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Premise Floral spurs are key innovations associated with elaborate pollination mechanisms that have evolved independently several times across angiosperms. Spur formation can shift the floral symmetry from radial to bilateral, as it is the case in Tropaeolum, the only member of the Brassicales with floral nectar spurs. The genetic mechanisms underlying both spur and bilateral symmetry in the family have not yet been investigated. Methods We studied flower development and morphoanatomy of Tropaeolum longifolium. We also generated a reference transcriptome and isolated all candidate genes involved in adaxial-abaxial differential growth during spur formation. Finally, we evaluated the evolution of the targeted genes across Brassicales and examined their expression in dissected floral parts. Results Five sepals initiate spirally, followed by five petals alternate to the sepals, five antesepalous stamens, three antepetalous stamens, and three carpels. Intercalary growth at the common base of sepals and petals forms a floral tube. The spur is an outgrowth from the adaxial region of the tube, lined up with the medial sepal. We identified Tropaeolum specific duplications in the TCP3/4L and STM gene lineages, which are critical for spur formation in other taxa. In addition, we found that TM6 (MADS-box), RL2 (RAD-like7), and KN2/6L2 and OSH6L (KNOX1 genes), have been lost in core Brassicales but retained in Tropaeolum. Conclusions Three genes are pivotal during the extreme adaxial-abaxial asymmetry of the floral tube, namely, TlTCP4L2 restricted to the adaxial side where the spur is formed, and TlTCP12 and TlSTM1 to the abaxial side, lacking a spur.

espanolEste trabajo optimizo la temperatura y el tiempo de pasteurizacion de los tuberculos de mashua (Tropaeolum tuberosumRuiz y Pavon) morada en funcion de su capacidad antioxidante, fenolicos totales, flavonoides totales, contenido de antocianinas y aceptabilidad. Para esto, se pesaron, lavaron, pelaron, cortaron y colocaron en un extractor de jugo los tuberculos mashua morados de la provincia de Acobamba-Huancavelica-Peru. La bebida de los tuberculos se preparo usando el jugo de los tuberculos,el tratamiento termico de las muestras se realizo a temperaturas entre 75 y 85 °C y tiempo de 10 a 25 min. El diseno se planifico utilizando un diseno central compuesto rotacionaly se realizo un analisis de superficie de respuesta en las muestras obtenidas. El contenido de fenoles totales, antocianinas, flavonoides totales y capacidad antioxidante fue influenciado por el tiempo y latemperatura de la pasteurizacion. A temperaturas mas altas y tiempos de pasteurizacion hubo una reduccion en el contenido de fenoles totales, antocianinas, flavonoides totales y capacidad antioxidante. La mayor aceptabilidad general de los panelistas fue a altas temperaturas y tiempos de pasteurizacion. La temperatura de 77 °C y tiempo de 13 minutos mostraron unvalor maximo de las variables de respuesta con baja degradacion de los compuestos antioxidantes y buena aceptabilidad de las bebidas de extracto de mashua. EnglishThis work optimized the temperature and time of pasteurization of purple mashua (Tropaeolum tuberosum Ruiz and Pavon) tubers drink based on its antioxidant capacity, total phenolics, total flavonoids, anthocyanins content, and acceptability. For this, purple mashua tubers from the province of Acobamba-Huancavelica-Peru were weighed, washed, peeled, cut and placed in a juice extractor. The beverage of tubers was prepared using the juice of tubers, the thermal treatment of the samples was carried out at temperatures between 75 to 85 °C and time 10 to 25 min. The design was planned using a Rotational Composite Central Design and a response surface analysis was performed on the samples obtained. The content of total phenolics, anthocyanins, total flavonoids, and antioxidant capacity was influenced by the time and temperature of pasteurization. At higher temperatures and pasteurization times there was a reduction in the content of total phenolics, anthocyanins, total flavonoids, and antioxidant capacity. The greatest general acceptability of the panelists was at high pasteurization temperatures and times. The temperature of 77 °C and time of 13 minutes showed the maximum value of response variables with low degradation of the antioxidant compounds and good acceptability of mashua extract drinks.

Tropaeolum tuberosum, commonly known as Mashua, is an herbal remedy used in traditional Andean medicine for the relief of kidney and bladder pain, as well as contusions. This study aimed to evaluate the fractions and isolated compounds from T. tuberosum with analgesic activity mediated by the transient receptor potential vanilloid-1 receptor. A bioguided phytochemical analysis based on NMR/MS was performed to identify the compounds of the n-heptane fractions from samples of purple tubers of T. tuberosum. The transient receptor potential vanilloid-1 agonist and antagonist activity were assessed through the measurement of intracellular Ca2+ in HEK001 cells. The chemical structure determination led to the identification of two alkamides: N-(2-hydroxyethyl)-7Z,10Z,13Z,16Z-docosatetraenamide (1) and N-oleoyldopamine (2). Both compounds induced increased intracellular calcium flow with IC50 values of 3.2 nM and 7.9 nM, respectively, thus activating the transient receptor potential vanilloid-1 receptor. Our research is the first report to show that these two compounds isolated from T. tuberosum can act as agonists of the transient receptor potential vanilloid-1 receptor, providing scientific evidence for the traditional use of this species in pain relief.

Complete plastome sequence of Tropaeolum pentaphyllum revealed molecular markers, hotspots of nucleotide polymorphism, RNA editing sites and phylogenetic aspects Tropaeolaceae Juss. ex DC. comprises approximately 95 species across North and South Americas. Tropaeolum pentaphyllum Lam. is an unconventional and endangered species with occurrence in some countries of South America. Although this species presents nutritional, medicinal and ornamental uses, genetic studies involving natural populations or promising genotypes are practically non-existent. Here, we report the nucleotide sequence of T. pentaphyllum plastome. It represents the first complete plastome sequence of the family Tropaeolaceae to be fully sequenced and analyzed in detail. The sequencing data revealed that the T. pentaphyllum plastome is highly similar to the plastomes of other Brassicales. Notwithstanding, our analyses detected some specific features concerning events of IR expansion and structural changes in some genes such as matK, rpoA, and rpoC2. We also detected 251 SSR loci, nine hotspots of nucleotide polymorphism, and two specific RNA editing sites in the plastome of T. pentaphyllum. Moreover, plastid phylogenomic inference indicated a closed relationship between the families Tropaeolaceae and Akaniaceae, which formed a sister group to Moringaceae-Caricaceae. Finally, our data bring new molecular markers and evolutionary features to be applied in the natural population, germplasm collection, and genotype selection aiming conservation, genetic diversity evaluation, and exploitation of this endangered species.

Eco-friendly biosynthesis of nanoparticles from medicinal plants as reducing agent has gained importance due to its potential therapeutic uses. In the present study Silver nanoparticles (AgNPs) were eco-friendly synthesized using the leaf extracts of the medicinal plant Tropaeolum majus. The obtained AgNPs were characterized by UV - visible spectrum, FTIR, SEM and XRD which clearly showed the reduction of Ag+ ions to Ag0. In addition, the aqueous and ethanolic extracts were analyzed for phytochemicals and its antioxidant activities. GC–MS spectrum showed the presence of 25 compounds with benzeneacetic acid as the dominant contents. The synthesized AgNPs revealed maximum absorption spectrum at 463 nm and FTIR vibrational peaks at 3357.46, 21,966.52, 2118.42, 1637.27, 658.571 and 411.728 cm−1 respectively. SEM and XRD studies evidenced the nature of nanocrystalline with face centered cubic (fcc) crystal structure. Both AgNPs and plant extracts showed more inhibition activity against Pseudomonas aeroginosa compared to other bacteria with MIC value of 6.25 μg/ml. Antifungal activities was higher for Penicilium notatum with MIC value 31.2 μg/ml. The IC50 values for MCF7 for aqueous extract were found to be 4.68 μg/ml, ethanol extract 7.5 μg/ml, AgNPs 2.49 μg/ml, and doxorubicin 1.4 μg/ml. The IC50 values for VERO cell line for aqueous extract was 8.1 μg/ml, ethanol extract with 6.8 μg/ml, silver nanoparticles 5.3 μg/ml and doxorubicin 2.6 μg/ml respectively. Conclusively, the antibacterial, antifungal, antioxidant and anticancer properties of the synthesized AgNPs from Tropaeolum majus act as major therapeutic drug for microbial infectious disease and other health associated disorders.

