomes of unfertilized female inflorescences, it is surprising that older techniques of hashish making have received less technological advancement. The purpose of this study was to employ organically grown cannabis and to isolate pure trichomes from freshly picked flowers via exposure to vapor from solid CO 2 , commonly known as "dry ice", followed by their isolation via sifting through a 150 µ screens while maintaining the cold chain. Biochemical analysis was undertaken on fresh flower, frozen-sifted flower by-products, treated trichomes (Kryo-Kief™), dried flower, dried sifted flower by-product and dried kief. The dry ice process successfully concentrated cannabinoid content as high as 60.7%, with corresponding concentration and preservation of monoterpenoids encountered in fresh flower that are usually lost during the conventional cannabis drying and curing process. The resulting dried sifted flower by-product after dry ice processing remains a usable commodity. This approach may be of interest to pharmaceutical companies and supplement producers pursuing cannabis-based medicine development with an eye toward full synergy of ingredients harnessing the entourage effect.

The aim of this study was to conduct a histochemical analysis to localize lipids, terpenes, essential oil, and iridoids in the trichomes of the L. album subsp. album corolla. Morphometric examinations of individual trichome types were performed. Light and scanning electron microscopy techniques were used to show the micromorphology and localization of lipophilic compounds and iridoids in secretory trichomes with the use of histochemical tests. Additionally, the content of essential oil and its components were determined using gas chromatography-mass spectrometry (GC-MS). Qualitative analyses of triterpenes carried out using high-performance thin-layer chromatography (HPTLC) coupled with densitometric detection, and the iridoid content expressed as aucubin was examined with spectrophotometric techniques. We showed the presence of iridoids and different lipophilic compounds in papillae and glandular and non-glandular trichomes. On average, the flowers of L. album subsp. album yielded 0.04 mL/kg of essential oil, which was dominated by aldehydes, sesquiterpenes, and alkanes. The extract of the L. album subsp. album corolla contained 1.5 × 10 -3 ± 4.3 × 10 -4 mg/mL of iridoid aucubin and three triterpenes: oleanolic acid, β-amyrin, and β-amyrin acetate. Aucubin and β-amyrin acetate were detected for the first time. We suggest the use of L. album subsp. album flowers as supplements in human nutrition.

Background: Perilla frutescens (L.) Britt is a medicinal and edible plant widely cultivated in Asia. Terpenoids, flavonoids and phenolic acids are the primary source of medicinal ingredients. Glandular trichomes with multicellular structures are known as biochemical cell factories which synthesized specialized metabolites. However, there is currently limited information regarding the site and mechanism of biosynthesis of these constituents in P. frutescens. Herein, we studied morphological features of glandular trichomes, metabolic profiling and transcriptomes through different tissues., Results: Observation of light microscopy and scanning electron microscopy indicated the presence of three distinct glandular trichome types based on their morphological features: peltate, capitate, and digitiform glandular trichomes. The oil of peltate glandular trichomes, collected by custom-made micropipettes and analyzed by LC-MS and GC-MS, contained perillaketone, isoegomaketone, and egomaketone as the major constituents which are consistent with the components of leaves. Metabolomics and transcriptomics were applied to explore the bioactive constituent biosynthesis in the leaves, stem, and root of P. frutescens. Transcriptome sequencing profiles revealed differential regulation of genes related to terpenoids, flavonoids, and phenylpropanoid biosynthesis, respectively with most genes expressed highly in leaves. The genes affecting the development of trichomes were preliminarily predicted and discussed., Conclusions: The current study established the morphological and chemical characteristics of glandular trichome types of P. frutescens implying the bioactive constituents were mainly synthesized in peltate glandular trichomes. The genes related to bioactive constituents biosynthesis were explored via transcriptomes, which provided the basis for unraveling the biosynthesis of bioactive constituents in this popular medicinal plant.

Cannabis sativa has been cultivated since antiquity as a source of fibre, food and medicine. The recent resurgence of C. sativa as a cash crop is mainly driven by the medicinal and therapeutic properties of its resin, which contains compounds that interact with the human endocannabinoid system. Compared to other medicinal crops of similar value, however, little is known about the biology of C. sativa. Glandular trichomes are small hair-like projections made up of stalk and head tissue and are responsible for the production of the resin in C. sativa. Trichome productivity, as determined by C. sativa resin yield and composition, is only beginning to be understood at the molecular level. In this study the proteomes of glandular trichome stalks and heads, were investigated and compared to the proteome of the whole flower tissue, to help further elucidate C. sativa glandular trichome biochemistry. The data suggested that the floral tissue acts as a major source of carbon and energy to the glandular trichome head sink tissue, supplying sugars which drive secondary metabolite biosynthesis. The trichome stalk seems to play only a limited role in secondary metabolism and acts as both source and sink., Competing Interests: First author LJK received a partial stipend from the commercial entity Cann Group Limited. The authors have declared that no competing interests exist. This does not alter our adherence to PLOS ONE policies on sharing data and materials.

