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Cannabis sativa (C. sativa L.) has garnered significant attention worldwide due to its widespread use as a pharmaceutical agent. With the increasing clinical application of C. sativa and cannabinoid therapeutics, there is strong interest in the development of superior plant varieties and optimisation of growth conditions to enhance secondary metabolite yield. Our RNA sequencing analysis revealed differential expression of hormone-related transcripts in developing C. sativa trichomes, suggesting the involvement of hormone signalling pathways in cannabinoid production. Leveraging the potency of exogenous hormones on plants, this study sought to determine if the application of cytokinin (CK), gibberellic acid (GA) and jasmonic acid (JA) modified trichome morphology and the cannabinoid profile over an 8-week period following the induction of flowering. Exogenous hormone application led to alterations in trichome morphology, with each treatment significantly reducing trichome head width by the final week of assessment. Interestingly, GA application also resulted in a significant reduction in the concentration of Δ-9-tetrahydrocannabinol (THC), Δ-9-tetrahydrocannabinolic acid (THCA), cannabidiol (CBD) and cannabidiolic acid (CBDA) by week 8 post floral induction, however, JA and CK treatment did not consistently modulate the accumulation of these cannabinoids. The minor cannabinoids, cannabidivaranic acid (CBDVA), cannabicyclolic acid (CBLA), cannabicyclol (CBL), cannabichromene (CBC), cannabigerolic acid (CBGA) and cannabigerol (CBG), were also affected by hormone treatments, with varying degrees of accumulation observed. These findings underscore the intricate interplay between phytohormones and secondary metabolite biosynthesis in C. sativa. Our study highlights the potential of hormone modulation as a strategy to enhance cannabinoid yield and offers some insights into the regulatory mechanisms governing cannabinoid biosynthesis in C. sativa trichomes. [ABSTRACT FROM AUTHOR]

Abstract Background The glandular trichomes of tobacco (Nicotiana tabacum) can efficiently produce secondary metabolites. They act as natural bioreactors, and their natural products function to protect plants against insect-pests and pathogens and are also components of industrial chemicals. To clarify the molecular mechanisms of tobacco glandular trichome development and secondary metabolic regulation, glandular trichomes and glandless trichomes, as well as other different developmental tissues, were used for RNA sequencing and analysis. Results By comparing glandless and glandular trichomes with other tissues, we obtained differentially expressed genes. They were obviously enriched in KEGG pathways, such as cutin, suberine, and wax biosynthesis, flavonoid and isoflavonoid biosynthesis, terpenoid biosynthesis, and plant–pathogen interaction. In particular, the expression levels of genes related to the terpenoid, flavonoid, and wax biosynthesis pathway mainly showed down-regulation in glandless trichomes, implying that they lack the capability to synthesize certain exudate compounds. Among the differentially expressed genes, 234 transcription factors were found, including AP2-ERFs, MYBs, bHLHs, WRKYs, Homeoboxes (HD-ZIP), and C2H2-ZFs. These transcription factor and genes that highly expressed in trichomes or specially expressed in GT or GLT. Following the overexpression of R2R3-MYB transcription factor Nitab4.5_0011760g0030.1 in tobacco, an increase in the number of branched glandular trichomes was observed. Conclusions Our data provide comprehensive gene expression information at the transcriptional level and an understanding of the regulatory pathways involved in glandular trichome development and secondary metabolism.

This study investigates the micromorphology and morphometrics of aggregate fruit (ripe hypanthium and achene) of eight wild Rosa species collected throughout Serbia. Field research was conducted to find roses at different habitats, collecting 1,836 aggregate fruits with 25,999 achenes from 137 shrubs. Multicellular glandular trichomes are found on the hypanthium surface, while long and short stalk capitate glandular trichomes were observed on the fruits of R. gallica, R. pendulina, R. tomentosa and R. villosa. Numerous unicellular non-glandular trichomes were found inside the urceolate hypanthium and on the surface of most studied achenes, except for R. spinosissima species. Micromorphological analysis revealed three types of exocarp ornamentation: scalariform (R. dumetorum, R. tomentosa and R. micrantha), reticulate (R. gallica, R. arvensis, R. spinosissima and R. villosa) and reticulate-smooth (R. pendulina). Morphometric analysis indicated significant differences among the 14 morphological characters. Achene length was the most significant morphometric feature for identifying Rosa species (P˂0.05). The results contribute to our poor knowledge of rose fruit morphology and provide some well-defined microcharacters that could be used as additional features for species delimitation. [ABSTRACT FROM AUTHOR]

Cotton crops are routinely threatened by emerging fungal diseases. Fungal endophytes also can be considered latent phytopathogens. In this study we tested the hypothesis that an endophytic strain of Diaporthe, isolated from chlorotic leaves of cotton (Gossypium hirsutum), could trigger physiological effects of biotic stress in this oilseed plant. We also assessed the histopathological aspects of the mycelial interaction of the endophyte with the adaxial surface of G. hirsutum leaves. Thus, we studied the synthesis of photosynthetic pigments, pattern of gas exchange, and photochemistry of cotton plants subjected to inoculation with Diaporthe ueckerae via root and leaf at three different phenological stages (vegetative, reproductive, and maturation). Additionally, we histopathologically analyzed infected leaves using electron microscopy to study the process of leaf colonization by this endophytic fungus. We evidenced that D. ueckerae inoculation negatively affected the synthesis of photosynthetic pigments in plants at vegetative and reproductive stages. Moreover, inoculation also negatively affected the photosynthetic rate and carboxylation efficiency of these plants. We also found that the presence of the endophyte increased transpiration and decreased water use efficiency in the plants. Furthermore, foliar inoculation negatively affected stomatal conductance, whereas inoculation via leaf or root reduced the photochemical performance of cotton. We also observed that D. ueckerae colonizes the leaf tissues of G. hirsutum via glandular trichomes and forces penetration into the epidermis using appressoria, and the plant responds by closing the stomata. The observed physiological alterations are indicative of biotic stress, confirming the hypothesis that D. ueckerae may be an opportunistic phytopathogen for cotton plants. [ABSTRACT FROM AUTHOR]

Chemical fertilizers used in plant development and differentiation have become a global problem affecting the entire ecosystem, especially soil pollution. Food production demand with the increasing population has encouraged scientists to use biogenic nanoparticles in the agricultural field. Evaluation of growth, development, and differentiation processes of sweet basil (Ocimum basilicum L.) seedlings at gradually increasing concentrations of biogenic iron oxide nanoparticles (BIO-NPs) were identified by morphological and physiological parameters in this study. The results showed that growth parameters reached the maximum value at 100 mg/L but were less at other concentrations. At similar concentration, the stomatal density of the leaf was the maximum, while the stomatal area showed the lowest value. The levels of H2O2 and malondialdehyde (MDA) decreased in the treated seedlings. BIO-NPs increased the antioxidant defense and supported its growth by changing the antioxidant enzyme activities, H2O2, and MDA contents. The BIO-NP treatment provided positive improvements in phytochemical content in parallel with the growth and development of sweet basil seedlings. Different growth parameters, physiological results, supporting enzyme activities, and biochemical data revealed the contribution of the BIO-NP treatments to the growth and development of sweet basil seedlings. BIO-NPs improved higher phytochemical production of sweet basil, which may be suitable for its propagation on a commercial scale. [ABSTRACT FROM AUTHOR]