This research was funded by the CTS-107 and AGR145 Groups from the Granada University. M. Fuel obtained all plants of the study from Ecuador. A part of this study has been funded by the 2018 Research Initiation Grants Program for Master Students of the Vice-Rectorate for Research and Knowledge Transfer of the University of Granada., Moringa oleifera, Tropaeolum tuberosum and Annona cherimola are medicinal plants traditionally used in Ecuador. However, their therapeutic properties are not completely known. We analyzed chromatographically ethanolic extracts of the seeds of M. oleifera, A. cherimola and the tubers of T. tuberosum; all presented a high content of polyphenols. The extract of A. cherimola showed the highest antioxidant activity and M. oleifera had the highest capacity to enhance the activity of detoxifying enzymes such as glutathione S-transferase and quinone oxidoreductase. The antitumor effect of these extracts was evaluated in vitro with colorectal cancer (CRC) cell lines T84, HCT-15, SW480 and HT-29, as well as with cancer stem cells (CSCs). A. cherimola and M. oleifera extracts presented the lowest IC50 in T-84 and HCT-15 (resistant) cells, respectively, as well as the highest level of inhibition of proliferation in multicellular tumor spheroids of HCT-15 cells. The inhibitory effect on CSCs is noteworthy because in vivo, these cells are often responsible for cancer recurrences and resistance to chemotherapy. Moreover, all extracts showed a synergistic activity with 5-Fu. The antiproliferative mechanism of the extracts was related to overexpression of caspases 9, 8 and 3 and increased production of reactive oxygen species. In addition, we observed cell death by autophagy in M. oleifera and T. tuberosum extracts. Therefore, these ethanolic extracts are excellent candidates for future molecular analysis of the presence of bioactive compounds and in vivo studies which could improve colon cancer therapy., Granada Universit CTS-107 AGR145, 2018 Research Initiation Grants Program for Master Students of the Vice-Rectorate for Research and Knowledge Transfer of the University of Granada

Two alkaloids were isolated and identified for the first time in the black tubers of Tropaeolum tuberosum, collected from the Titicani-Taca, Ingavi province in La Paz, Bolivia. Their structures were elucidated by extensive NMR and MS spectroscopic analyses. The isolated compounds were evaluated for their cytotoxicity and apoptotic capacity against four human cancer cell lines. 2-Benzyl-3-thioxohexahydropyrrolo[1,2-c]imidazole-1-one (1) showed slight cytotoxic activity against all the cancer cell lines which were tested, with IC50 values ranging from 27.45 ± 0.80 to 31.07 ± 0.87 μM. Moreover, N-(4-acetyl-5-methyl-5-phenyl-4,5-dihydro-1,3,4-thiadiazol-2-yl) acetamide (2) showed significant anti-cancer potential, with IC50 values between 1.26 ± 0.57 μM and 1.37 ± 0.09 μM against all human cancer cell lines which were tested. Treatment of tumour cell lines with the compounds caused an increase in the apoptotic rate of these cells, observing that compound 2 presented an apoptotic effect which was double with respect to the control (Dimethylenastron).

Summary Lysophosphatidate acyltransferase (LPAAT) catalyses the second step of the Kennedy pathway for triacylglycerol (TAG) synthesis. In this study we expressed Trapaeolum majus LPAAT in Brassica napus (B. napus) cv 12075 to evaluate the effects on lipid synthesis and estimate the flux control coefficient for LPAAT.We estimated the flux control coefficient of LPAAT in a whole plant context by deriving a relationship between it and overall lipid accumulation, given that this process is a exponential.Increasing LPAAT activity resulted in greater TAG accumulation in seeds of between 25% and 29%; altered fatty acid distributions in seed lipids (particularly those of the Kennedy pathway); and a redistribution of label from 14C‐glycerol between phosphoglycerides.Greater LPAAT activity in seeds led to an increase in TAG content despite its low intrinsic flux control coefficient on account of the exponential nature of lipid accumulation that amplifies the effect of the small flux increment achieved by increasing its activity. We have also developed a novel application of metabolic control analysis likely to have broad application as it determines the in planta flux control that a single component has upon accumulation of storage products.

In this study we aim to show how the peltate leaves of Colocasia fallax Schott and Tropaeolum majus L., despite their compact design, achieve a rigid connection between petiole and lamina. We have combined various microscopy techniques and computed tomography (CT) scanning for the analysis of the basic structure of the plant’s stabilization system. Mechanical tests yielded key mechanical parameters and allowed us to assess the mode of failure. The results of the tests were further processed in a finite element method (FEM) analysis. We were able to show that both plants are able to endure high loads irrespective of the different composition of the supporting structure. C. fallax forms many separate branched strands, whereas T. majus forms fewer strands of greater diameter interconnected in the centre of the transition area, forming a bundle of irregular orientation. This results in different ways to dissipate loads on the lamina. In C. fallax we observed the outer strands of the strengthening tissue under high stress while the inner bundle carries little load. In T. majus the load is distributed more evenly through the juncture in the middle of the transition area. Potential applications include the construction of biomimetical flying roofs.

Mashua (Tropaeolum tuberosum) is an important food in certain areas of the Andean region, where it is popularly believed to possess medicinal properties. Several studies have previously shown the potential of this tuber as a source of bioactive compounds. Traditionally, the tuber is exposed to the sun before consumption, in order to reduce its bitterness. The present work aims to study, at the proteome level, the differential abundance of proteins in tubers subjected to different postharvest treatments: sun-exposure (SUN), shade (SHA), refrigeration (COLD) and shade combined with sun-exposure (SHA-SUN) compared to recently harvested tubers (INIT). Results showed that sun exposure for prolonged times (9 days) resulted in increased abundance of proteins classified as heat shock proteins, intracellular traffic, disease/defense and protein degradation. Our results reflect that the sun treatment activates defense systems and osmoprotection adjustment against water loss and reactive oxygen species.