Urban green infrastructure is closely linked to the alleviation of pollution from atmospheric particulate matter. Although particle deposition has been shown to depend on leaf characteristics, the findings from earlier studies are sometimes ambiguous due to the lack of controlling variables. In this study, we investigated the impact of leaf morphological characteristics on PM 2.5 dry deposition velocity by employing a control-variable approach. We focused on four indices: trichome density, petiole length, aspect ratio (width-to-length ratio), and fractal deviation. For each index, tree species were chosen from the same family or genus to minimize the influence of other factors and make a group of treatments for an individual index. The dry deposition velocities of PM 2.5 were determined through application of an indirect method. The results revealed that the presence of leaf trichomes had a positive effect on PM 2.5 dry deposition velocity, and a higher trichome density also led to a greater particle deposition velocity. Lower leaf aspect ratio, shorter petioles, and higher leaf fractal deviation were associated with greater PM 2.5 dry deposition velocity. The control-variable approach allows to investigate the correlation between deposition velocity and a certain leaf characteristic independently while minimizing the effects of others. Thus, our study can clarify how a single leaf characteristic affects particle deposition velocity, and expound its potential mechanism more scientifically than the published studies. Our research points out the importance of controlling variables, and also provides ideas for future researches on related factors to be found. Meanwhile the results would help provide insight into design improvements or adaptive management for the alleviation of air pollution., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Main Conclusion: Three types of the glandular trichomes are developed on the flowers and leaves of Millingtonia hortensis. Morphology, cell ultrastructure and content of the volatile compounds are specific to each trichome type. The aim of this study was to characterize the structural and histochemical features of the glandular trichomes (GTs) of two types localized on the different flower parts and leaves in Millingtonia hortensis, as well as to identify the composition of the internal pool of metabolites. The peltate GTs are most common; they are founded on peduncle, calyx, ovary, and leaves. GTs consist of 12-24-cell disk-shaped head and a single-celled neck. The capitate GTs are located on corolla tube and have four to eight-cell head, single-celled neck and a wide multicellular stalk. A series of histochemical reactions and fluorescent microscopy revealed the various substances in the chemical composition of GTs. Acid polysaccharides are predominately identified in the capitate trichomes of the corolla tube and peltate trichomes of calyx, terpenes present in larger quantity in the trichomes of the corolla tube and ovary, whilst phenolic substances prevail in the trichomes of the calyx and ovary. GTs of each type are characterized by specific ultrastructural traits. Smooth endoplasmic reticulum (SER) and leucoplasts prevail in the peltate trichomes of peduncle, calyx and ovary; Golgi apparatus is the common organelle in the capitate trichomes of the corolla tube and peltate trichomes of calyx; the huge aggregates of the RER cisterns there are in cytoplasm of all leaf trichomes. Synthesized secretion accumulates in the subcuticular cavity of all GTs except the leaf peltate trichomes. In the trichomes of the leaves secretion is stored in the thick upper cell wall with the wide cutinized layer. For the first time content of the internal pool of metabolites from the flowers and leaves was identified by GC-MS. Seventeen compounds, including alcohols, fatty acid derivatives, monoterpenes, sesquiterpenes, and benzenoids were identified. 1-octen 3-ol, 3-carene, methyl salicylate, p-hydroxybenzeneethanol and 1-hydroxy-2,4-di-tertbutyl-benzene were the main compounds of the flower scent. We consider GTs of the reproductive organs in M. hortensis synthesizing acid polysaccharides and volatile compounds as secretory structures attracting of pollinators, whereas the leaf peltate trichomes accumulating predominately non-volatile phenols, protect young vegetative shoots against small herbivorous insects and pathogens.

Flos Lamii albi has a high biological activity and is widely used in herbal medicine. The aim of the study was to characterize the secretory structures present in Lamium album subsp. album corolla and the location of phenolic compounds. Additionally, we carried out qualitative phytochemical analyses of flavonoids and phenolic acids. Light, fluorescence, and scanning electron microscopy were used to analyze the structure of the floral organs. The main classes of phenolic compounds and their localization were determined histochemically. Phytochemical analyses were performed with high-performance thin-layer chromatography (HPTLC) and high-performance liquid chromatography (HPLC). Six types of glandular trichomes were found which contained flavonoids, phenolic acids, and tannins. The phytochemical studies demonstrated the presence of caffeic, chlorogenic, ferulic, gallic, p-coumaric, protocatechuic, syringic, gentisic, and vanillic phenolic acids as well as rutoside, isoquercetin, and quercetin flavonoids. The corolla in L. album subsp. album has antioxidant properties due to the presence of various polyphenols, as shown by the histo- and phytochemical analyses. The distribution and morphology of trichomes and the content of phenolic compounds in the corolla have taxonomic, pharmacognostic, and practical importance, facilitating the identification of the raw material.

Main Conclusion: Permanent glandular trichomes of Robinia viscosa var. hartwigii produce viscous secretion containing several secondary metabolites, as lipids, mucilage, flavonoids, proteins and alkaloids. Robinia viscosa var. hartwigii (Hartweg's locust) is an ornamental tree with high apicultural value. It can be planted in urban greenery and in degraded areas. The shoots, leaves, and inflorescences of this plant are equipped with numerous persistent glandular trichomes producing sticky secretion. The distribution, origin, development, morphology, anatomy, and ultrastructure of glandular trichomes of Hartweg's locust flowers as well as the localisation and composition of their secretory products were investigated for the first time. To this end, light, scanning, and transmission electron microscopy combined with histochemical and fluorescence techniques were used. The massive glandular trichomes differing in the distribution, length, and stage of development were built of a multicellular and multiseriate stalk and a multicellular head. The secretory cells in the stalk and head had large nuclei with nucleoli, numerous chloroplasts with thylakoids and starch grains, mitochondria, endoplasmic reticulum profiles, Golgi apparatus, vesicles, and multivesicular bodies. Many vacuoles contained phenolic compounds dissolved or forming various condensed deposits. The secretion components were transported through symplast elements, and the granulocrine and eccrine modes of nectar secretion were observed. The secretion was accumulated in the subcuticular space at the trichome apex and released through a pore in the cuticle. Histochemical and fluorescence assays showed that the trichomes and secretion contained lipophilic and polyphenol compounds, polysaccharides, proteins, and alkaloids. We suggest that these metabolites may serve an important function in protection of plants against biotic stress conditions and may also be a source of phytopharmaceuticals in the future.

This study presented a micromorphological and phytochemical survey on Lavandula dentata L. cultivated at the Ghirardi Botanic Garden (Toscolano Maderno, BS, Italy). The morphological investigation revealed the presence of peltate, short- and medium-stalked capitate trichomes. The histochemical survey showed terpene production by peltates and medium-stalked capitates, hydrophilic secretions by short-stalked capitates. The phytochemical survey was developed on leaf and flower volatile organic compounds (VOCs) and on the essential oil (EO) from the flowering aerial parts. The VOC profiles represented an element of novelty and were dominated by oxygenated monoterpenes, among which 1,8-cineole and β-pinene were the most abundant (77.40 %, 7.11 % leaves; 81.08 %, 10.46 % flowers). The EO of L. dentata was dominated by oxygenated monoterpenes with a high percentage of 1,8-cineole (69.08 %), followed by β-pinene, trans-pinocarveol and myrtenal. Evaluations about the ecological role, the potential biological activity and the sensory attributes were proposed, based on literature contributions., (© 2020 Wiley-VHCA AG, Zurich, Switzerland.)