Background: The glandular trichomes of tobacco (Nicotiana tabacum) can efficiently produce secondary metabolites. They act as natural bioreactors, and their natural products function to protect plants against insect-pests and pathogens and are also components of industrial chemicals. To clarify the molecular mechanisms of tobacco glandular trichome development and secondary metabolic regulation, glandular trichomes and glandless trichomes, as well as other different developmental tissues, were used for RNA sequencing and analysis. Results: By comparing glandless and glandular trichomes with other tissues, we obtained differentially expressed genes. They were obviously enriched in KEGG pathways, such as cutin, suberine, and wax biosynthesis, flavonoid and isoflavonoid biosynthesis, terpenoid biosynthesis, and plant–pathogen interaction. In particular, the expression levels of genes related to the terpenoid, flavonoid, and wax biosynthesis pathway mainly showed down-regulation in glandless trichomes, implying that they lack the capability to synthesize certain exudate compounds. Among the differentially expressed genes, 234 transcription factors were found, including AP2-ERFs, MYBs, bHLHs, WRKYs, Homeoboxes (HD-ZIP), and C2H2-ZFs. These transcription factor and genes that highly expressed in trichomes or specially expressed in GT or GLT. Following the overexpression of R2R3-MYB transcription factor Nitab4.5_0011760g0030.1 in tobacco, an increase in the number of branched glandular trichomes was observed. Conclusions: Our data provide comprehensive gene expression information at the transcriptional level and an understanding of the regulatory pathways involved in glandular trichome development and secondary metabolism. [ABSTRACT FROM AUTHOR]

Rocky outcrop environments at high altitudes have nutrient‐poor soil, where species are exposed to water scarcity and high solar radiation. Baccharis platypoda DC. occurs in such an environment and has a rigid and transparent secretion that covers the entire inflorescence. We analysed and compared the secretory structures and their chemical composition in female and male inflorescences of B. platypoda, a dioecious species, to explore chemodiversity within this species and assess potential differences between individuals. Our investigation also aims to understand the occurrence of these substances in the genus Baccharis L. Chemical compounds and secretory structures were similar in female and male inflorescences. There are glandular trichomes on the epidermis of the abaxial surface of bracts, and secretory ducts in the axis of the inflorescence, as well as in sepals, petals, and bracts. Histochemical tests were positive for phenolic compounds, flavonoids, proteins, pectin, and lipids, but not for mucilage. Flavonoid content varied between 6.24% and 9.81%, being higher in female inflorescences. Chromatography revealed the presence of several phenolic compounds, some terpenes, and other less frequent classes in both female and male inflorescences. We highlight that trichomes found on these surfaces produce abundant phenolic compounds. These act as natural defence agents, absorbing UV radiation and minimizing oxidative stress to plant cells. The chemical composition of the secretion covering the inflorescences may reflect adaptation and survival mechanisms of these organisms under extreme sun exposure. [ABSTRACT FROM AUTHOR]

Trichomes are well-known efficient plant defense mechanisms to limit arthropod herbivory, especially in Solanaceae. The present study aims to evaluate the impact of trichome types on the development, survival and dispersal of Tetranychus urticae, and the phytoseiid predatory mite Typhlodromus (Anthoseius) recki. Six Solanum lycopersicum cultivars and two wild Solanum species, S. cheesmaniae and S. peruvianum, presenting contrasting densities and types of trichomes, were considered. Cultivars and species were characterized by counting each trichome type on leaves, petioles and stems. Mites stuck on petiole and stem and alive mites on the leaflet used for mite release and in the whole plant were counted three weeks after T. urticae plant infestation. Tetranychus urticae settlement and dispersal were differently affected by trichomes. Trichome types V and VI did not affect settlement and dispersal, whereas trichome types I and IV on the petiole had the highest impacton mites. Trichomes on leaves slightly affected mite establishment, there appears to be a repellent effect of trichome types I and IV. The low densities of both T. urticae and its predator detected for the cv. Lancaster could not be clearly associated to the trichome types here considered. The predator did not seem to be affected by plant characteristics, but rather by T. urticae numbers on the plant. The trichome traits unfavorable to T. urticae, did not affect the predator which showed high efficiency to control this pest on all the plant genotypes considered, but at a favorable predator:prey ratio (1:1). Altogether, these results are encouraging for the use of T. (A.) recki as a biological control agent of T. urticae regardless of the trichome structure of the tomato cultivars, but other conditions should be tested to conclude on practical implementations. [ABSTRACT FROM AUTHOR]

Chemical fertilizers used in plant development and differentiation have become a global problem affecting the entire ecosystem, especially soil pollution. Food production demand with the increasing population has encouraged scientists to use biogenic nanoparticles in the agricultural field. Evaluation of growth, development, and differentiation processes of sweet basil (Ocimum basilicum L.) seedlings at gradually increasing concentrations of biogenic iron oxide nanoparticles (BIO-NPs) were identified by morphological and physiological parameters in this study. The results showed that growth parameters reached the maximum value at 100 mg/L but were less at other concentrations. At similar concentration, the stomatal density of the leaf was the maximum, while the stomatal area showed the lowest value. The levels of H2O2 and malondialdehyde (MDA) decreased in the treated seedlings. BIO-NPs increased the antioxidant defense and supported its growth by changing the antioxidant enzyme activities, H2O2, and MDA contents. The BIO-NP treatment provided positive improvements in phytochemical content in parallel with the growth and development of sweet basil seedlings. Different growth parameters, physiological results, supporting enzyme activities, and biochemical data revealed the contribution of the BIO-NP treatments to the growth and development of sweet basil seedlings. BIO-NPs improved higher phytochemical production of sweet basil, which may be suitable for its propagation on a commercial scale.

Abstract Artemisinin is primarily synthesized and stored in the subepidermal space of the glandular trichomes of Artemisia annua. The augmentation of trichome density has been demonstrated to enhance artemisinin yield. However, existing literature lacks insights into the correlation between the stratum corneum and trichomes. This study aims to unravel the involvement of TrichomeLess Regulator 3 (TLR3), which encodes the transcription factor, in artemisinin biosynthesis and its potential association with the stratum corneum. TLR3 was identified as a candidate gene through transcriptome analysis. The role of TLR3 in trichome development and morphology was investigated using yeast two-hybrid, pull-down analysis, and RNA electrophoresis mobility assay. Our research revealed that TLR3 negatively regulates trichome development. It modulates the morphology of Arabidopsis thaliana trichomes by inhibiting branching and inducing the formation of abnormal trichomes in Artemisia annua. Overexpression of the TLR3 gene disrupts the arrangement of the stratum corneum and reduces artemisinin content. Simultaneously, TLR3 possesses the capacity to regulate stratum corneum development and trichome follicle morphology by interacting with TRICHOME AND ARTEMISININ REGULATOR 1, and CycTL. Consequently, our findings underscore the pivotal role of TLR3 in the development of glandular trichomes and stratum corneum biosynthesis, thereby influencing the morphology of Artemisia annua trichomes.