Ethnopharmacological relevance Tropaeolum tuberosum Ruiz & Pavon (Tropaeolaceae). Sim (commonly called Mashua) is an indigenous plant that has medicinal values for various ethnic groups of the regions of the Andes mountain range of South America, which use it for the treatment of diseases venereal, lung and skin; for the healing of internal and external wounds; and as an analgesic for kidney and bladder pain. Aim of the review: We critically summarised the current evidence on the botanic characterisation and distribution, ethnopharmacology, secondary metabolites, pharmacological activities, qualitative and quantitative analysis, and toxicology of T. tuberosum. Materials and methods The relevant information on T. tuberosum was gathered from worldwide accepted scientific databases via electronic search (Google scholar, Elsevier, SciFinder, ScienceDirect, PubMed, SpringerLink, Web of Science, Scopus, Wiley Online, Mendeley, Scielo and Dialnet electronic databases). Information was also obtained from the literature and books as well as PhD and MSc dissertations. Plant names were validated by ‘The Plant List’ ( www.theplantlist.org ). Results T. tuberosum has diverse uses in local and popular medicine, specifically for relieving pain and infections in humans. Regarding its biological activities, polar extracts (aqueous, hydroalcoholic) and isolated compounds from the tubers have exhibited a wide range of in vitro and in vivo pharmacological effects, including antibacterial, antioxidant, anti-inflammatory activities. Quantitative analysis (e.g., NMR, HPLC, GC-MS) indicated the presence of a set of secondary metabolites, including hydroxybenzoic acids, tannins, flavanols, anthocyanins, glucosinolates, isothiocyanates, phytosterols, fatty acids and alkamides in the tubers of T. tuberosum. Likewise, glucosinolates have been identified in the seeds and isothiocyanates have been detected in leaves, flowers and seeds. Conclusions T. tuberosum has been tested for various biological activities and the extracts (tubers in particular) demonstrated a promising potential as an antibacterial, antioxidant, anti-inflammatory and inhibitors of benign prostatic hyperplasia. A lack of alignment between the ethno-medicinal uses and existing biological screenings was observed, indicating the need to explore its potential for the treatment against respiratory affections, urinary affections and blood diseases. Likewise, it is necessary to analyse deeply the relationship that exists between the different tuber colours of T. tuberosum and its use for the treatment of certain diseases. Validation of clinical studies of the antibacterial, antioxidant/anti-inflammatory, anti-spermatogenic activities and as inhibitors of benign prostatic hyperplasia is required. Moreover, studies on the toxicity, bioavailability, and pharmacokinetics, in addition to clinical trials, are indispensable for assessing the safety and efficacy of the active metabolites or extracts obtained from T. tuberosum. Other areas that need investigation are the development of future applications based on their active metabolites, such as neurodegenerative diseases (Alzheimer's disease, Parkinson's disease, Huntington's disease). Finally, the work purposes to motivate other research groups to carry out a series of scientific studies that can fill the gaps that exist with respect to Mashua properties, and thus be able to change the focus of T. tuberosum (Mashua) that currently has in the consumer society.

Mashua (Tropaeolum tuberosum) is a tuber crop indigenous to the Andean highlands and of economic value to its native people as a food and medicinal crop. Field plants often exhibit symptoms typical of virus infection, including interveinal chlorosis, leaf cupping, distortion, and stunting. Using an herbaceous host range, at least 1 mechanically transmissible virus was detected in each of 10 accessions. All virus isolates tested reacted to potyvirus group antiserum in indirect enzyme-linked immunosorbent assays (ELISA). Results of monoclonal antibody testing using antigen coated plate (ACP) ELISA, where virions are disrupted to expose cryptotopes, indicated that isolate Tt 4 has epitopes in common with maize dwarf mosaic potyvirus A (MDMV-A). Polyclonal antiserum produced to purified preparations of the Tt 4 isolate reacted in indirect ELISA to Tt 4, the common strain of potato potyvirus Y (PVY-C), less strongly to tobacco etch potyvirus (TEV) and potato potyvirus A (PVA), but not to 11 other potyviruses, including MDMV-A. Conversely, the Tt 4 isolate reacted strongly in indirect ELISA to antisera to bean common mosaic potyvirus (BCMV) and watermelon mosaic potyvirus 2 (WMV-2), but not antisera to 12 other potyviruses. Our data suggest that isolate Tt 4 is a distinct potyvirus closely related to and sharing external epitopes with PVY-C, TEV, PVA, WMV-2, and BCMV, and buried epitopes with MDMV-A. The virus has been tentatively named Tropaeolum mosaic potyvirus (TropMV). A survey of T. tuberosum in Ecuador revealed that TropMV was widespread, being detected by ACP-ELISA in 34 of 46 accessions at the National Institute for Agricultural Research's Santa Catalina Research Station, Quito, and all eight production fields surveyed.

Background Nasturtium (Indian cress, Tropaeolum majus) is known for its pharmacological value in the treatment of bacterial infections of the upper air tract and urinary bladder. However, scientific data on the anti-inflammatory potency in human-derived cells is missing. Purpose The aim of this study was to investigate the potential of nasturtium to inhibit the lipopolysaccharide (LPS) induced inflammatory response in primary human cells of the immune system. Study design The anti-inflammatory activities of nasturtium and its fractions were evaluated via regulation of arachidonic acid (AA) pathway and MAPK kinase cascade. Fraction H4 which was responsible for the anti-inflammatory effects was further characterized. Methods Human peripheral blood mononuclear cells (PBMC) were either treated with plant extracts or fractions thereof, stimulated with LPS and/or N-formyl-methionyl-leucyl-phenylalanine (fMLP) and analysed for COX and LOX, release of prostaglandin PGE2, leukotriene LTB4, TNF-alpha and ERK signaling pathway activation. The plant extracts were separated into four fractions by HPLC; fraction H4 was subjected to UHPLC-ToF/MS analysis to identify potential bioactive compounds. Results We found that aqueous extracts of nasturtium did exert strong concentration dependent suppression of LPS-triggered TNF-alpha release and COX pathway signaling, including PGE2 synthesis. Whereas COX-1 protein expression was not impacted, LPS-triggered COX-2 protein expression was concentration dependently blocked by the plant extract but not COX-2 enzyme activity. These findings suggest a mechanism of action for the plant extract which is different from non-steroidal anti-inflammatory drugs (NSAIDs). Moreover, the plant extract blocked leukotriene LTB4 release, the major end product of the 5-LOX pathway from PBMC. Down-regulation of ERK1/2 and c-Jun activation preceded COX-2 suppression upon plant extract treatment in the presence of LPS. Using HPLC separation of the aqueous extract followed by metabolomic analysis we could limit the number of relevant bioactive compounds in the extract to about 50. Conclusions This study provides a rationale for the anti-inflammatory efficacy of nasturtium observed in man and gives first insight into the underlying molecular mechanisms.

Morphological, technological and nutritional analyses were done in two scarcely studied starches from Andean tubers (mashua and melloco). The low sedimentation values, and the high zeta potential of mashua and melloco starches in cold dispersions, as consequence of their electronegativity, indicated a better behaviour as stabilizer than potato starch. During heating, mashua and melloco starches presented much higher viscosity than potato starch, associated with their higher average particle size and greater amylose content. DSC and TGA analyses indicated that melloco starch had the highest gelatinization enthalpy ΔH (12.32 J g) and degradation temperature (270 °C), in comparison with potato starch, which are indicators of a better thermal resistance. Consequently, extracted mashua and melloco starches could be excellent and cost-effective thickening or gelling agents in both foods and a wide range of biomaterials. Mashua and melloco starches exhibited a digestion rate close to 80%, which agreed with the low resistant starch content., The authors are grateful for the financing of MINECO-Spain (AGL2014-53445-R), the Community of Madrid (ALIBIRD-CM S-2013/ABI-272), the support of the National Secretary of Higher Education, Science, Technology and Innovation of Ecuador (SENESCYT), and the collaboration of Nerea Muñoz (CIAL, CSIC-UAM) and Sergey Gorbatov (Vrije Universiteit Amsterdam).

Ethno–pharmacological relevance Tropaeolum tuberosum, commonly known as “Mashua”, is one of the plants most frequently used by Andean (Peruvian–Bolivian) people as food and medicine. It is used as a remedy against a wide range of diseases, especially those related with inflammation. Objectives This study aims to identify compounds active against inflammatory related conditions. Materials and methods A bioassay–guided isolation of anti–inflammatory compounds from black and purple tubers of T. tuberosum was performed measuring TNF–α and NF–κB production in THP–1 monocytic cells. Results The bioassay–guided isolation led to one active compound from purple T. tuberosum, N–oleoyldopamine (1), and another active compound from black T. tuberosum, N–(2–Hydroxyethyl)–7Z,10Z,13Z,16Z–docosatetraenamide (2). Both compounds displayed anti–TNF–α activity with IC50 values of 3.12 ± 0.19 μM and 1.56 ± 0.15 μM, respectively. Also, both compounds suppressed NF–κB with IC50 of 3.54 ± 0.02 μM and 1.77 ± 0.07 μM, respectively. Conclusions We identified bioactive compounds from purple and black Tropaeolum tuberosum responsible for their anti–inflammatory activity: N–oleoyldopamine (1) and N–(2–Hydroxyethyl)–7Z,10Z,13Z,16Z–docosatetraenamide (2). This is the first report which isolates these compounds from T. tuberosum and describes their anti–inflammatory activities.