Tea trichomes contain special flavor-determining metabolites; however, little is known about how and why tea trichomes produce them. Integrated metabolite and transcriptome profiling on tea trichomes in comparison with that on leaves showed that trichomes contribute to tea plant defense and tea flavor and nutritional quality. These unicellular, nonglandular, and unbranched tea trichomes produce a wide array of tea characteristic metabolites, such as UV-protective flavonoids, insect-toxic caffeine, herbivore-defensive volatiles, and theanine, as evidenced by the expression of whole sets of genes involved in different metabolic pathways. Both dry and fresh trichomes contain several volatiles and flavonols that were not found or at much low levels in trichome-removed leaves, including benzoic acid derivatives, lipid oxidation derivatives, and monoterpene derivatives. Trichomes also specifically expressed many disease signaling genes and various antiherbivore or antiabiotic peptides. Trichomes are one of the domestication traits in tea plants. Tea trichomes contribute to tea plant defenses and tea flavors.

Genus Ocimum is known to have species possessing important therapeutic essential oil. The major phytoconstituents of essential oil in Ocimum species are phenylpropanoids and terpenoids. The essential oil is accumulated in the trichomes; the specialized structures predominantly found on leaves and other tissues. The development of trichome is integrated with development of plant and leaf and also tightly coordinated with the primary and secondary metabolic pathways producing essential oil constituents. In continuation to our studies on elucidating/understanding the mechanism of biosynthesis of  essential oil pathways in Ocimum species, we have performed comparative transcriptome analysis to investigate the role of trichome-related gene expression in the regulation of biosynthetic pathways of essential oil. The essential oil biogenesis is tightly integrated with primary metabolic activities, the analysis for the expression pattern of genes related to primary metabolism and its relationship with secondary metabolism was evaluated in comparative manner. Physiological parameters in relation to primary metabolism such as photosynthetic pigment content, soluble sugar content, and invertase enzymes along with morphological parameters were analysed in O. basilicum and O. sanctum. Differential expression profiling uncovered about 8116 and 2810 differentially expressed transcripts in O. basilicum and O. sanctum, respectively. Enrichment of differentially expressed genes were analysed in relation to metabolic pathways, primary metabolism and secondary metabolism. Trichome related genes identified from the Ocimum species vis-à-vis their expression profiles suggested higher expression in O. basilicum. The findings in this study provide interesting insights into the role of trichome-related transcripts in relation to essential oil content in Ocimum species. The study is valuable as this is the first study on revealing the transcripts and their role in trichome development and essential oil biogenesis in two major species of Ocimum.

Interspecific New Rice for Africa (NERICA) varieties have been recently developed and used in Sub-Saharan Africa but herbivore resistance properties of these plants remain poorly understood. Here we report that, compared to a local Japanese cultivar Nipponbare, NERICA 1, 4 and 10 are significantly more damaged by insect herbivores in the paddy fields. In contrast to high levels of leaf damage from rice skippers and grasshoppers, constitutive and induced volatile organic compounds for indirect plant defense were higher or similar in NERICAs and Nipponbare. Accumulation of direct defense secondary metabolites, momilactones A and B, and p-coumaroylputrescine (CoP) was reduced in NERICAs, while feruloylputrescine accumulated at similar levels in all varieties. Finally, we found that Nipponbare leaves were covered with sharp nonglandular trichomes impregnated with silicon but comparable defense structures were virtually absent in herbivory-prone NERICA plants. As damage to the larval gut membranes by Nipponbare silicified trichomes that pass intact through the insect digestive system, occurs, and larval performance is enhanced by trichome removal from otherwise chemically defended Nipponbare plants, we propose that silicified trichomes work as an important defense mechanism of rice against chewing insect herbivores., (© 2020 John Wiley & Sons Ltd.)

Glandular trichomes (GTs) are defensive structures that produce and accumulate specialized metabolites and protect plants against herbivores, pathogens, and abiotic stress. GTs have been extensively studied in angiosperms for their roles in defense and biosynthesis of high-value metabolites. In contrast, trichomes of gymnosperms have been described in fossilized samples, but have not been studied in living plants. Here, we describe the characterization of GTs on young stems of a hybrid white spruce. Metabolite and histological analysis of spruce GTs support a glandular function with accumulation of a diverse array of mono-, sesqui- and diterpenes including diterpene methylesters. Methylated diterpenes have previously been associated with insect resistance in white spruce. Headspeace analysis of spruce GTs showed a profile of volatiles dominated by monoterpenes and a highly diverse array of sesquiterpenes. Spruce GTs appear early during shoot growth, prior to the development of a lignified bark and prior to accumulation of terpenes in needles. Spruce GTs may provide an early, terpene-based chemical defense system at a developmental stage when young shoots are particularly vulnerable to foliage and shoot feeding insects, and before the resin duct system characteristic of conifers has fully developed.

The removal of particulate matters (PM) has emerged as one of the most significant issues in public health and environment worldwide. Environmentalists have proposed the use of indoor air-purifying plants as an eco-friendly strategy to resolve PM-related problems and effectively remove fine particulate matter (PM 2.5 ). Among air-purifying plants, Tillandsia usneoides (L.) L. (T. usneoides) has been used as a biomonitor for heavy metals and air pollutants. However, the PM removal effect of T. usneoides and its primary mechanism remain unclear. Here, we investigated the PM removal performance of T. usneoides in a closed chamber under flow conditions, the effects of trichomes, and the array density according to the different types of PM. The chamber with bulk T. usneoides under flow conditions exhibited 16.5 % and 9.2 % higher removal efficiency in PM 2.5 T. usneoides for incense and A1 rigid PM, respectively, than that without T. usneoides. T. usneoides with trichome structure exhibited larger removal efficiencies of 7% and 2% in PM 2.5 and PM 10 , respectively, than without trichome for incense particles. In addition, the increase in total effective surface was effective for the deposition of both PM types. The increase in effective surface area by trichome structure and array density of T. usneoides is a crucial factor for the deposition of PM., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Ocimum is one of the most revered medicinally useful plants which have various species. Each of the species is distinct in terms of metabolite composition as well as the medicinal property. Some basil types are used more often as an aromatic and flavoring ingredient. It would be informative to know relatedness among the species which though belong to the same genera while exclusively different in terms of metabolic composition and the operating pathways. In the present investigation the similar effort has been made in order to differentiate three commonly occurring Ocimum species having the high medicinal value, these are Ocimum sanctum, O. gratissimum and O. kilimandscharicum. The parameters for the comparative analysis of these three Ocimum species comprised of temporal changes in number leaf trichomes, essential oil composition, phenylpropanoid pathway genes expression and the activity of important enzymes. O. gratissimum was found to be richest in phenylpropanoid accumulation as well as their gene expression when compared to O. sanctum while O. kilimandscharicum was found to be accumulating terpenoid. In order to get an overview of this qualitative and quantitative regulation of terpenes and phenylpropenes, the expression pattern of some important transcription factors involved in secondary metabolism were also studied.