SUMMARY: Cannabis glandular trichomes (GTs) are economically and biotechnologically important structures that have a remarkable morphology and capacity to produce, store, and secrete diverse classes of secondary metabolites. However, our understanding of the developmental changes and the underlying molecular processes involved in cannabis GT development is limited. In this study, we developed Cannabis Glandular Trichome Detection Model (CGTDM), a deep learning‐based model capable of differentiating and quantifying three types of cannabis GTs with a high degree of efficiency and accuracy. By profiling at eight different time points, we captured dynamic changes in gene expression, phenotypes, and metabolic processes associated with GT development. By integrating weighted gene co‐expression network analysis with CGTDM measurements, we established correlations between phenotypic variations in GT traits and the global transcriptome profiles across the developmental gradient. Notably, we identified a module containing methyl jasmonate (MeJA)‐responsive genes that significantly correlated with stalked GT density and cannabinoid content during development, suggesting the existence of a MeJA‐mediated GT formation pathway. Our findings were further supported by the successful promotion of GT development in cannabis through exogenous MeJA treatment. Importantly, we have identified CsMYC4 as a key transcription factor that positively regulates GT formation via MeJA signaling in cannabis. These findings provide novel tools for GT detection and counting, as well as valuable information for understanding the molecular regulatory mechanism of GT formation, which has the potential to facilitate the molecular breeding, targeted engineering, informed harvest timing, and manipulation of cannabinoid production. [ABSTRACT FROM AUTHOR]

Artemisinin is primarily synthesized and stored in the subepidermal space of the glandular trichomes of Artemisia annua. The augmentation of trichome density has been demonstrated to enhance artemisinin yield. However, existing literature lacks insights into the correlation between the stratum corneum and trichomes. This study aims to unravel the involvement of TrichomeLess Regulator 3 (TLR3), which encodes the transcription factor, in artemisinin biosynthesis and its potential association with the stratum corneum. TLR3 was identified as a candidate gene through transcriptome analysis. The role of TLR3 in trichome development and morphology was investigated using yeast two-hybrid, pull-down analysis, and RNA electrophoresis mobility assay. Our research revealed that TLR3 negatively regulates trichome development. It modulates the morphology of Arabidopsis thaliana trichomes by inhibiting branching and inducing the formation of abnormal trichomes in Artemisia annua. Overexpression of the TLR3 gene disrupts the arrangement of the stratum corneum and reduces artemisinin content. Simultaneously, TLR3 possesses the capacity to regulate stratum corneum development and trichome follicle morphology by interacting with TRICHOME AND ARTEMISININ REGULATOR 1, and CycTL. Consequently, our findings underscore the pivotal role of TLR3 in the development of glandular trichomes and stratum corneum biosynthesis, thereby influencing the morphology of Artemisia annua trichomes. [ABSTRACT FROM AUTHOR]

Artemisia arborescens is a Mediterranean evergreen shrub, with silver grey-green tomentose leaves and a strong scent. It has various ethnopharmacological uses and its secondary metabolites have demonstrated antimicrobial, antiviral, pharmaceutical, phytotoxic, and insecticidal activities. Different extracts obtained from aerial parts of this species are known for their allelopathic effect, but similar studies on its essential oil (EO) are lacking. Therefore, we carried out a pharmacognostic study, obtaining the characterization of the secretory structures and the EO produced. Trans-thujone and camphor are the main components, followed by aromadendrene, camphene, and 8-cedren-13-ol. EO phytotoxic activity was tested on weed plants (Lolium multiflorum Lam. and Sinapis arvensis L.) and crops (Raphanus sativus L. and Cucumis sativus L.), showing inhibition on both germination and radical growth of the two weeds tested. The effects of the EO against the bacterial plant pathogens Xanthomonas campestris pv. campestris (Gram−) and Pseudomonas syringae pv. tomato (Gram+) was also assayed. The minimum inhibitory concentration (MIC) was observed when it was used undiluted [100% v/v], and growth inhibition when diluted at different doses. The antimicrobial activity was also confirmed by the cellular material release and biofilm formation assays. The overall data show that A. arborescens EO can find application as a potential alternative biocontrol product against weeds and plant pathogens. This goal is particularly important from the perspective of replacing synthetic pesticides with natural products, which safeguard both the environment and the health of consumers. [ABSTRACT FROM AUTHOR]

Trichome is a hair-like structure involved in mechanical and chemical defenses of plants against herbivorous insects. Nicotiana benthamiana, a wild tobacco plant, has well-developed glandular trichomes that secrete sugar esters with potent repellent and insecticidal properties. However, there is a lack of information about the effectiveness of trichome extract in the control of plant-sapping insects. The objective of this study was to investigate the effects of N. benthamiana trichome exudates on Bemisia tabaci MED (Gennadius) (Hemiptera: Aleyrodidae), a highly destructive insect pest that poses significant threats to both vegetable and ornamental plants globally. First, we determined the host preference and mortality of B. tabaci adults using the choice test and feeding assay towards tomato and N. benthamiana plants. B. tabaci preferred tomato over N. benthamiana plants. Second, we extracted N. benthamiana trichome exudates by washing the leaves with either water or ethanol and evaluated their oral toxicities against B. tabaci adults using a parafilm feeding chamber containing 20% sucrose solution. Oral ingestion of both extracts significantly increased mortality in a concentration-dependent manner. Oral ingestion of ethanol-washed 10% trichome extract caused >60% mortality in B. tabaci adults after 36 h. Third, trichome exudates were concentrated by drying to obtain a powder form, which was more potent in killing whiteflies than the liquid form. Oral ingestion of 1% trichome powder was completely lethal to B. tabaci within 36 h. N. benthamiana trichome exudates are highly toxic to B. tabaci through oral ingestion, suggesting that N. benthamiana can be used as a potential natural pesticide for whitefly management.

Cannabis sativa (C. sativa L.) has garnered significant attention worldwide due to its widespread use as a pharmaceutical agent. With the increasing clinical application of C. sativa and cannabinoid therapeutics, there is strong interest in the development of superior plant varieties and optimisation of growth conditions to enhance secondary metabolite yield. Our RNA sequencing analysis revealed differential expression of hormone-related transcripts in developing C. sativa trichomes, suggesting the involvement of hormone signalling pathways in cannabinoid production. Leveraging the potency of exogenous hormones on plants, this study sought to determine if the application of cytokinin (CK), gibberellic acid (GA) and jasmonic acid (JA) modified trichome morphology and the cannabinoid profile over an 8-week period following the induction of flowering. Exogenous hormone application led to alterations in trichome morphology, with each treatment significantly reducing trichome head width by the final week of assessment. Interestingly, GA application also resulted in a significant reduction in the concentration of Δ-9-tetrahydrocannabinol (THC), Δ-9-tetrahydrocannabinolic acid (THCA), cannabidiol (CBD) and cannabidiolic acid (CBDA) by week 8 post floral induction, however, JA and CK treatment did not consistently modulate the accumulation of these cannabinoids. The minor cannabinoids, cannabidivaranic acid (CBDVA), cannabicyclolic acid (CBLA), cannabicyclol (CBL), cannabichromene (CBC), cannabigerolic acid (CBGA) and cannabigerol (CBG), were also affected by hormone treatments, with varying degrees of accumulation observed. These findings underscore the intricate interplay between phytohormones and secondary metabolite biosynthesis in C. sativa. Our study highlights the potential of hormone modulation as a strategy to enhance cannabinoid yield and offers some insights into the regulatory mechanisms governing cannabinoid biosynthesis in C. sativa trichomes.