The objective of this study was to characterize the nutritional compounds of interest present in vegetables known as non-conventional, in Brazil. The following evaluations were carried out: antioxidant activity, phenolic compounds, vitamin C, calories, carbohydrates, humidity, lipids, proteins, fiber, acidity and quantification of minerals (P, K, Ca, Mg, S, Cu, Fe, Mn, Zn and B). The species studied were Amaranthus hybridus L., Amaranthus viridis L., Basella alba L., Eryngium campestre L., Hibiscus sabdariffa L., Lactuca canadensis L., Rumex acetosa L., Stachys byzantina K. Koch, Tropaeolum majus L. and Xanthosoma sagittifolium L. Representative samples of plant structures of interest were harvested from each species suitable for human consumption such as leaves, flowers and flower buds. The results were submitted to multivariate analysis - principal components analysis (PCA). All the species present nutritional compounds of interest in different levels among the evaluated structures.

SCOPE Benzyl isothiocyanate (BITC), which occurs in Brassicales, has demonstrated chemopreventive potency and cancer treatment properties in cell and animal studies. However, fate of BITC in human body is not comprehensively studied. Therefore, the present human intervention study investigates the metabolism of the glucosinolate (GSL) glucotropaeolin and its corresponding BITC metabolites. Analyzing BITC metabolites in plasma and urine should reveal insights about resorption, metabolism, and excretion. METHODS AND RESULTS Fifteen healthy men were randomly recruited for a cross-over study and consumed 10 g freeze-dried Indian cress as a liquid preparation containing 1000 μmol glucotropaeolin. Blood and urine samples were taken at several time points and investigated by LC-ESI-MS/MS after sample preparation using SPE. Plasma contained high levels of BITC-glutathione (BITC-GSH), BITC-cysteinylglycine (BITC-CysGly), and BITC-N-acetyl-L-cysteine (BITC-NAC) 1-5 h after ingestion, with BITC-CysGly appearing as the main metabolite. Compared to human plasma, the main urinary metabolites were BITC-NAC and BITC-Cys, determined 4-6 h after ingestion. CONCLUSION This study confirms that consumption of Indian cress increases the concentration of BITC metabolites in human plasma and urine. The outcome of this human intervention study supports clinical research dealing with GSL-containing innovative food products or pharmaceutical preparations.

Xyloglucan XG (molecular weight 462 kDa, 1,4-/1,4,6-(pGlc) linked backbone, side chains of 1-pXyl, 1,2-pXyl, 1-p-Gal) was isolated from the seeds of Tropaeolum majus. XG (100 μg/mL) induced terminal cellular differentiation of human keratinocytes, as demonstrated by immunofluorescence staining and Western blot using cytokeratin 10 and involucrin as marker proteins. Differentiation was also induced by XG-derived oligosaccharides (degree of polymerization 7-9). Quantitative real-time polymerase chain reaction (qPCR) revealed the induction of gene expression of typical differentiation markers (cytokeratin, filaggrin, involucrin, loricrin, transglutaminase) in a time-dependent manner. Whole human genome microarray indicated that most of upregulated genes were related to differentiation processes. Microarray findings on selected genes were subsequently confirmed by qPCR. For identification of the molecular target of xyloglucan PAGE of keratinocyte membrane preparations was performed, followed by blotting with fluorescein isothiocyanate-labeled XG. XG interacting proteins were characterized by MS. Peptide fragments of epidermal growth factor receptor (EGFR) and integrin β4 were identified. Subsequent phospho-kinase array indicated that phosphorylation of EGFR and transcription factor cAMP response element-binding protein (CREB) was decreased in the XG-treated cells. Thus, the XG-induced differentiation of keratinocytes is proposed to be mediated by the inhibition of the phosphorylation of EGFR, leading to a dimished CREB activation, which is essential for the activation of cellular differentiation.

Two new South American species of Tropaeolum L. (Tropaeolaceae), T. slanisii Bulacio from Salta, Argentina, and T. kieslingii Bulacio from Tarija, Bolivia, found during a revision of the genus for the Flora of Argentina, are described, illustrated, and compared with related species. Both new species belong to Tropaeolum sect. Tropaeolum, which is characterized by the strongly heteromorphic petals, usually ciliate on the margins, and the entire or more or less deeply incised leaf blades.

Background: Tropaeolum pentaphyllum Lam. tubers (Tropaeolaceae) are known and used as a condiment and for the treatment of skin infections in Southern Brazil. However, its activity and composition has not yet been investigated. Thus, different extracts and the essential oil from the tubers were tested against a range of microorganisms. The most active extracts were submitted to chromatographic analysis. Methods: Hydroalcoholic extract (70%), fractions of it, and the essential oil from the tubers were tested against several bacteria, yeasts and molds, furnishing the corresponding inhibitory, bactericidal and fungicidal minimal concentration values. The most active extracts were submitted to GC-MS investigation. Results: The strongest effects against different strains of microorganisms, such as Gram-positive and negative bacteria, Candida spp. and dermatophytes were observed for the essential oil and the chloroform fraction, with minimal inhibitory concentrations (MICs) well below 200 µg/mL. GC-MS analysis revealed that the major essential oil constituent is benzyl isothiocyanate (BITC), while the chloroform fraction is constituted of BITC, amides, sulfur, fatty acids and its esters, all compounds that may be related to the demonstrated activity. Conclusions: Overall, the results support the popular use of the plant for the treatment of skin infections, and revealed the main active compounds.

A deep-sequencing approach was pursued utilizing 454 and Illumina sequencing methods to discover new genes involved in xyloglucan biosynthesis. cDNA sequences were generated from developing nasturtium (Tropaeolum majus) seeds, which produce large amounts of non-fucosylated xyloglucan as a seed storage polymer. In addition to known xyloglucan biosynthetic genes, a previously uncharacterized putative xyloglucan galactosyltransferase was identified. Analysis of an Arabidopsis thaliana mutant line defective in the corresponding ortholog (AT5G62220) revealed that this gene shows no redundancy with the previously characterized xyloglucan galactosyltransferase, MUR3, but is required for galactosyl-substitution of xyloglucan at a different position. The gene was termed XLT2 for Xyloglucan L-side chain galactosylTransferase position 2. It represents an enzyme in the same subclade of glycosyltransferase family 47 as MUR3. A double mutant defective in both MUR3 (mur3.1) and XLT2 led to an Arabidopsis plant with xyloglucan that consists essentially of only xylosylated glucosyl units, with no further substitutions.

Phenolic compounds from mashua tuber were evaluated as potential antioxidants to retard the oxidation of crude soybean oil submitted to accelerated storage and frying. During the accelerated storage, an ethanolic crude extract, a purified extract, an aqueous fraction and an ethyl acetate fraction from mashua containing different gallic acid equivalent concentrations (100, 300 and 600 ppm) in oil were evaluated at 55 °C. After 15 days of storage, better effects were evidenced against soybean oil oxidation at 300 and 600 ppm of ethyl acetate fraction in comparison to 200 ppm butylated hydroxytoluene and the control (no antioxidant added). During the frying process at ∼180 °C, principal component analysis revealed that the content of trienes and dienes were strongly correlated with the frying batch. Ethyl acetate fraction at 200 ppm showed the highest efficacy against oil oxidation in terms of polar compound values, free fatty acids and conjugated dienes and trienes in comparison to the oil containing 200 ppm tert-butylhydroquinone and control. Differential scanning calorimetry corroborated the efficacy of ethyl acetate fraction phenolic and it is strongly recommended as method for validation of results. This study provides strong evidence related to the excellent protective effects against soybean oil oxidation of mashua phenolics. This crop could be utilized as an alternative source of natural antioxidants by the oil industry.