Trichomes, which develop from epidermal cells, are regarded as one of the key features that are involved in the evaluation of tea quality and tea germplasm resources. The metabolites from trichomes have been well characterized in tea products. However, little is known regarding the metabolites in fresh tea trichomes and the molecular differences in trichomes and tea leaves per se. In this study, we developed a method to collect trichomes from tea plant tender shoots, and their main secondary metabolites, including catechins, caffeine, amino acids, and aroma compounds, were determined. We found that the majority of these compounds were significantly less abundant in trichomes than in tea leaves. RNA-Seq was used to investigate the differences in the molecular regulatory mechanism between trichomes and leaves to gain further insight into the differences in trichomes and tea leaves. In total, 52.96 Gb of clean data were generated, and 6560 differentially expressed genes (DEGs), including 4471 upregulated and 2089 downregulated genes, were identified in the trichomes vs. leaves comparison. Notably, the structural genes of the major metabolite biosynthesis pathways, transcription factors, and other key DEGs were identified and comparatively analyzed between trichomes and leaves, while trichome-specific genes were also identified. Our results provide new insights into the differences between tea trichomes and leaves at the metabolic and transcriptomic levels, and open up new doors to further recognize and re-evaluate the role of trichomes in tea quality formation and tea plant growth and development.

Plant nanobiotechnology has the potential to revolutionize agriculture. However, the lack of effective methods to deliver nanoparticles (NPs) to the precise locations in plants where they are needed impedes these technological innovations. Here, model gold nanoparticles (AuNP) were coated with citrate, bovine serum albumin (BSA) as a protein control, or LM6-M, an antibody with an affinity for functional groups unique to stomata on leaf surfaces to deliver the AuNPs to stomata. One-month-old Vicia faba leaves were exposed via drop deposition to aqueous suspensions of LM6-M-coated AuNPs and allowed to air dry. After rinsing, Au distribution on the leaf surface was investigated by enhanced dark-field microscopy and X-ray fluorescence mapping. While citrate-coated AuNPs randomly covered the plant leaves, LM6M-AuNPs strongly adhered to the stomata and remained on the leaf surface after rinsing, and BSA-AuNPs specifically targeted trichome hairs. To the authors' knowledge, this is the first report of active targeting of live leaf structures using NPs coated with molecular recognition molecules. This proof-of-concept study provides a strategy for future targeted nanopesticide delivery research.

Obesity that is highly associated with numerous metabolic diseases has become a global health issue nowdays. Plant sesterterpenoids are an important group of natural products with great potential; thus, their bioactivities deserve extensive exploration. RNA-seq analysis indicated that leucosceptroid B, a sesterterpenoid previously discovered from the glandular trichomes of Leucosceptrum canum, significantly regulated the expression of 10 genes involved in lipid metabolism in Caenorhabditis elegans. Furthermore, leucosceptroid B was found to reduce fat storage, and downregulate the expression of two stearoyl-CoA desaturase (SCD) genes fat-6 and fat-7, and a fatty acid elongase gene elo-2 in wild-type C. elegans. In addition, leucosceptroid B significantly decreased fat accumulation in both fat-6 and fat-7 mutant worms but did not affect the fat storage of fat-6; fat-7 double mutant. These findings indicated that leucosceptroid B reduced fat storage depending on the downregulated expression of fat-6, fat-7 and elo-2 and thereby inhibiting the biosynthesis of the corresponding unsaturated fatty acid. These findings provide new insights into the development and utilization of plant sesterterpenoids as potential antilipemic agents., (Copyright © 2019 China Pharmaceutical University. Published by Elsevier B.V. All rights reserved.)

Background: One of the roles of plant trichomes is thought to be reducing feeding damage from herbivores. Among trichomes, glandular trichomes play a role in chemical defense systems in plants by means of stored biologically active phytochemicals. These phytochemicals act as pest repellents. They show antimicrobial and insecticidal activities, and they have also been isolated and identified from wild plants., Results: The Asteraceae species Heterotheca subaxillaris has many glandular trichomes on the leaf surface, and these contain sesquiterpene carboxylates, which show insect antifeedant activity. Because these sesquiterpene carboxylates are major constituents of glandular trichomes, they may act as a chemical defense in H. subaxillaris. The Orobanchaceae species Parentucellia viscosa also has many glandular trichomes on the leaf surface and produces an insect antifeedant clerodane-type diterpene, kolavenic acid, in these trichomes. Additionally, two other Orobanchaceae species, Bellardia trixago and Parentucellia latifolia, also have many glandular trichomes, but the constituents of these glandular trichomes did not show biological activities against test insects. However, the seco-labdane diterpene alcohol trixagol and its hemi-malonate were major constituents in B. trixago, and these terpenes may act as physical defenses against herbivores by interfering with feeding due to their viscosity., Conclusion: The secondary metabolites from glandular trichomes of H. subaxillaris and P. viscosa showed insect antifeedant activity, and these secondary metabolites were presumed to act as chemical defenses for these plant species. On the other hand, non-biologically active secondary metabolites produced by two other Orobanchaceae, B. trixago and P. latifolia, were presumed to act as physical defenses due to their viscosity. Defense systems such as these may be applicable to new crop breeding to enhance protection against insect pests. © 2019 Society of Chemical Industry., (© 2019 Society of Chemical Industry.)