Premise: Glandular trichomes are implicated in direct and indirect defense of plants. However, the degree to which glandular and non‐glandular trichomes have evolved as a consequence of herbivory remains unclear, because their heritability, their association with herbivore resistance, their trade‐offs with one another, and their association with other functions are rarely quantified. Methods: We conducted a phylogenetic comparison of trichomes and herbivore resistance against the generalist caterpillar, Heliothis virescens, among tarweed species (Asteraceae: Madiinae) and a genetic correlation study comparing those same traits among maternal half‐sibs of three tarweed species. Results: Within a tarweed species, we found no evidence that herbivore growth rate decreased on tarweed individuals or maternal sib groups with more glandularity or denser trichomes. However, tarweed species with more glandularity and fewer non‐glandular trichomes resulted in slower‐growing herbivores. Likewise, a trade‐off between glandular and non‐glandular trichomes was apparent among tarweed species, but not among individuals or sib groups within a species. Conclusions: Our results suggest that this key herbivore does not select for trichomes as a direct defense in tarweed species. However, trichomes differed substantially among species and likely affect herbivore pressure on those species. Our results demonstrate that trade‐offs among plant traits, as well as inference on the function of those traits, can depend on scale. [ABSTRACT FROM AUTHOR]

Fungal diseases constitute a major challenge in the cultivation of the world's most important textile plant, cotton. Opportunistic fungi attack plants and promote physiological alterations and tissue damage, thereby causing losses in productivity. Herein, the hypothesis that Gossypium hirsutum L. plants are physiologically affected, depending on the phenological stage and site of infection (root system or leaf), when affected by the opportunistic phytopathogen Fusarium equiseti was tested. To this end, the photosynthetic pigment production, gas exchange, chlorophyll a fluorescence, and histopathological mechanisms involved in the plant-phytopathogen interaction were evaluated. This study aimed to diagnose the most aggressive route of infection by comparing infection via root with infection via the leaf, and to identify the developmental stage with the highest susceptibility by comparing different phenological stages. Cotton plants show metabolic losses at the maturation stage, resulting in low photosynthetic pigment concentrations, low photosynthetic rates, and photochemical stress. However, when infected by F. equiseti in the root region, the photosynthesis and photochemistry of these plants are more affected than plants inoculated in the leaf region. Low net photosynthesis (A) and water use efficiency (WUE) rates, but high light absorption flux (ABS/RC) and energy dissipation flux (DI0/RC) values were observed. Regardless of the phenological stage, the presence of F. equiseti in the roots of cotton plants caused high transpiratory rates in the plants, suggesting non-stomatic water loss and photochemical damage. In contrast, non-inoculated plants produced more photosynthetic pigments and showed higher WUE and photochemical efficiency, as indicated by the high PIABS values. Leaf histopathology assessment revealed more fragile regions of the epidermis, such as the contact region between the guard cells and adjacent epidermal cells, as well as the basal region of glandular trichomes as important access routes for F. equiseti to the inner leaf layers at all developmental stages. This study confirmed that the root colonization of cotton plants with F. equiseti can more intensely impact the physiological quality of the plants, regardless of the developmental stage. [ABSTRACT FROM AUTHOR]

Main conclusion: In flowers multiple secretory systems cooperate to deliver specialized metabolites to support specific roles in defence and pollination. The collective roles of cell types, enzymes, and transporters are discussed. The interplay between reproductive strategies and defense mechanisms in flowering plants has long been recognized, with trade-offs between investment in defense and reproduction predicted. Glandular trichomes and secretory cavities or ducts, which are epidermal and internal structures, play a pivotal role in the secretion, accumulation, and transport of specialized secondary metabolites, and contribute significantly to defense and pollination. Recent investigations have revealed an intricate connection between these two structures, whereby specialized volatile and non-volatile metabolites are exchanged, collectively shaping their respective ecological functions. However, a comprehensive understanding of this profound integration remains largely elusive. In this review, we explore the secretory systems and associated secondary metabolism primarily in Asteraceous species to propose potential shared mechanisms facilitating the directional translocation of these metabolites to diverse destinations. We summarize recent advances in our understanding of the cooperativity between epidermal and internal secretory structures in the biosynthesis, secretion, accumulation, and emission of terpenes, providing specific well-documented examples from pyrethrum (Tanacetum cinerariifolium). Pyrethrum is renowned for its natural pyrethrin insecticides, which accumulate in the flower head, and more recently, for emitting an aphid alarm pheromone. These examples highlight the diverse specializations of secondary metabolism in pyrethrum and raise intriguing questions regarding the regulation of production and translocation of these compounds within and between its various epidermal and internal secretory systems, spanning multiple tissues, to serve distinct ecological purposes. By discussing the cooperative nature of secretory structures in flowering plants, this review sheds light on the intricate mechanisms underlying the ecological roles of terpenes in defense and pollination. [ABSTRACT FROM AUTHOR]

In this work, we studied Tanacetum vulgare, Tanacetum parthenium, and Tanacetum corymbosum (Asteraceae) cultivated at the Ghirardi Botanic Garden (Toscolano Maderno, Brescia, Northern Italy) of the University of Milan. An integrative research approach was adopted: microscopic and histochemical, with special focus on the secretory structures responsible for the productivity of secondary metabolites; phytochemical, with the analysis of the essential oil (EO) profiles from the air-dried, flowered aerial parts collected in June 2021; bio-ecological, with emphasis, based on literature data, on the ecology and biological activity of the main EO components. In all three species, two basic trichome morphotypes (flagellar non-glandular and biseriate glandular) occurred with different distribution patterns. The glandular ones produced terpenes, along with flavonoids. A high level of chemical variability in the EO compositions emerged, specifically for qualitative data. T. vulgare profile was more complex and heterogeneous than those obtained from T. parthenium and T. corymbosum, with camphor as the predominant compound, followed by farnesol and α-santalone, respectively. Finally, the obtained scientific findings were made available to the visitors of the botanic garden through new dissemination labeling that highlights the "invisible", microscopic features of the plants, from an Open Science perspective ("Botanic Garden, factories of molecules...work in progress"—Lombardy Region Project Lr. 25/2016, year 2021). [ABSTRACT FROM AUTHOR]