Ethnopharmacological relevance Previous studies have shown that the extracts obtained from Tropaeolum majus L., and its main compound isoquercitrin (ISQ), exhibit pronounced diuretic effects, supporting the ethnopharmacological use of this plant. The aim of this study was to evaluate the efficacy and mechanisms underlying the diuretic action of an ethanolic extract of Tropaeolum majus (HETM), its purified fraction (TMLR), and its main compound ISQ, in spontaneously hypertensive rats (SHR). Materials and methods The diuretic effects of HETM (300 mg/kg; p.o. ), TMLR (100 mg/kg; p.o. ), and ISQ (10 mg/kg; p.o. ), were compared with classical diuretics in 7 days repeated-dose treatment. The urinary volume, sodium, potassium, chloride, bicarbonate, conductivity, pH and density were estimated in the sample collected for 15 h. The plasmatic concentration of sodium, potassium, urea, creatinine, aldosterone, vasopressin, nitrite and angiotensin converting enzyme (ACE) activity were measured in samples collected at the end of the experiment (seventh day). Using pharmacological antagonists or inhibitors, we determine the involvement of bradykinin, prostaglandin and nitric oxide (NO) in ISQ-induced diuresis. In addition, reactive oxygen species (ROS) and the activity of erythrocytary carbonic anhydrase and renal Na + /K + /ATPase were evaluated in vitro . Results HETM, TMLR and ISQ increased diuresis similarly to spironolactone and also presented K + -sparing effects. All groups presented both plasmatic aldosterone levels and ACE activity reduced. Previous treatment with HOE-140 (a B2-bradykinin receptor antagonist), or indomethacin (a cyclooxygenase inhibitor), or L-NAME (a NO synthase inhibitor), fully avoided the diuretic effect of ISQ. In addition, the 7 days treatment with ISQ resulted in increased plasmatic levels of nitrite and reducing ROS production. Moreover, the renal Na + /K + /ATPase activity was significantly decreased by ISQ. Conclusion Our results suggest that the mechanisms through ISQ and extracts of Tropaeolum majus increase diuresis in SHR rats are mainly related to ACE inhibition, increased bioavailability of bradykinin, PGI2, and nitric oxide, besides an inhibitory effect on Na + /K + -ATPase.

Transcriptome analysis based on deep expressed sequence tag (EST) sequencing allows quantitative comparisons of gene expression across multiple species. Using pyrosequencing, we generated over 7 million ESTs from four stages of developing seeds of Ricinus communis, Brassica napus, Euonymus alatus and Tropaeolum majus, which differ in their storage tissue for oil, their ability to photosynthesize and in the structure and content of their triacylglycerols (TAG). The larger number of ESTs in these 16 datasets provided reliable estimates of the expression of acyltransferases and other enzymes expressed at low levels. Analysis of EST levels from these oilseeds revealed both conserved and distinct species-specific expression patterns for genes involved in the synthesis of glycerolipids and their precursors. Independent of the species and tissue type, ESTs for core fatty acid synthesis enzymes maintained a conserved stoichiometry and a strong correlation in temporal profiles throughout seed development. However, ESTs associated with non-plastid enzymes of oil biosynthesis displayed dissimilar temporal patterns indicative of different regulation. The EST levels for several genes potentially involved in accumulation of unusual TAG structures were distinct. Comparison of expression of members from multi-gene families allowed the identification of specific isoforms with conserved function in oil biosynthesis. In all four oilseeds, ESTs for Rubisco were present, suggesting its possible role in carbon metabolism, irrespective of light availability. Together, these data provide a resource for use in comparative and functional genomics of diverse oilseeds. Expression data for more than 350 genes encoding enzymes and proteins involved in lipid metabolism are available at the 'ARALIP' website (http://aralip.plantbiology.msu.edu/).

Fluorescently labeled derivatives of a xyloglucan (XG) nonasaccharide Glc(4)Xyl(3)Gal(2) (XLLG) were used as glycosyl acceptors in assays of xyloglucan endotransglycosylase (XET) from germinated nasturtium (Tropaeolum majus) seeds. We have investigated how the type of the oligosaccharide label influences the kinetic parameters of the reaction. The fluorescent probes used to label XLLG were anthranilic acid (AA), 8-aminonaphtalene-1,3,6-trisulfonic acid (ANTS), fluorescein isothiocyanate (FITC), and sulforhodamine (SR), respectively. The obtained data were compared with those of the reactions where aldose and/or alditol forms of tritium-labeled xyloglucan-derived nonasaccharide served as the respective acceptors. Modification at C-1 of the reducing-end glucose in XLLG by substitution with the fluorophore markedly affected the kinetic parameters of the reaction. The Michaelis constants K(m) for individual acceptors increased in the order [1-(3)H]XLLGXLLG-SR[1-(3)H]XLLGolXLLG-FITCXLLG-ANTSXLLG-AA, while the turnover numbers characterized by k(cat) decreased in the order XLLG-FITCXLLG-SRXLLG-ANTS[1-(3)H]XLLGol[1-(3)H]XLLGXLLG-AA. Catalytic efficiency (expressed as k(cat)/K(m)) with XLLG labeled with SR or FITC was 15 and 28 times, respectively, higher than with the tritium-labeled natural substrate [1-(3)H]XLLG. Comparison of the kinetic parameters found with acceptors labeled with different types of labels enables to select the most effective substrates for the high-throughput assays of XET.

The species of Tropaeolum sect. Chilensia Sparre, are discussed. Their phytogeography, ecology, evolution and pollination are considered. Photographs of habitats, and of most of the species taken in the wild, are shown. A synopsis of all the species in the section, and a new arrangement of subsections is provided.

Tropaeolum austropurpureum (J.M. Watson & A.R. Flores) J.M. Watson & A.R. Flores, stat. nov., based on Tropaeolum hookerianum Barneoud subsp. austropurpureum J.M. Watson & A.R. Flores, is described, and the new rank is published. Details of the discovery, ecology, and possible evolution of this species are discussed. Instructions for its cultivation are given.

Five forms of xyloglucan endotransglycosylase/hydrolase (XTH) differing in their isoelectric points (pI) were detected in crude extracts from germinating nasturtium seeds. Without further fractionation, all five forms behaved as typical endotransglycosylases since they exhibited only transglycosylating (XET) activity and no xyloglucan-hydrolysing (XEH) activity. They all were glycoproteins with identical molecular mass, and deglycosylation led to a decrease in molecular mass from approximately 29 to 26.5 kDa. The major enzyme form having pI 6.3, temporarily designated as TmXET(6.3), was isolated and characterized. Molecular and biochemical properties of TmXET(6.3) confirmed its distinction from the XTHs described previously from nasturtium. The enzyme exhibited broad substrate specificity by transferring xyloglucan or hydroxyethylcellulose fragments not only to oligoxyloglucosides and cello-oligosaccharides but also to oligosaccharides derived from beta-(1,4)-d-glucuronoxylan, beta-(1,6)-d-glucan, mixed-linkage beta-(1,3; 1,4)-d-glucan and at a relatively low rate also to beta-(1,3)-gluco-oligosaccharides. The transglycosylating activity with xyloglucan as donor and cello-oligosaccharides as acceptors represented 4.6%, with laminarioligosaccharides 0.23%, with mixed-linkage beta-(1,3; 1,4)-d-gluco-oligosaccharides 2.06%, with beta-(1,4)-d-glucuronoxylo-oligosaccharides 0.31% and with beta-(1,6)-d-gluco-oligosaccharides 0.69% of that determined with xyloglucan oligosaccharides as acceptors. Based on the sequence homology of tryptic fragments with the sequences of known XTHs, the TmXET(6.3) was classified into group II of the XTH phylogeny of glycoside hydrolase family GH16.