Plasma membrane (PM) lipid composition and domain organization are modulated by polarized exocytosis. Conversely, targeting of secretory vesicles at specific domains in the PM is carried out by exocyst complexes, which contain EXO70 subunits that play a significant role in the final recognition of the target membrane. As we have shown previously, a mature Arabidopsis trichome contains a basal domain with a thin cell wall and an apical domain with a thick secondary cell wall, which is developed in an EXO70H4-dependent manner. These domains are separated by a cell wall structure named the Ortmannian ring. Using phospholipid markers, we demonstrate that there are two distinct PM domains corresponding to these cell wall domains. The apical domain is enriched in phosphatidic acid (PA) and phosphatidylserine, with an undetectable amount of phosphatidylinositol 4,5-bisphosphate (PIP 2 ), whereas the basal domain is PIP 2 -rich. While the apical domain recruits EXO70H4, the basal domain recruits EXO70A1, which corresponds to the lipid-binding capacities of these two paralogs. Loss of EXO70H4 results in a loss of the Ortmannian ring border and decreased apical PA accumulation, which causes the PA and PIP 2 domains to merge together. Using transmission electron microscopy, we describe these accumulations as a unique anatomical feature of the apical cell wall-radially distributed rod-shaped membranous pockets, where both EXO70H4 and lipid markers are immobilized.

Plant growth promoting bacteria (PGPB) are agriculturally important soil bacteria that increase plant growth. We subjected peppermint to inoculation with three species of PGPB. After inoculation, the plants were sprayed with methyl jasmonate solution (MeJA) or SA (salicylic acid). Then, the plants were harvested and the plant growth parameters, trichome density, EO content and endogenous phytohormones were measured. Shoot fresh weight was reduced in plants inoculated and treated with MeJA whereas EO content varied depending on the MeJA concentration applied. Plants inoculated and treated with MeJA 2 mM showed the maximum increase in EO production, revealing a synergism between PGPB and MeJA. SA treatments also enhanced EO yield. The increased growth and EO production observed upon PGPB application were at least partly due to an increase in the JA and SA concentrations in the plant, as well as to an associated rise in the glandular trichome density., (Copyright © 2019 Elsevier Masson SAS. All rights reserved.)

Artemisia annua is the only natural source of the sesquiterpenoid artemisinin, which is widely used to treat malaria. The phytohormone jasmonic acid (JA) can significantly promote artemisinin biosynthesis in A. annua. AabHLH1 can bind and activate artemisinin biosynthetic genes, such as AaADS and AaCYP71AV1. In this study, we proved that AabHLH1 was responsive to MeJA treatment and highly expressed in glandular trichome-enriched tissues, and that its expression profile was similar to that of AaADS. Yeast two-hybrid assays showed that AabHLH1 interacted with all nine AaJAZ proteins in A. annua. Functional analysis with transgenic plants showed that several artemisinin biosynthetic genes were upregulated in AabHLH1-OE transgenic A. annua lines and downregulated in AabHLH1-EAR lines; furthermore, the artemisinin content was increased in the AabHLH1-OE lines and decreased in the AabHLH1-EAR lines. These results demonstrate that the JA-induced AabHLH1 positively regulates artemisinin biosynthesis by regulating the biosynthetic genes, and thus provide new insight into the regulatory mechanism of JA-induced artemisinin biosynthesis in A. annua., (© 2019 International Union of Biochemistry and Molecular Biology, Inc.)

Key Message: QTLs for insect resistance parameters, trichome type IV development, and more than 200 non-volatile metabolites, including 76 acyl sugars, all co-locate at the end of Chromosome 2 of Solanum galapagense. Host plant resistance is gaining importance as more and more insecticides are being banned due to environmental concerns. In tomato, resistance towards insects is found in wild relatives and has been attributed to the presence of glandular trichomes and their specific phytochemical composition. In this paper, we describe the results from a large-scale QTL mapping of data from whitefly resistance tests, trichome phenotyping and a comprehensive metabolomics analysis in a recombinant inbred line population derived from a cross between the cultivated Solanum lycopersicum and the wild relative S. galapagense, which is resistant to a range of pest insects. One major QTL (Wf-1) was found to govern the resistance against two different whitefly species. This QTL co-localizes with QTLs for the presence of trichomes type IV and V, as well as all 76 acyl sugars detected and about 150 other non-volatile phytochemicals, including methyl esters of the flavonols myricetin and quercetin. Based on these results, we hypothesize that Wf-1 is regulating the formation of glandular trichome type IV on the leaf epidermis, enabling the production and accumulation of bioactive metabolites in this type of trichomes.

Olive is one of the most important fruit crop trees in the history of Mediterranean because of the high quality oil. Olive oil has a well-balanced fatty acid composition along with biophenols, which make it exceptional in human diet and provide an exceptional value to the olive oil. Leaf non-glandular peltate trichomes are specialized cell types representing a protective barrier against acute environmental conditions. To characterize the proteome of this highly differentiated cell type, we performed a comparative proteomic analysis among isolated trichomes and trichome-less leaves. Proteins were separated and identified using the 2-DE MALDI-TOF/MS method. A number of enzymes involved in abiotic and biotic stress responses are present and may be responsible for the adaptation to prolonged adverse environmental conditions. The results show that this highly differentiated cell type is physiologically active fulfilling the demands of the trichomes in furnishing the leaf with a highly protective mechanism., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

Background: Cannabis possesses a rich spectrum of phytochemicals i.e. cannabinoids, terpenes and phenolic compounds of industrial and medicinal interests. Most of these high-value plant products are synthesised in the disk cells and stored in the secretory cavity in glandular trichomes. Conventional trichome analysis was so far based on optical microscopy, electron microscopy or extraction based methods that are either limited to spatial or chemical information. Here we combine both information to obtain the spatial distribution of distinct secondary metabolites on a single-trichome level by applying Coherent anti-Stokes Raman scattering (CARS), a microspectroscopic technique, to trichomes derived from sepals of a drug- and a fibre-type., Results: Hyperspectral CARS imaging in combination with a nonlinear unmixing method allows to identify and localise Δ 9 -tetrahydrocannabinolic acid (THCA) in the secretory cavity of drug-type trichomes and cannabidiolic acid (CBDA)/myrcene in the secretory cavity of fibre-type trichomes, thus enabling an easy discrimination between high-THCA and high-CBDA producers. A unique spectral fingerprint is found in the disk cells of drug-type trichomes, which is most similar to cannabigerolic acid (CBGA) and is not found in fibre-type trichomes. Furthermore, we differentiate between different cell types by a combination of CARS with simultaneously acquired two-photon fluorescence (TPF) of chlorophyll a from chloroplasts and organic fluorescence mainly arising from cell walls enabling 3D visualisation of the essential oil distribution and cellular structures., Conclusion: Here we demonstrate a label-free and non-destructive method to analyse the distribution of secondary metabolites and distinguish between different cell and chemo-types with high spatial resolution on a single trichome. The record of chemical fingerprints of single trichomes offers the possibility to optimise growth conditions as well as guarantee a direct process control for industrially cultivated medicinal Cannabis plants. Moreover, this method is not limited to Cannabis related issues but can be widely implemented for optimising and monitoring all kinds of natural or biotechnological production processes with simultaneous spatial and chemical information.