To study the genes regulating the synthesis of sugar esters in tobacco, transcriptome sequencing analysis was performed on tobacco glandular hair cells and bundle sheath cells, and RT-qPCR was used for verification. The results showed that 11 962 differentially expressed genes were obtained, of which 5145 were up-regulated and 6817 were down-regulated. GO annotation of differentially expressed genes showed that the up-regulated genes mainly included biological processes (2265), molecular functions (1169), and cellular components (363), with a total of 3793 GO terms. Based on the assignment of KEGG pathway, the sequence was mapped to 122 metabolic pathways. Glycolysis/gluconeogenesis, starch and sucrose metabolism, phenylpropanoid biosynthesis related to glycoester synthesis were highly enriched, and acyl sugar pathway in glandular hairs was significantly up-regulated. By comparing with the tobacco reference genome, 12 up-regulated acyltransferase genes were identified in glandular hair cells. RT-qPCR results showed that the expression of 11 genes was higher than that of the control. The expression of gene 63826, gene 16194 and gene 37511 increased by 195.79 times, 166.23 times and 132.65 times, respectively. The above 11 genes may be involved in regulating the synthesis of sugar esters in glandular hair cells, providing a basis for further verification of gene function. [ABSTRACT FROM AUTHOR]

Achillea millefolium L. herb and flowers have high biological activity; hence, they are used in medicine and cosmetics. The aim of this study was to perform morpho-anatomical analyses of the raw material, including secretory tissues, histochemical assays of the location of lipophilic compounds, and quantitative and qualitative analysis of essential oil (EO). Light and scanning electron microscopy techniques were used to analyse plant structures. The qualitative analyses of EO were carried out using gas chromatography-mass spectrometry (GC/MS). The results of this study showed the presence of exogenous secretory structures in the raw material, i.e., conical cells (papillae) on the adaxial surface of petal teeth and biseriate glandular trichomes on the surface flowers, bracts, stems, and leaves. Canal-shaped endogenous secretory tissue was observed in the stems and leaves. The histochemical assays revealed the presence of total, acidic, and neutral lipids as well as EO in the glandular trichome cells. Additionally, papillae located at the petal teeth contained neutral lipids. Sesquiterpenes were detected in the glandular trichomes and petal epidermis cells. The secretory canals in the stems were found to contain total and neutral lipids. The phytochemical assays demonstrated that the A. millefolium subsp. millefolium flowers contained over 2.5-fold higher amounts of EO (6.1 mL/kg) than the herb (2.4 mL/kg). The EO extracted from the flowers and herb had a similar dominant compounds: β-pinene, bornyl acetate, (E)-nerolidol, 1,8-cineole, borneol, sabinene, camphor, and α-pinene. Both EO samples had greater amounts of monoterpenes than sesquiterpenes. Higher amounts of oxygenated monoterpenes and oxygenated sesquiterpenoids were detected in the EO from the herb than from the flowers. [ABSTRACT FROM AUTHOR]

This study investigates the impact of priming on the resistance of tomato plants to the potato aphid Macrosiphum euphorbiae, focusing on the role of glandular trichomes. Glandular trichomes are specific hairs that provide protection to tomato plants against herbivorous insects. The experimental priming conducted in this study revealed that prior infestation by Spodoptera littoralis caterpillars increased the plant's resistance against M. euphorbiae, pointing at the jasmonic acid (JA) signaling pathway in regulating this plant-aphid interaction. Glandular trichomes type IV were effective against aphids regardless of the previous infestation. Using JA-deficient tomato (spr2), we observed that M. euphorbiae multiplication increased, while the number of aphids on salicylic -deficient NahG plants was lower than in the wildtype Moneymaker. These findings emphasize the crucial role of the JA signaling pathway in tomato plant resistance to aphids and the importance of glandular trichomes to enhance plant defences against pests. [ABSTRACT FROM AUTHOR]

Trichome is a hair-like structure involved in mechanical and chemical defenses of plants against herbivorous insects. Nicotiana benthamiana, a wild tobacco plant, has well-developed glandular trichomes that secrete sugar esters with potent repellent and insecticidal properties. However, there is a lack of information about the effectiveness of trichome extract in the control of plant-sapping insects. The objective of this study was to investigate the effects of N. benthamiana trichome exudates on Bemisia tabaci MED (Gennadius) (Hemiptera: Aleyrodidae), a highly destructive insect pest that poses significant threats to both vegetable and ornamental plants globally. First, we determined the host preference and mortality of B. tabaci adults using the choice test and feeding assay towards tomato and N. benthamiana plants. B. tabaci preferred tomato over N. benthamiana plants. Second, we extracted N. benthamiana trichome exudates by washing the leaves with either water or ethanol and evaluated their oral toxicities against B. tabaci adults using a parafilm feeding chamber containing 20% sucrose solution. Oral ingestion of both extracts significantly increased mortality in a concentration-dependent manner. Oral ingestion of ethanol-washed 10% trichome extract caused >60% mortality in B. tabaci adults after 36 h. Third, trichome exudates were concentrated by drying to obtain a powder form, which was more potent in killing whiteflies than the liquid form. Oral ingestion of 1% trichome powder was completely lethal to B. tabaci within 36 h. N. benthamiana trichome exudates are highly toxic to B. tabaci through oral ingestion, suggesting that N. benthamiana can be used as a potential natural pesticide for whitefly management. [ABSTRACT FROM AUTHOR]

ABSTRACTThis study investigates the impact of priming on the resistance of tomato plants to the potato aphid Macrosiphum euphorbiae, focusing on the role of glandular trichomes. Glandular trichomes are specific hairs that provide protection to tomato plants against herbivorous insects. The experimental priming conducted in this study revealed that prior infestation by Spodoptera littoralis caterpillars increased the plant's resistance against M. euphorbiae, pointing at the jasmonic acid (JA) signaling pathway in regulating this plant-aphid interaction. Glandular trichomes type IV were effective against aphids regardless of the previous infestation. Using JA-deficient tomato (spr2), we observed that M. euphorbiae multiplication increased, while the number of aphids on salicylic -deficient NahG plants was lower than in the wildtype Moneymaker. These findings emphasize the crucial role of the JA signaling pathway in tomato plant resistance to aphids and the importance of glandular trichomes to enhance plant defences against pests.

The recent introduction of Bemisia tabaci (Gennadius) MED (Hemiptera: Aleyrodidae) in the main tomato-producing areas in Brazil is concerning due to its high infestation capacity and to the numerous lethal viruses that this insect can transmit to plants. In addition, B. tabaci MED has a history of lower susceptibility to several groups of insecticides. Thus, this study evaluated the preference of B. tabaci MED among 39 tomato genotypes, belonging to seven Solanum species, as well as the physical and morphological factors related to the preference. The tomato genotypes LA 716, IAC 294, PI 127,826, PI 134,418, PI 134,417, Tainara, PI 126,925 and LA 116 expressed resistance to B. tabaci MED. The density of glandular trichomes was negatively correlated with the number of adults, B. tabaci MED eggs/cm² and nymphs/cm². On the other hand, the density of non-glandular trichomes and the slope of the trichomes stimulated the colonization by adults of B. tabaci MED. Substrate color was not correlated with genotype preference. The resistant genotypes and their characteristics can be explored in genetic breeding programs with the aim of developing cultivars resistant to whiteflies. [ABSTRACT FROM AUTHOR]