SUMMARYEcuadorean mashua (Tropaeolum tuberosum) germplasm has been found to be widely infected with the potyvirus Tropaeolum mosaic virus (TropMV). The objective of this research was to produce virus-tested (VT) germplasm to compare growth in vitro and in the field with virus-infected (V) germplasm. Twenty-three of 25 apical dome-derived clones tested free of virus based on bioassays using Nicotiana benthamiana and Chenopodium quinoa. In vitro-generated plant tissue was just as effective for determining VT status as greenhouse-generated plant tissue. Genotype rather than virus-infection status appeared to have a greater effect on in vitro proliferation. There were no differences in in vitro rooting among the genotypes or between the VT clones compared to the V clones, with at least 90% of the microcuttings rooting. However, rooted microcuttings of V clones were taller than rooted microcuttings of VT clones. Plants were readily re-established in a greenhouse at the USDA, Foreign Disease-Weed Science Research Unit at Fort Detrick, USA. In field experiment 1, ca. 75% of the plants survived field transplanting and VT plants of genotype 1147 had greater tuber weight (928 g) than V plants (235 g). In field experiment 2, plant mortality was high one month after field transplanting. Genotypes 1093 (59%) and 1141 (54%) had higher survival than genotype 1147 (44%); however, survival did not differ between the VT (46%) and V (59%) plants of all genotypes. No differences were noted in field performance for the three genotypes after 10 months of growth. Although overall tuber yield among the V, VT and VTR (reinfected-VT plants) did not differ, V plants produced big tubers that weighed more than those from VT plants. Thirty-three percent of the VT plants became reinfected and 42% of the V plants tested negative after 10 months in the field based on double-antibody-sandwich ELISA.

Tropaeolum majus L. (Tropaeolaceae), popularly known as “chaguinha”, is well recognized in Brazilian traditional medicine as diuretic agent, although no scientific data have been published to support this effect. Aim of the study To evaluate the diuretic activity of the infusion and the hydroethanolic extract (HETM) of Tropaeolum majus , and possible mechanism of action. Material and methods The infusions (2,5 – 10%) and the HETM doses (150, 300 mg/kg) were orally administered to rats. Urinary excretion, the electrolytes levels, and urea and creatinine were measured in of saline-loaded rats. Results The oral administration of 10% (corresponding to 500 mg/kg) of the infusion increased significantly the urinary Na + excretion. Only the oral administration of 300 mg/kg of HETM increased significantly the urinary and Na + excretion. Prolonged administration of the HETM (300 mg/kg) significantly increased diuresis and the urinary excretion of Na + , but others parameters were unaffected. To gain some evidence in possible involvement of prostaglandins system in diuretic action, the oral administration of HETM (300 mg/kg) in association indomethacin (5 mg/kg) reduced the urinary and sodium excretion when compared only HETM group. Conclusion The results suggest that HETM could present compound(s) responsible for diuretic activities with no signs of toxicity, and the mechanism could involve prostaglandin system.

International audience; Indian cress (Tropaeolum majus L.) absolute was studied by GC-olfactometry (VIDEO-Sniff method) in order to identify odor-active aroma compounds. Because of its fruity-sulfury odor note, a compound that has never been identified in plant extracts before stood out: O,S-diethyl thiocarbonate, present at 0.1% (percentage of the total GC/FID area) in the extract. GCxGC-TOFMS allowed for a clean mass spectrum to be obtained, and isolation by preparative GC followed by NMR studies allowed its identification. Here, we report on the first detection of O,S-diethyl thiocarbonate in Indian cress absolute by GC-olfactometry/VIDEO-Sniff and on its isolation and identification. The synthesis and odor evaluation of its homologues are presented.

Tropaeolum tuberosum Ruiz & Pavon, along with other several species, is an edible-tuber crop that grows in the Andean region. Folk medicine describes the use of mashua to reduce reproductive function in men. The present study aimed to evaluate the effect of mashua (1 g kg(-1)) on sperm production in rats during 7, 12, 21 and 42 days of treatment. The following parameters were assessed: reproductive organ weights, spermatid count and daily sperm production (DSP), sperm count in epididymis and sperm transit and serum testosterone levels. Freeze-dried extract of mashua had 3.7 g 100 g(-1) of benzyl glucosinolate. Mashua-treated rats showed a reduction in testicular spermatid number and DSP from day 12 to day 42; meanwhile, the effect of mashua was noted in epididymal sperm count after 12 and 42 days of treatment. In addition, epididymal sperm transit time was delayed at day 7 and it was accelerated on days 12 and 21 of treatment. No differences in serum testosterone levels were found between rats treated with vehicle and mashua after 42 days of treatment. Finally, mashua reduces testicular function after one spermatogenic cycle by reducing spermatid and sperm number, DSP and epididymal sperm transit time.

Summary A full-length cDNA encoding a putative diacylglycerol acyltransferase 1 (DGAT1, EC 2.3.1.20) was obtained from Tropaeolum majus (garden nasturtium). The 1557-bp open reading frame of this cDNA, designated TmDGAT1, encodes a protein of 518 amino acids showing high homology to other plant DGAT1s. The TmDGAT1 gene was expressed exclusively in developing seeds. Expression of recombinant TmDGAT1 in the yeast H1246MATα quadruple mutant (DGA1, LRO1, ARE1, ARE2) restored the capability of the mutant host to produce triacylglycerols (TAGs). The recombinant TmDGAT1 protein was capable of utilizing a range of 14C-labelled fatty acyl-CoA donors and diacylglycerol acceptors, and could synthesize 14C-trierucin. Collectively, these findings confirm that the TmDGAT1 gene encodes an acyl-CoA-dependent DGAT1. In plant transformation studies, seed-specific expression of TmDGAT1 was able to complement the low TAG/unusual fatty acid phenotype of the Arabidopsis AS11 (DGAT1) mutant. Over-expression of TmDGAT1 in wild-type Arabidopsis and high-erucic-acid rapeseed (HEAR) and canola Brassica napus resulted in an increase in oil content (3.5%–10% on a dry weight basis, or a net increase of 11%–30%). Site-directed mutagenesis was conducted on six putative functional regions/motifs of the TmDGAT1 enzyme. Mutagenesis of a serine residue in a putative SnRK1 target site resulted in a 38%–80% increase in DGAT1 activity, and over-expression of the mutated TmDGAT1 in Arabidopsis resulted in a 20%–50% increase in oil content on a per seed basis. Thus, alteration of this putative serine/threonine protein kinase site can be exploited to enhance DGAT1 activity, and expression of mutated DGAT1 can be used to enhance oil content.