Variation in the induction of plant defenses along the plant canopy can determine distribution and colonization of arthropod herbivores within the plant. In tomato, type VI glandular trichomes, which are epidermal defensive structures, and their derived volatiles are induced by the phytohormone jasmonic acid (JA). How JA-mediated induction of these trichome-associated chemical defenses depends on the leaf developmental stage and correlates with resistance against herbivory is unknown. We showed that application of JA reduced thrips-associated damage, however the amplitude of this response was reduced in the fully developed leaves compared to those still developing. Although JA increased type-VI trichome densities in all leaf developmental stages, as well as JA-inducible defensive proteins, these increases were stronger in developing leaves. Remarkably, the concentration of trichome-derived volatiles was induced by JA to a larger degree in developing leaves than in fully developed leaves. In fully developed leaves, the increase in trichome-derived volatiles was explained by an enhanced production per trichome, while in developing leaves this was mainly caused by increases in type-VI trichome densities. Together, we showed that JA-mediated induction of trichome density and chemistry depends on leaf development stage, and it might explain the degree of thrips-associated leaf damage in tomato., (Copyright © 2018 The Authors. Published by Elsevier B.V. All rights reserved.)

Plants have developed a plethora of signals to interact with other organisms, finally building up a sophisticate language for communication. In this context, we investigated Salvia verticillata L. (Lamiaceae), with the primary goal to link secondary metabolites and actual biotic relationships. We specifically analysed the volatile organic compounds (VOC) spontaneously emitted by leaves and flowers and determined the composition of the essential oils obtained from the aerial parts across 2015 and 2016. We merged information of chemical analyses to a micromorphological investigation on the glandular indumentum and to focal observations on the pollinator assemblage. The VOC profiles were highly variable, with the floral bouquet being the most complex. Flowers and leaves showed 37 and 20 exclusive compounds, dominated by 1,8-cineole (10.4%) and germacrene D (38.4%), respectively. Sesquiterpene hydrocarbons prevailed (83.3% leaves; 73.7% flowers) and 19 common compounds were detected. The oil profiles proved to be consistent across the two years: sesquiterpene hydrocarbons invariably dominated, with germacrene D, bicyclogermacrene and β-caryophyllene as main compounds. The whole plant epidermis is thickly covered by two types of glandular hairs: peltates and small capitates, both responsible for the synthesis of terpenes, finally resulting in the VOC emission and in the essential oil production. S. verticillata attracted mainly bees belonging to two functional groups: medium-sized and large bees, notwithstanding the small size of its flowers. At the site, Apis mellifera and different Bombus species were recorded, mainly interested in feeding on nectar. The literature survey on the isolated volatile compounds confirmed the hypotheses on the seduction strategies towards Apoidea., (Copyright © 2018. Published by Elsevier Ltd.)

Subarctic vegetation is composed of mountain birch [Betula pubescens ssp. czerepanovii (MB)] forests with shrubs and other species growing in the understorey. The effects of the presence and density of one understorey shrub, Rhododendron tomentosum (RT), on the volatile emissions of MB, were investigated in a Finnish subarctic forest site in early and late growing season. Only MB trees with an RT-understorey emitted the RT-specific sesquiterpenoids, palustrol, ledol and aromadendrene. Myrcene, which is the most abundant RT-monoterpene was also emitted in higher quantities by MB trees with an RT-understorey. The effect of RT understorey density on the recovery of RT compounds from MB branches was evident only during the late season when sampling temperature, as well as RT emissions, were higher. MB sesquiterpene and total emission rates decreased from early season to late season, while monoterpene emission rate increased. Both RT and MB terpenoid emission rates were linked to density of foliar glandular trichomes, which deteriorated over the season on MB leaves and emerged with new leaves in the late season in RT. We show that sesquiterpene and monoterpene compounds emitted by understorey vegetation are adsorbed and re-released by MB, strongly affecting the MB volatile emission profile.

Naturally occurring and semisynthetic ergot alkaloids play a role in health care or as recreational drugs in Western and indigenous Mexican societies. Evidence is summarized that ergot alkaloids present in Central American Convolvulaceae like Turbina corymbosa, Ipomoea violacea , and Ipomoea asarifolia are colonized by different species of a newly described clavicipitaceous fungal genus named Periglandula . The fungi are associated with peltate glandular trichomes on the adaxial leaf surface of its host plants. The Periglandula fungi are not yet culturable in vitro but were demonstrated to have the capacity to synthesize ergot alkaloids. The alkaloids do not remain in the fungal mycelium but are translocated via the glandular trichomes into their plant host. Both fungi and host benefit from a symbiotic lifestyle. In evolutionary terms the alkaloid biosynthetic gene cluster in the Periglandula/Ipomoea symbiosis is likely to have a conserved (basic) structure while biosynthetic ergot gene clusters within the genera Claviceps and Epichloe were under ecological selection for alkaloid diversification., Competing Interests: The authors declare no conflict of interest., (Georg Thieme Verlag KG Stuttgart · New York.)