Plant defense traits such as trichomes along with biocontrol agents may provide alternatives to insecticide use in tomatoes, Solanum lycopersicum L. (Solanaceae). However, plant‐herbivore‐natural enemy interactions are not always complementary. In a series of greenhouse and field experiments, we explored whether augmented defense traits (i.e., production of acylsugars) in tomato plants could reduce sweet potato whitefly, Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae), populations and aid the establishment of the predatory mite Amblyseius swirskii Athias‐Henriot (Acari: Phytoseiidae). In the field experiment, commercial tomato cultivars and acylsugar‐producing tomato lines received no predatory mites or mites released via three methods: dusting on top, dusting on bottom, or slow‐release sachets. In the first greenhouse experiment, predatory mites were released onto the commercial and acylsugar‐producing tomato plants via sachets. In the second greenhouse experiment using a similar design, we augmented the mite diet with an alternative non‐prey resource (i.e., cattail pollen). Our results indicated that acylsugar‐producing tomato plants supported significantly fewer whiteflies than the commercial lines in all experiments. However, in the field, despite lower whitefly numbers, Tomato yellow leaf curl virus (TYLCV) (Geminiviridae, Begomovirus) was detected at higher frequencies in acylsugar‐producing lines. Few mites were recovered from all commercial and acylsugar‐producing lines in the field or greenhouse experiments suggesting A. swirskii does not establish well on tomatoes, and acylsugar lines successfully decreased whitefly populations but not a viral disease transmitted even at low whitefly abundance. [ABSTRACT FROM AUTHOR]

SUMMARY: Several commercially important secondary metabolites are produced and accumulated in high amounts by glandular trichomes, giving the prospect of using them as metabolic cell factories. Due to extremely high metabolic fluxes through glandular trichomes, previous research focused on how such flows are achieved. The question regarding their bioenergetics became even more interesting with the discovery of photosynthetic activity in some glandular trichomes. Despite recent advances, how primary metabolism contributes to the high metabolic fluxes in glandular trichomes is still not fully elucidated. Using computational methods and available multi‐omics data, we first developed a quantitative framework to investigate the possible role of photosynthetic energy supply in terpenoid production and next tested experimentally the simulation‐driven hypothesis. With this work, we provide the first reconstruction of specialised metabolism in Type‐VI photosynthetic glandular trichomes of Solanum lycopersicum. Our model predicted that increasing light intensities results in a shift of carbon partitioning from catabolic to anabolic reactions driven by the energy availability of the cell. Moreover, we show the benefit of shifting between isoprenoid pathways under different light regimes, leading to a production of different classes of terpenes. Our computational predictions were confirmed in vivo, demonstrating a significant increase in production of monoterpenoids while the sesquiterpenes remained unchanged under higher light intensities. The outcomes of this research provide quantitative measures to assess the beneficial role of chloroplast in glandular trichomes for enhanced production of secondary metabolites and can guide the design of new experiments that aim at modulating terpenoid production. Significance Statement: Our interdisciplinary work provides a bioenergetic explanation for the added benefit of having photosynthetically active glandular trichomes. [ABSTRACT FROM AUTHOR]

Main conclusion: Morphological, phytochemical, and transcriptome analyses revealed candidate genes involved in the biosynthesis of volatile monoterpenes and development of glandular trichomes in Monarda citriodora. Monarda citriodora Cerv. ex Lag. is a valuable aromatic plant due to the presence of monoterpenes as major constituents in its essential oil (EO). Thus, it is of sheer importance to gain knowledge about the site of the biosynthesis of these terpenoid compounds in M. citriodora, as well as the genes involved in their biosynthesis. In this study, we studied different types of trichomes and their relative densities in three different developmental stages of leaves, early stage of leaf development (L1), mid-stage of leaf development (L2), and later stage of leaf development (L3) and the histochemistry of trichomes for the presence of lipid and terpenoid compounds. Further, the phytochemical analysis of this plant through GC–MS indicated a higher content of monoterpenes (thymol, thymoquinone, γ-terpinene, p-cymene, and carvacrol) in the L1 stage with a substantial decrease in the L3 stage of leaf development. This considerable decrease in the content of monoterpenes was attributed to the decrease in the trichome density from L1 to L3. Further, we developed a de novo transcriptome assembly by carrying out RNA sequencing of different plant parts of M. citriodora. The transcriptome data revealed several putative unigenes involved in the biosynthesis of specialized terpenoid compounds, as well as regulatory genes involved in glandular trichome development. The data generated in the present study build a strong foundation for further improvement of M. citriodora, in terms of quantity and quality of its essential oil, through genetic engineering. [ABSTRACT FROM AUTHOR]

Micro-morphological characteristics of leaves such as stomata and trichomes are reliable indicators of plant response to environmental conditions. The aim of this study was to determine the adaptation strategies of 10 woody species growing in semi-arid Zagros forests in western Iran based on leaf microstructures, focusing on trichomes and stomata using light and scanning electron microscopy. Different types of trichomes and stomata were recognized. Stomata were generally more visible on the abaxial sides of leaves, a feature that helps reduce water loss from leaf surfaces exposed to direct airflow and radiation. Trichomes were classified into two different types: glandular trichomes and non-glandular trichomes with subgroups such as solitary, two-armed T-shaped unicellular, crypt, hooked hairs with cystoliths, fasciculate, stellate, long coiled, tufted stellate, long multiradiate, short bristles with cystoliths, peltate scales, and dendritic hairs. The abundance of trichomes on the leaves indicates their role as biological control parameters. Eucalyptus camaldulensis was the only introduced species studied that lacked trichomes on both sides of the leaf. The most frequently observed types of stomata were "Anomocytic" and "Paracytic". The striking differences between the micro-morphological characteristics of the leaves of different species are an expression of the plants adaptations to the environmental ecological conditions.

Abstract Background Glandular capitate trichomes which form on bract tissues of female inflorescences of high THC-containing Cannabis sativa L. plants are important sources of terpenes and cannabinoids. The influence of plant age and cannabis genotype on capitate trichome development, morphology, and maturation has not been extensively studied. Knowledge of the various developmental changes that occur in trichomes over time and the influence of genotype and plant age on distribution, numbers, and morphological features should lead to a better understanding of cannabis quality and consistency. Methods Bract tissues of two genotypes—“Moby Dick” and “Space Queen”—were examined from 3 weeks to 8 weeks of flower development using light and scanning electron microscopy. Numbers of capitate trichomes on upper and lower bract surfaces were recorded at different positions within the inflorescence. Observations on distribution, extent of stalk formation, glandular head diameter, production of resin, and extent of dehiscence and senescence were made at various time points. The effects of post-harvesting handling and drying on trichome morphology were examined in an additional five genotypes. Results Two glandular trichome types—bulbous and capitate (sessile or stalked)—were observed. Capitate trichome numbers and stalk length were significantly (P = 0.05) greater in “Space Queen” compared to “Moby Dick” at 3 and 6 weeks of flower development. Significantly more stalked-capitate trichomes were present on lower compared to upper bract surfaces at 6 weeks in both genotypes, while sessile-capitate trichomes predominated at 3 weeks. Epidermal and hypodermal cells elongated to different extents during stalk formation, producing significant variation in length (from 20 to 1100 μm). Glandular heads ranged from 40 to 110 μm in diameter. Maturation of stalked-capitate glandular heads was accompanied by a brown color development, reduced UV autofluorescence, and head senescence and dehiscence. Secreted resinous material from glandular heads appeared as droplets on the cuticular surface that caused many heads to stick together or collapse. Trichome morphology was affected by the drying process. Conclusion Capitate trichome numbers, development, and degree of maturation were influenced by cannabis genotype and plant age. The observations of trichome development indicate that asynchronous formation leads to different stages of trichome maturity on bracts. Trichome stalk lengths also varied between the two genotypes selected for study as well as over time. The variability in developmental stage and maturation between genotypes can potentially lead to variation in total cannabinoid levels in final product. Post-harvest handling and drying were shown to affect trichome morphology.