Nasturtium (Tropaeolum majus L.) contains high concentrations of benzylglcosinolate. We found that a hydrolysis product of benzyl glucosinolate—the benzyl isothiocyanate (BITC)—modulates the intracellular localization of the transcription factor Forkhead box O 1 (FOXO1). FoxO transcription factors can antagonize insulin effects and trigger a variety of cellular processes involved in tumor suppression, longevity, development and metabolism. The current study evaluated the ability of BITC—extracted as intact glucosinolate from nasturtium and hydrolyzed with myrosinase—to modulate i) the insulin-signaling pathway, ii) the intracellular localization of FOXO1 and, iii) the expression of proteins involved in gluconeogenesis, antioxidant response and detoxification. Stably transfected human osteosarcoma cells (U-2 OS) with constitutive expression of FOXO1 protein labeled with GFP (green fluorescent protein) were used to evaluate the effect of BITC on FOXO1. Human hepatoma HepG2 cell cultures were selected to evaluate the effect on gluconeogenic, antioxidant and detoxification genes and protein expression. BITC reduced the phosphorylation of protein kinase B (AKT/PKB) and FOXO1; promoted FOXO1 translocation from cytoplasm into the nucleus antagonizing the insulin effect; was able to down-regulate the gene and protein expression of gluconeogenic enzymes; and induced the gene expression of antioxidant and detoxification enzymes. Knockdown analyses with specific siRNAs showed that the expression of gluconeogenic genes was dependent on nuclear factor (erythroid derived)-like2 (NRF2) and independent of FOXO1, AKT and NAD- dependent deacetylase sirtuin-1 (SIRT1). The current study provides evidence that BITC might have a role in type 2 diabetes T2D by reducing hepatic glucose production and increasing antioxidant resistance.

Despite the central role of xyloglucan (XyG) in plant cell wall structure and function, important details of its biosynthesis are not understood. To identify the gene(s) responsible for synthesizing the β-1,4 glucan backbone of XyG, we exploited a property of nasturtium ( Tropaeolum majus ) seed development. During the last stages of nasturtium seed maturation, a large amount of XyG is deposited as a reserve polysaccharide. A cDNA library was produced from mRNA isolated during the deposition of XyG, and partial sequences of 10,000 cDNA clones were determined. A single member of the C subfamily from the large family of cellulose synthase-like ( CSL ) genes was found to be overrepresented in the cDNA library. Heterologous expression of this gene in the yeast Pichia pastoris resulted in the production of a β-1,4 glucan, confirming that the CSLC protein has glucan synthase activity. The Arabidopsis CSLC4 gene, which is the gene with the highest sequence similarity to the nasturtium CSL gene, is coordinately expressed with other genes involved in XyG biosynthesis. These and other observations provide a compelling case that the CSLC gene family encode proteins that synthesize the XyG backbone.

Long-term memory (LTM) formation usually requires repeated, spaced learning events and is achieved by the synthesis of specific proteins. Other memory forms require a single learning experience and are independent of protein synthesis. We investigated in two closely related parasitic wasp species,Cotesia glomerataandCotesia rubecula, whether natural differences in foraging behaviour are correlated with differences in LTM acquisition and formation. These parasitic wasp species lay their eggs in young caterpillars of pierid butterflies and can learn to associate plant odours with a successful egg laying experience on caterpillars on the odour-producing plant. We used a classical conditioning set-up, while interfering with LTM formation through translation or transcription inhibitors. We show here thatC. rubeculaformed LTM after three spaced learning trials, whereasC. glomeratarequired only a single trial for LTM formation. After three spaced learning trials, LTM formation was complete within 4 h inC. glomerata, whereas inC. rubecula, LTM formation took 3 days. Linking neurobiology with ecology, we argue that this species-specific difference in LTM acquisition and formation is adaptive given the extreme differences in both the number of foraging decisions of the two wasp species and in the spatial distributions of their respective hosts in nature.

Maximum parsimony and Bayesian likelihood analysis of ribosomal DNA internal transcribed spacer (ITS) sequences from 55 samples representing 25 species, subspecies, and/or hybrids of Tropaeolum sect. Chilensia yielded a tree with a large proportion of well-supported nodes at the interspecific and intersubspecific level. However, the tree shows samples from two species, T.azureum and T. brachyceras, arising in two different positions. Samples representing subspecies of T. hookerianum and T. leptophyllum suggest that these species are polyphyletic. The data corroborate evidence for hybridization between T. brachyceras and T. tricolor. Consideration of interfertility data, past and present Chilean ecology, and empirical evidence for the behavior of phenotypic and genotypic characters in known hybrids suggest a high likelihood that reticulate evolution has played a role in the diversification of T. sect. Chilensia. This reticulate evolution may explain the discordance between the ITS tree and the conventional taxonomy. High divergence in ITS sequences between T. sect. Chilensia and other members of Tropaeolum prohibits reliable outgroup-rooting, but midpoint rooting places the root at a partition comprising taxa whose distribution conforms to a relictual eastern-western South American disjunction described for other taxa. Within the Chilean taxa, the analysis suggests that biogeographic diversification has been from the mesophytic southern habitats northward to central mediterranean and northern arid zones.

Using combinations of different polysaccharides as glycosyl donors and of oligosaccharides fluorescently labeled by sulforhodamine (SR) as glycosyl acceptors, we screened for the presence of transglycosylating activities in extracts from nasturtium (Tropaeolum majus). Besides xyloglucan endotransglycosylase/hydrolase (XTH/XET, EC 2.4.1.207) activity, which transfers xyloglucanosyl residues from xyloglucan (XG) to XG-derived oligosaccharides (XGOs), a glycosyl transfer from XG to SR-labeled cellooligosaccharides and laminarioligosaccharides has been detected. The XGOs also served as acceptors for the glycosyl transfer from soluble cellulose derivatives carboxymethyl cellulose and hydroxyethylcellulose. The effectivity of these polysaccharides as glycosyl donors for transfer to XG-derived octasaccharide [1-3H]XXLGol decreased in the order XG > HEC > CMC. Isoelectric focusing in polyacrylamide gels showed that bands corresponding to hetero-transglycosylase activities coincided with zones corresponding to XTH/XET. These results can be explained as due either to substrate non-specificity of certain isoenzymes of XTH/XET or to existence of enzymes catalyzing a hetero-transfer, that is the formation of covalent linkages between different types of carbohydrate polymers.

The contents and profile of polyphenols were analyzed in edible petals of nasturtium flowers (Tropaeolum majus) of three colors, and their oxygen radical absorbance capacities (ORAC) were compared. Three primary anthocyanins (ACNs) and 15 non-ACN phenolic compounds including hydroxycinammic acids (HCAs) and flavonoids (myricetin, quercetin, and kaempferol derivatives) were detected. Anthocyanin concentration was within 31.9 ± 21.7 and 114.5 ± 2.3 mg cyanidin-3-glucoside (cy-3-glu)/100 g fresh weight (FW) in yellow and red petals, respectively. The concentration of HCAs varied between 33.3 ± 7.1 and 235.6 ± 8.1 mg chlorogenic acid equivalents/100 g FW for red and yellow flowers, respectively. Red flowers had the highest level of flavonoids (315.1 ± 2.4 mg myricetin equivalents/100 g FW) and the highest ORAC radical-scavenging activity. These results show the diversity and abundance of polyphenolic compounds in nasturtium flowers, which could be the basis for applications in functional foods, cosmetics, and pharmaceuticals.

Edible flowers are commonly used in human nutrition and their consumption has increased in recent years. The aim of this study was to ascertain the nutritional composition and the content and profile of phenolic compounds of three edible flowers, monks cress (Tropaeolum majus), marigold (Tagetes erecta) and paracress (Spilanthes oleracea), and to determine the relationship between the presence of phenolic compounds and the antioxidant capacity. Proximate composition, total dietary fibre (TDF) and minerals were analysed according to official methods: total phenolic compounds (TPC) were determined with Folin-Ciocalteu’s reagent, whereas antioxidant capacity was evaluated using Trolox Equivalent Antioxidant Capacity (TEAC) and Oxygen Radical Absorbance Capacity (ORAC) assays. In addition, phenolic compounds were characterised by HPLC-DAD-MSn. In relation to the nutritional value, the edible flowers had a composition similar to that of other plant foods, with a high water and TDF content, low protein content and very low proportion of total fat—showing significant differences among samples. The levels of TPC compounds and the antioxidant capacity were significantly higher in T. erecta, followed by S. oleracea and T. majus. Thirty-nine different phenolic compounds were tentatively identified, with flavonols being the major compounds detected in all samples, followed by anthocyanins and hydroxycynnamic acid derivatives. In T. erecta small proportions of gallotannin and ellagic acid were also identified.