The volatile profiles (VOC) and the essential oil (EO) composition from the aerial parts of Salvia broussonetii were analysed. Sesquiterpene hydrocarbons dominate the VOCs from leaves (95.7%) and flowers (67.6%), followed by monoterpene hydrocarbons (2.6 and 29.7%, respectively). The main common compounds are germacrene D, β-bourbonene, α-pinene, α-copaene and α-gurjunene, even if with divergent relative abundances. In the leaf EOs the sesquiterpenes prevail, even if not overwhelmingly (about 50.0%), followed by monoterpenes (23.0-35.0%) and by minor fractions of diterpene hydrocarbons and non-terpene derivates. The most abundant common compounds across the two sampling periods are α-pinene, β-pinene, isobornyl acetate, α-gurjenene, germacrene D and bifloratriene. A morphological characterisation of the trichomes responsible for the productivity in terpenes was also performed. Four different morphotypes were observed on the above ground organs of S. brussonetii: peltates and capitates of type II and III resulted the only producers of volatile substances.

Schizonepeta tenuifolia Briquet, commonly known as Japanese catnip, is used for the treatment of colds, headaches, fevers, and skin rashes in traditional Asian medicine (China, Japan and Korea). The volatile oil and its constituents have various demonstrated biological activities, but there is currently limited information regarding the site of biosynthesis. Light microscopy and scanning electron microscopy indicated the presence of three distinct glandular trichome types which, based on their morphological features, are referred to as peltate, capitate and digitiform glandular trichomes. Laser scanning microscopy and 3D reconstruction demonstrated that terpenoid-producing peltate glandular trichomes contain a disk of twelve secretory cells. The oil of peltate glandular trichomes, collected by laser microdissection or using custom-made micropipettes, was demonstrated to contain (-)-pulegone, (+)-menthone and (+)-limonene as major constituents. Digitiform and capitate glandular trichomes did not contain appreciable levels of terpenoid volatiles. The yield of distilled oil from spikes was significantly (44%) higher than that from leaves, while the composition of oils was very similar. Oils collected directly from leaf peltate glandular trichomes over the course of a growing season contained primarily (-)-pulegone (>80% at 32 days after germination) in young plants, while (+)-menthone began to accumulate later (>75% at 80 days after germination), at the expense of (-)-pulegone (the levels of (+)-limonene remained fairly stable at 3-5%). The current study establishes the morphological and chemical characteristics of glandular trichome types of S. tenuifolia, and also provides the basis for unraveling the biosynthesis of essential oil in this popular medicinal plant., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The purpose of this investigation was to determine the influence of colored shade nets on the growth, anatomy and essential oil content, yield and chemical composition of Pogostemon cablin. The plants were cultivated under full sunlight, black, blue and red nets. The harvesting was performed 5 months after planting and it was followed by the analysis of plant growth parameters, leaf anatomy, essential oil content, yield and chemical composition. The plants grown under red net have produced more leaf, shoot, total dry weight and leaf area. Plants cultivated under colored nets showed differences in morphological features. Plants maintained under red net had a higher leaf blade thickness and polar and equatorial diameter of the stomata ratio. Additionally, higher yield of essential oil in the leaves was observed under red and blue colored shade net. The essential oil of the plants grown under red net showed the highest relative percentage of patchoulol (66.84%). Therefore, it is possible using colored shade nets to manipulate P. cablin growth, as well as its essential oil production with several chemical compositions. The analyses of principal components allowed observing that pogostol has negative correlation with α-guaiene and α-bulnesene. There was difference in total dry weight and patchoulol content when the patchouli is cultured under the red colored shade nets.

Background and Aims: Boraginales are often characterized by a dense cover of stiff, mineralized trichomes, which may act as a first line of defence against herbivores. Recent studies have demonstrated that the widely reported silica and calcium carbonate in plant trichomes may be replaced by calcium phosphate. The present study investigates mineralization patterns in 42 species from nine families of the order Boraginales to investigate detailed patterns of mineralization and the possible presence of a phylogenetic signal in different mineralization patterns., Methods: The distribution of biominerals was analysed by scanning electron microscopy (SEM) including cryo-SEM and energy-dispersive X-ray analyses with element mapping. The observed distribution of biominerals was plotted onto a published phylogeny of the Boraginales. Three colours were selected to represent the principal elements: Si (red), Ca (green) and P (blue)., Key Results: Calcium carbonate was present in the mineralized trichomes of all 42 species investigated, silica in 30 and calcium phosphate in 25; multiple mineralization with calcium carbonate and silica or calcium phosphate was found in all species, and 13 of the species were mineralized with all three biominerals. Trichome tips featured the most regular pattern - nearly all were exclusively mineralized with either silica or calcium phosphate. Biomineralization of the trichome shafts and bases was found to be more variable between species. However, the trichome bases were also frequently mineralized with calcium phosphate or silica, indicating that not only the tip is under functional constraints requiring specific patterns of chemical heterogeneity. The complete absence of either silica or phosphate may be an additional feature with systematic relevance., Conclusions: This study demonstrates that complex, site-specific and differential biomineralization is widespread across the order Boraginales. Calcium phosphate, only recently first reported as a structural plant biomineral, is common and appears to be functionally analogous to silica. A comparison with the phylogeny of Boraginales additionally reveals striking phylogenetic patterns. Most families show characteristic patterns of biomineralization, such as the virtual absence of calcium phosphate in Cordiaceae and Boraginaceae, the triple biomineralization of Heliotropiaceae and Ehretiaceae, or the absence of silica in Namaceae and Codonaceae. The complex chemical and phylogenetic patterns indicate that trichome evolution and functionalities are anything but simple and follow complex functional and phylogenetic constraints.

The distribution and ultrastructure of capitate glandular trichomes (GTs) in Flourensia species (Asteraceae) have been recently elucidated, but their metabolic activity and potential biological function remain unexplored. Selective nonvolatile metabolites from isolated GTs were strikingly similar to those found on leaf surfaces. The phytotoxic allelochemical sesquiterpene (-)-hamanasic acid A ((-)-HAA) was the major constituent (ca. 40%) in GTs. Although GTs are quaternary ammonium compounds (QACs)-accumulating species, glycine betaine was not found in GTs; it was only present in the leaf mesophyll. Two (-)-HAA accompanying surface secreted products: compounds 4-hydroxyacetophenone (piceol; 1) and 2-hydroxy-5-methoxyacetophenone (2), which were isolated and fully characterized (GC/MS, NMR), were present in the volatiles found in GTs. The essential oils of fresh leaves revealed ca. 33% monoterpenes, 26% hydrocarbon- and 30% oxygenated sesquiterpenes, most of them related to cadinene and bisabolene derivatives. Present results suggest a main role of GTs in determining the volatile and nonvolatile composition of F. campestris leaves. Based on the known activities of the compounds identified, it can be suggested that GTs in F. campestris would play key ecological functions in plant-pathogen and plant-plant interactions. In addition, the strikingly high contribution of compounds derived from cadinene and bisabolene pathways, highlights the potential of this species as a source of high-valued bioproducts., (© 2018 Wiley-VHCA AG, Zurich, Switzerland.)