Artemisia arborescens is a Mediterranean evergreen shrub, with silver grey-green tomentose leaves and a strong scent. It has various ethnopharmacological uses and its secondary metabolites have demonstrated antimicrobial, antiviral, pharmaceutical, phytotoxic, and insecticidal activities. Different extracts obtained from aerial parts of this species are known for their allelopathic effect, but similar studies on its essential oil (EO) are lacking. Therefore, we carried out a pharmacognostic study, obtaining the characterization of the secretory structures and the EO produced. Trans-thujone and camphor are the main components, followed by aromadendrene, camphene, and 8-cedren-13-ol. EO phytotoxic activity was tested on weed plants (Lolium multiflorum Lam. and Sinapis arvensis L.) and crops (Raphanus sativus L. and Cucumis sativus L.), showing inhibition on both germination and radical growth of the two weeds tested. The effects of the EO against the bacterial plant pathogens Xanthomonas campestris pv. campestris (Gram−) and Pseudomonas syringae pv. tomato (Gram+) was also assayed. The minimum inhibitory concentration (MIC) was observed when it was used undiluted [100% v/v], and growth inhibition when diluted at different doses. The antimicrobial activity was also confirmed by the cellular material release and biofilm formation assays. The overall data show that A. arborescens EO can find application as a potential alternative biocontrol product against weeds and plant pathogens. This goal is particularly important from the perspective of replacing synthetic pesticides with natural products, which safeguard both the environment and the health of consumers.

Artemisia annua is a medicinal plant particularly known for the production of a sesquiterpene lactone artemisinin; a specialty metabolite known for its efficacy in the treatment of malaria by killing different strains of Plasmodium falciparum due to radicals released upon the cleavage of its endoperoxide motif. Considering these facts and the immense medicinal value of artemisinin, the enhancement of in planta production of artemisinin is highly desirable. As strigolactones are known to regulate various aspects of plant growth and development, the effects of foliar spray of different concentrations of synthetic strigolactone analog GR24 (0, 0.5, 1, 2, 4, and 8 µM) on A. annua were studied. As compared to the control group, the foliar application of GR24 had a positive impact on general growth, photosynthesis, and other physiological indices with 4 µM GR24 showing the best results. The results indicate that GR24 application increased the plant biomass and various attributes related to photosynthesis, like total chlorophyll content, chlorophyll fluorescence, stomatal conductance, internal CO2, and net photosynthetic rate. Moreover, the activity of various enzymes related to photosynthesis like carbonic anhydrase, nitrate reductase, and RuBisCO was escalated. The GR24 also improved certain attributes related to glandular trichomes, with a significant enhancement in content and yield of artemisinin as compared to untreated plants. [ABSTRACT FROM AUTHOR]

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. LAY DESCRIPTION: For millennia humans have grown and consumed cannabis flowers for use in traditional medicine and intoxication, due to the plant's active ingredients called cannabinoids. The most well‐known cannabinoid is tetrahydrocannabidiol (THC). Despite this long history, and recent relaxation of cannabis uses for medicinal and recreational consumption in many parts of the world, little is known about how the plant makes the cannabinoids in its cells. Small mushroom‐shaped structures on the flower surface, called glandular trichomes, produce and store the cannabinoids. To image inside the cells of THC‐rich cannabis flowers, it is necessary to use electron microscopy, and this has revealed complex internal membrane networks and specialised cell structures in these tiny biofactories. In this study, we explored different ways to preserve the cannabis glandular trichomes for electron microscopy, using either cryogenic preservation or chemical crosslinking, in order to determine the benefits and drawbacks of each method. The chemical crosslinking method produced dramatic re‐arrangement of cell contents that were not seen in the cryofixation, where the near‐instantaneous immobilisation of the cell structure led to exceptional preservation of the THC‐producing cells. Despite the benefits of using cryofixation for preserving the complex cell structures, it was not as useful for seeing details of the cell walls surrounding the glandular trichome. Together, our data show stark differences in cannabis glandular trichome cell structure that result from cryofixation and chemical fixation. This is important because the chemical fixation protocol was historically used and informed previous models of cannabinoid production and trafficking. Cryofixation produces exciting new models of THC production in cannabis, due to its high‐fidelity preservation of cell structures. In contrast, studies of cell walls may still benefit from the traditional chemical cross‐linking technique of sample preservation for electron microscopy. https://www.cell.com/current‐biology/fulltext/S0960‐9822(22)01115‐0 [ABSTRACT FROM AUTHOR]

Negative frequency-dependent selection (NFDS) has been shown to maintain polymorphism in a diverse array of traits. The action of NFDS has been confirmed through modeling, experimental approaches, and genetic analyses. In this study, we investigated NFDS in the wild using morph-frequency changes spanning a 20-year period from over 30 dimorphic populations of Datura wrightii. In these populations, plants either possess glandular (sticky) or non-glandular (velvety) trichomes, and the ratio of these morphs varies substantially among populations. Our method provided evidence that NFDS, rather than drift or migration, is the primary force maintaining this dimorphism. Most populations that were initially dimorphic remained dimorphic, and the overall mean and variance in morph frequency did not change over time. Furthermore, morph-frequency differences were not related to geographic distances. Together, these results indicate that neither directional selection, drift, or migration played a substantial role in determining morph frequencies. However, as predicted by negative frequency-dependent selection, we found that the rare morph tended to increase in frequency, leading to a negative relationship between the change in the frequency of the sticky morph and its initial frequency. In addition, we found that morph-frequency change over time was significantly correlated with the damage inflicted by two herbivores: Lema daturaphila and Tupiochoris notatus. The latter is a specialist on the sticky morph and damage by this herbivore was greatest when the sticky morph was common. The reverse was true for L. daturaphila, such that damage increased with the frequency of the velvety morph. These findings suggest that these herbivores contribute to balancing selection on the observed trichome dimorphism.

Glandular trichomes (GTs) are one of the epidermal tissues of medicinal plants which function in the synthesis, storage, and secretion of secondary metabolites. The active ingredients of Chinese medicinal materials are mostly secondary metabolites of plants. Accordingly, it is of great research value to explore the quality of medicinal materials using the GTs of medicinal plants as the starting point. However, most of the current studies on GTs of medicinal plants are still at the simple morphological identification stage, and there are few studies on the compounds secreted by GTs and secondary metabolic processes. Here, we reviewed the literature, summarized the morphological types of medicinal plant GTs, separation and purification technology, analysis technology, and biological activities of secondary metabolites, and established a research approach to medicinal plant GTs. We hope to provide a reference for future research on GT inclusions and secondary metabolism.