The floral ontogeny of three species of Tropaeolum was studied using scanning electron microscopy to find morphological evidence for discussing the systematic position of the family. The initiation of the androecium is highly unusual: there are always eight stamens which arise (1) either in a spiral sequence starting with the stamen opposite sepal four, running in a direction opposite to the sequence of the sepals, and with reversals in the direction of the spiral, or (2) as a sequence of paired and unpaired stamens. The floral symmetry changes twice during the development of the flower, from polysymmetrical at sepal and petal initiation, through oblique monosymmetry at stamen initiation, and ending with median monosymmetry in later developmental stages. The occurrence of median monosymmetry is a late-developmental event and is caused by the initiation of a hypanthial spur, and the unequal growth of the petals and styles. The origin for the unusual sequence of stamen initiation reflects a trend affecting the whole flower which is linked with the changing patterns of floral symmetry. Octandry is enhanced by multiple causes, such as the loss of two stamens in an originally diplostemonous androecium and the regulating pressure of the gynoecium. The change in symmetry during ontogeny is significant for discussing the systematic position of Tropaeolaceae in comparison with the glucosinolate-producing taxa and the Sapindales. The combination of an androecium with eight stamens and oblique monosymmetry is either a single event in evolution and links Tropaeolum with the Sapindales, or it has evolved at least twice, once in the Sapindales, and once in a clade comprising Tropaeolaceae, Akaniaceae and Bretschneideraceae. Morphological data support a sister group relationship of the three latter families, which is in line with macromolecular studies.

A new species, Tropaeolum sparrei Stahl (Tropaeolaceae), is described from submontane cloud forest habitat in western Ecuador. It differs from T. papillosum Hughes (the species it resembles most) by having thinner, tomentose stems, smaller, glabrous or subglabrous leaves with the petiole inserted relatively closer to the lower leaf margin, and flowers with uniformly coloured and straight calyx spurs. In leaf shape it is also similar to T. repandum Heilborn, from which it differs in its larger flowers with entirely black petals and blue anthers.

Trophaeastrum should be reduced to synonyms of Tropaeolum. The two new combinations Tropaeolum porifolium (Cav.) L.Andersson & S.Andersson and T. trialatum (Suess.) L.Andersson & S.Andersson are made. Further, although most of the sections of Tropaeolum that were recognised in Sparre and Andersson's monograph are monophyletic, only two entities can be distinguished that are both monophyletic and morphologically diagnosable. Therefore, the genus Tropaeolum should be subdivided into two sections, T. sect. Tropaeolum and T. sect. Chilensia. Tropaeolum sect. Tropaeolum should include the formerly recognised sections Bicolora, Dipetala, Mucoronata, Schizotrophaeum, Serratociliata, Tropaeolum, and Umbellata. Tropaeolum sect. Chilensia should include the formerly recognised genera Magallana and Trophaeastrum, and the formerly recognised T. sect. Chymocarpus.

Quechua farmers have cultivated mashua (Tropaeolum tuberosum Ruiz & Pavon) and other tuber crops for thousands of years. The practice of trading seed tubers may have contributed to dispersal of viral diseases, such as the tropaeolum mosaic virus (TropMV). We surveyed 17 accessions of mashua collected from Quechua farmers in the provinces of Cuzco and Ayacucho, Peru. Most cross-reacted with the TropMV antibody and showed viral disease symptoms. Significant differences were observed between accessions from Cuzco and Ayacucho, with respect to virus infection and tuber yield under greenhouse conditions. Of the accessions from Cuzco, 87% displayed viral symptoms, while only 22% from Ayacucho showed symptoms. Fewer tubers from Cuzco generated mature plants. In turn, those mature plants produced lower tuber yields. The practice of trading seed tubers may be advantageous for promoting crop diversity but can be harmful when diseased seed tubers are being traded. A program to generate and distribute virus-free seed tubers among Andean farmers would contribute to higher crop yields while preserving local customs and crop diversity.

Ethnopharmacological relevance The use of medicinal plants for the treatment of diseases usually comes from the belief that they present low toxicity due their natural origin. However, it is necessary a toxicological and pharmacological evaluation for these plants. Tropaeolum majus is a medicinal plant used in popular medicine to treat several diseases, including cardiovascular disorders, urinary tract infections and asthma. Even though several studies proved its therapeutic effects, there are few toxicological studies with this species. Aim of the study The present study was carried out to evaluate the subchronic toxicity of the hydroethanolic extract obtained from leaves of T. majus (HETM) in Wistar rats. Material and methods Male and female Wistar rats received three doses of HETM (75, 375 and 750 mg/kg) for 28 days. After the treatments biochemical, hematological and histopathological parameters were analyzed. Results No significant alterations in the animal's body weight gain, relative organs weight, serum biochemical analysis, hematological or histopathological analyses of liver, kidneys and spleen were observed. Conclusions These results demonstrate the absence of subchronic toxicity due to oral treatment with HETM for 28 days in Wistar rats. However, other toxicological studies are necessary to evaluate the total safety of this plant.

Nasturtium (Tropaeolum majus L.) oil contains the highest levels of erucic acid of known seed oils (75–80%). A significant portion of the acid is attached to the 2-position of the glycerol, and trierucin is a major component (ca. 50%) of the oil. Seeds from eleven varieties of commercially available garden nasturtium (T. majus) were screened for oil content, erucic acid levels and fatty acid distribution. Oil contents ranged fromca. 6 to 11%, and erucic acid levels in the oils ranged from 62 to 80%. One sample ofT. speciosum was also analyzed, and contained 28% oil, fatty acids from C16 to C28 and triglycerides up to C72.

Aim of the study Previous studies have shown that the extracts obtained from Tropaeolum majus L. exhibit pronounced diuretic properties. In the present study, we assessed whether the hypotensive and/or antihypertensive mechanism of hydroethanolic extract (HETM), semi-purified fraction (TMLR) obtained from T. majus and the flavonoids isoquercitrin (ISQ) and kaempferol (KPF) can be mediated by their interaction with angiotensin converting enzyme (ACE). Methods and methods Firstly, to evaluate changes in mean arterial pressure (MAP), different groups of normotensive and spontaneously hypertensive rats (SHR) were orally and intraduodenally treated with HETM (10–300 mg/kg) and TMLR (12.5–100 mg/kg) and intravenously treated with ISQ and KPF being later anesthetized with ketamine (100 mg/kg) and xylazine (20 mg/kg). The left femoral vein and the right carotid artery were isolated, and polyethylene catheters were inserted for ISQ and KPF (0.5–4 mg/kg) administration and blood pressure recording, respectively. The plasmatic ACE activity was evaluated to indirect fluorimetry, in serum samples after orally treatment with HETM, TMLR, ISQ and KPF. Results The oral administration of the HETM and its TMLR significantly reduced, in a dose-dependent manner, the MAP in both normotensive and SHR. In addition, these preparations significantly decreased the MAP for up to 3 h after the administration of the extract. Additionally, the intravenous administration of ISQ, but not KPF, decreased MAP in rats. Otherwise, neither the extracts nor ISQ affected the heart rate. The oral administration of the HETM, TMLR or ISQ reduced ACE activity in serum samples at 90 min after administration. Finally, the intravenous administration of ISQ caused a significant reduction in the hypertensive response to angiotensin I, but not angiotensin II in normotensive rats. Conclusion Our results show that the hypotensive effects caused by the HETM, as well as by its TMLR, may be associated with the high levels of the flavonoid ISQ found in this plant. In addition, ISQ-induced hypotension in rats is an event dependent on the inhibition of angiotensin II generation by ACE.