Plant glandular trichomes are able to secrete and store large amounts of volatile organic compounds (VOCs). VOCs typically accumulate in dedicated extracellular spaces, which can be either subcuticular, as in the Lamiaceae or Asteraceae, or intercellular, as in the Solanaceae. Volatiles are retained at high concentrations in these storage cavities with limited release into the atmosphere and without re-entering the secretory cells, where they would be toxic. This implies the existence of mechanisms allowing transport of VOCs to the cavity but preventing their diffusion out once they have been delivered. The cuticle and cell wall lining the cavity are likely to have key roles in retaining volatiles, but their exact composition and the potential molecular players involved are largely unknown., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The commercially important essential oils of peppermint ( Mentha × piperita ) and its relatives in the mint family (Lamiaceae) are accumulated in specialized anatomical structures called glandular trichomes (GTs). A genome-scale stoichiometric model of secretory phase metabolism in peppermint GTs was constructed based on current biochemical and physiological knowledge. Fluxes through the network were predicted based on metabolomic and transcriptomic data. Using simulated reaction deletions, this model predicted that two processes, the regeneration of ATP and ferredoxin (in its reduced form), exert substantial control over flux toward monoterpenes. Follow-up biochemical assays with isolated GTs indicated that oxidative phosphorylation and ethanolic fermentation were active and that cooperation to provide ATP depended on the concentration of the carbon source. We also report that GTs with high flux toward monoterpenes express, at very high levels, genes coding for a unique pair of ferredoxin and ferredoxin-NADP + reductase isoforms. This study provides, to our knowledge, the first evidence of how bioenergetic processes determine flux through monoterpene biosynthesis in GTs., (© 2017 American Society of Plant Biologists. All Rights Reserved.)

At least three Leonurus species, Leonurus cardiaca, Leonurus japonicus , and Leonurus sibiricus , are used in various traditional medicinal systems for different therapeutic purposes. The plant names " L. sibiricus " and " L. japonicus " are often used as synonyms, which causes confusion in literature and implies that several studies may be based on improperly identified plant material. To avoid further confusion, the current status of the identification of these three species is illustrated. Characteristics for their unequivocal identification are presented using stereo and light microscopy as well as HPTLC.The establishment of the species-specific anatomical, morphological, and phytochemical characters was based on reference specimens in comparison with wild collected or commercially obtained material. Morphologically, the species differed in shape and length ratio of the upper and lower lip of the corolla. Differentiating anatomical characters involve the presence, density, and dimension of trichomes on different organs. Detailed anatomical descriptions of the leaves and corollae of L. japonicus and L. sibiricus are given and compared to L. cardiaca . These three sets of characters facilitate fast and reliable identification. Complementary HPTLC fingerprints show type-specific patterns that allow the differentiation of L. japonicus from the other two species. For the distinction of L. cardiaca and L. sibiricus , flower morphology has to be considered.The presented data contribute to the quality control of the three Leonurus species and additionally meet a definite and timely need regarding the introduction of L. japonicus to the European Pharmacopoeia in addition to the already existing monograph of L. cardiaca ., Competing Interests: Conflict of Interest: The authors declare no conflict of interest., (Georg Thieme Verlag KG Stuttgart · New York.)

Introduction: Glandular trichomes of plants are biochemical factories that could produce, store and secrete copious pharmaceutically important natural products. The Labiatae plant Leonurus japonicus is an important traditional Chinese medicine used to treat gynecological diseases, and has abundant peltate glandular trichomes (PGTs), in which the secondary metabolites accumulated are still unknown., Objective: To study the secondary metabolites specifically accumulated in the PGTs of L. japonicus and their biological activities, and provide a new way to pinpoint bioactive natural products from plants., Methodology: Morphology of the trichomes on L. japonicus were observed under a scanning electron microscope. The PGTs of L. japonicus were precisely collected using laser microdissection (LMD) and analysed for their secondary metabolites with ultra performance liquid chromatography-tandem mass spectrometry (UPLC-MS/MS). Targeted compounds were isolated with classical phytochemical methods, and their structures were elucidated by spectroscopic analysis. Biological activities were evaluated by in vitro assays., Results: Two labdane diterpenoids, leoheterin (1) and galeopsin (2), were localised in the PGTs of L. japonicus. Antithrombotic activity of 1 in anti-platelet aggregation assay induced by arachidonic acid was observed. Both compounds showed potential anti-inflammatory activity by inhibiting proinflammatory cytokine TNF-α. In addition, anti-proliferative effect of both compounds on several cancer cell lines was also detected., Conclusion: Two bioactive labdane diterpenoids were localised in the PGTs of L. japonicus. The findings suggested that it might be an efficient approach to explore bioactive natural products from the glandular trichomes of medicinal plants with LMD-UPLC/MS/MS. Copyright © 2017 John Wiley & Sons, Ltd., (Copyright © 2017 John Wiley & Sons, Ltd.)

Transmission electron microscopy was used to reveal a layer of multiply folded membranes that closely surrounded the tannin-accumulating vacuole in cells of honeysuckle petal trichomes. A huge amount of tannins were deposited in the peripheral region and the center of the vacuole. The prolific membranes extended to the tannins deposited along the vacuole periphery. It was difficult to distinguish the vacuole membrane, and it seemed as if it was the layer of multiply folded membranes that separated the vacuole lumen from the cytoplasm. In addition, there were also membrane assemblies in the cytoplasm away from the vacuole, which were continuous with the proliferated membranes bordering the vacuole. Therefore, the tannin-accumulating vacuole was in close association with a very large network of proliferated membranes. The occurrence of such a layer of multiply folded membranes around the tannin-accumulating vacuole might be a structural strategy for improvement of the efficiency of vacuolar accumulation of tannins., (Copyright © 2017 Elsevier Ltd. All rights reserved.)