Monoterpenoids are C10 isoprenoid natural products with diverse functions as volatile and non-volatile secondary metabolites. Their biosynthesis from acyclic diphosphate precursors is often accompanied by one or more ring closures, which stabilize the linear, carbocationic intermediates generated during typical terpene synthase reaction sequences. The subset that remains uncyclized following the elimination of the diphosphate group correspondingly retains higher reactivity than their cyclized counterparts, a property that may explain the additional transformations that geraniol and other acyclics undergo to yield monoterpene aldehydes, acids, esters, glycosides, and iridoids, many of which possess allelochemical activity as pollinator attractants, feeding deterrents, or insect repellents. Essential oils of lemon grass (or citronella) (Cymbopogon), rose-scented geraniums (Pelargonium), and lemon-scented gums (Corymbia) are rich sources of the acyclic monoterpene alcohols we collectively refer to in this review as 'citronelloids'. Their formation from geranyl diphosphate may be brought about by the tps-g or tps-b subclade of terpene synthases or, alternatively, by a subclade of Nudix hydrolases which act preferentially on prenyl diphosphate substrates, yielding monoterpene alcohols in a two step process that involves a monophosphate intermediate. In some members of the Lamiid and Campanulid clades, further oxidation of geraniol and a non-classical, reductive cyclization leads to the iridoids, a group of mostly non-volatile, bicyclic monoterpenes with potent anti-herbivory and anti-inflammatory properties. Here we review the recent literature on acyclic monoterpene biosynthesis, their natural distribution across the plant kingdom, and their emerging roles in medicine as biologically active natural products. [ABSTRACT FROM AUTHOR]

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−1DW. 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. [ABSTRACT FROM AUTHOR]

This study presents the first comparative analysis of the leaf secretory structures across Asteraceae. In this work, the leaf secretory structures of more than 500 species of 35 of the 40 tribes and 11 of the 13 subfamilies of Asteraceae are described and compared to evaluate their diversity at the tribe level and to identify evolutionary patterns. Leaf secretory structures are present in 28 of the 35 analyzed tribes and correspond to canals (recorded in 17 tribes), secretory cavities (1 tribe), hydathodes (19 tribes), laticifers (4 tribes) and glandular trichomes (24 tribes). Canals are mostly associated with vascular bundles and predominate in Asteroideae, while cavities were only present within Tageteae. Hydathodes occur in leaves without divisions and with well-developed teeth. Laticifers were observed only in the tribes of Cichorioideae. Seven glandular trichome morphotypes were differentiated by their cellular composition and shape. These observations together with the available information showed that secretory structures are found in 80% of the Asteraceae tribes. Four of the 40 tribes did not present any type of secretory structure. Our study reveals that almost all of the tribes possess one to three types of secretory structures, and are absent in some early-diverging clades. Character evolution analyses show that glandular trichomes are plesiomorphic in Asteraceae. This study found that secretory structures prevail in late-diverging lineages and were taxonomically informative at different levels. Our comparative study of the secretory structures in Asteraceae is essential for the standardization of its terminology and will provide a frame of reference for future studies. [ABSTRACT FROM AUTHOR]

In this work, we studied Tanacetum vulgare, Tanacetum parthenium, and Tanacetum corymbosum (Asteraceae) cultivated at the Ghirardi Botanic Garden (Toscolano Maderno, Brescia, Northern Italy) of the University of Milan. An integrative research approach was adopted: microscopic and histochemical, with special focus on the secretory structures responsible for the productivity of secondary metabolites; phytochemical, with the analysis of the essential oil (EO) profiles from the air-dried, flowered aerial parts collected in June 2021; bio-ecological, with emphasis, based on literature data, on the ecology and biological activity of the main EO components. In all three species, two basic trichome morphotypes (flagellar non-glandular and biseriate glandular) occurred with different distribution patterns. The glandular ones produced terpenes, along with flavonoids. A high level of chemical variability in the EO compositions emerged, specifically for qualitative data. T. vulgare profile was more complex and heterogeneous than those obtained from T. parthenium and T. corymbosum, with camphor as the predominant compound, followed by farnesol and α-santalone, respectively. Finally, the obtained scientific findings were made available to the visitors of the botanic garden through new dissemination labeling that highlights the “invisible”, microscopic features of the plants, from an Open Science perspective (“Botanic Garden, factories of molecules…work in progress”—Lombardy Region Project Lr. 25/2016, year 2021).

Hot air impingement drying (HAID) was employed to dry Amomum villosum (A. v.) fruit, one of the most popular traditional Chinese herbs and functional foods, under different drying temperatures, air velocities and the drying characteristics and quality were evaluated. The drying time decreased from 12 to 4 h, 10 to 8 h with the increase of temperature and velocity from 50 to 70 °C, 3 to 9 m/s, respectively. Weibull model precisely described the drying behavior. The highest total flavonoid content was obtained at 60 °C with low shell burst ratio. As for essential oil components, the relative content of monoterpenoids increased while sesquiterpenes decreased with increasing temperature. The acids and alcohols contents decreased with increase in velocity. By considering both drying time and quality, 60 °C and 6 m/s was proposed as the most favorable condition. The findings indicate HAID is a promising drying method for A. v. fruit. [ABSTRACT FROM AUTHOR]

Glandular trichomes (GTs) are one of the epidermal tissues of medicinal plants which function in the synthesis, storage, and secretion of secondary metabolites. The active ingredients of Chinese medicinal materials are mostly secondary metabolites of plants. Accordingly, it is of great research value to explore the quality of medicinal materials using the GTs of medicinal plants as the starting point. However, most of the current studies on GTs of medicinal plants are still at the simple morphological identification stage, and there are few studies on the compounds secreted by GTs and secondary metabolic processes. Here, we reviewed the literature, summarized the morphological types of medicinal plant GTs, separation and purification technology, analysis technology, and biological activities of secondary metabolites, and established a research approach to medicinal plant GTs. We hope to provide a reference for future research on GT inclusions and secondary metabolism. [ABSTRACT FROM AUTHOR]

Arnica montana L. is a very important medicinal plant and simultaneously a European endemic endangered plant species. The morphological features and details of seed development and achene variability are poorly recognized. The aim of this study was to determine the impact of the achene position in the infructescence and the location of the inflorescence on the plant on the (i) morphological characteristics and germination ability of achenes, and (ii) recruitment of seedlings and their biometric features. Infructescences containing fully ripe achenes were randomly collected from A. montana individuals for the measurements and the germination experiment. Scanning electron microscopy, fluorescence microscopy, and light microscopy were used for characterization of flowers and achenes. The morphological traits of achenes and reproductive characteristics of A. montana were determined by the position of the achenes in the infructescence and the location of the inflorescence on the plant. The surface of arnica achenes is equipped with non-glandular and glandular trichomes, which is very rarely presented in species of the family Asteraceae. It is possible that the fluid-containing glandular trichomes are a source of essential oils. The peripherally located achenes were longer, thinner, and lighter. They were characterized by lower embryo weight, lower embryo/achene weight ratio, and lower germination capacity in comparison to the centrally located ones. The results presented in this article fill the gap in the knowledge of the morphology of achenes and the biology of the species, and provide information that can help in breeding programs, active protection, and field cultivation. [ABSTRACT FROM AUTHOR]