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Affective disorders are frequently associated with disrupted circadian rhythms. The existence of rhythmic secretion of central serotonin (5-hydroxytryptamine, 5-HT) pattern has been reported; however, the functional mechanism underlying the circadian control of 5-HTergic mood regulation remains largely unknown. Here, we investigate the role of the circadian nuclear receptor REV-ERBα in regulating tryptophan hydroxylase 2 (Tph2), the rate-limiting enzyme of 5-HT synthesis. We demonstrate that the REV-ERBα expressed in dorsal raphe (DR) 5-HTergic neurons functionally competes with PET-1-a nuclear activator crucial for 5-HTergic neuron development. In mice, genetic ablation of DR 5-HTergic REV-ERBα increases Tph2 expression, leading to elevated DR 5-HT levels and reduced depression-like behaviors at dusk. Further, pharmacological manipulation of the mice DR REV-ERBα activity increases DR 5-HT levels and affects despair-related behaviors. Our findings provide valuable insights into the molecular and cellular link between the circadian rhythm and the mood-controlling DR 5-HTergic systems.
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Ritmo Circadiano , Núcleo Dorsal del Rafe , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares , Serotonina , Triptófano Hidroxilasa , Animales , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares/metabolismo , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares/genética , Núcleo Dorsal del Rafe/metabolismo , Serotonina/metabolismo , Serotonina/biosíntesis , Triptófano Hidroxilasa/metabolismo , Triptófano Hidroxilasa/genética , Ratones , Masculino , Afecto/fisiología , Ratones Noqueados , Ratones Endogámicos C57BL , Factores de Transcripción/metabolismo , Factores de Transcripción/genética , Depresión/metabolismoRESUMEN
This study aims to formulate a stable high internal phase double emulsion (HIPDE) using soybean protein isolate (SPI), gallic acid (GA), and xanthan gum (XG). To prepare HIPDE, W1/O was formulated with the water phase dispersed in the oil phase using polyglycerol polyricinoleate (PGPR) as a stabilizer. Thereafter, W1/O dispersed in W2 (SPI solution) was used. To stabilize the HIPDE, GA was added in W1 (0 or 1 %), XG was added in W2 (0 or 1 %), and the pH of the W phases was adjusted to acidic, neutral, and basic. The samples containing GA in W1 and XG in W2 did not phase out during the storage periods and maintained a higher ζ-potential value, a higher apparent viscosity, and a more sustainable droplet compared to others. These results were derived by the interaction between SPI and XG, SPI and GA, or GA and PGPR. Physicochemical crosslinks were formed, such as gallate-derived groups, SPI-GA complexation (Michael addition, Shiff base reaction), and hydrogen bonding. In conclusion, applying the SPI, GA, and XG to HIPDE would contribute to various industries such as food, medicine, and cosmetics.
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Ácido Gálico , Proteínas de Soja , Emulsiones/química , Polisacáridos Bacterianos/química , Agua/químicaRESUMEN
The increasing abundance of nanoplastics in the environment is a cause of serious concern and its acute and chronic effects on ecosystems need to be thoroughly investigated. Toward this end, this study investigated the parental transfer of nanoplastics by chronically exposing Pisum sativum (pea) plants to nanoplastics through soil medium. We observed the presence of nanoplastics in harvested fruits and a subsequent generation of plants replanted in uncontaminated soil using confocal laser scanning microscopy. The fluorescence was located in the cell wall of the vascular bundles, but not in the epidermis, indicating the parental transfer of nanoplastics. In addition, we determined the effects of nanoplastics on the health of subsequent plant generations by estimating the reproductive factors and measuring the content of individual nutrients in peas. Decreases in crop yield and fruit biomass, in addition to changes in nutrient content and composition, were noted. The transgenerational effects of nanoplastics on plants can profoundly impact terrestrial ecosystems, including both plant species and their predators, raising critical safety concerns. Our findings highlight the evidence of parental transfer of nanoplastics in the soil through plants and shows that the chronic effects of nanoplastics on plants may pose a threat to the food supply.
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Microplásticos , Pisum sativum , Microplásticos/farmacología , Ecosistema , Raíces de Plantas , Plantas , SueloRESUMEN
The addition of various oils to meat analog has been an important topic to improving its juiciness and tenderness. However, this causes a concern about oil leaching from the meat analog during long-term storage. Here, we aimed to assess the storage stability of vegetable-oil-supplemented meat analogs and analyze the effects of temperature and storage period on their physiochemical characteristics. The meat analogs were prepared by adding 30 g castor oil, orange oil, palm oil, shortening, or margarine vegetable oil based on 100 g of textured vegetable protein. They were then stored at -18 or -60 °C for 6 months and analyzed at one-month intervals. The meat analog supplemented with orange oil had the highest water content (64.85%; 66.07%), hardness (35.48 N; 34.05 N), and DPPH-radical-scavenging activity (30.01%; 30.87%) under -18 and -60 °C, respectively, as well as the highest liquid-holding capacity in different conditions. During frozen storage, temperature barely affected the meat quality. The storage stability of all meat analog samples was maintained for 6 months, although the quality was slightly reduced with an increase in storage duration. Coliform group bacteria were not detected regardless of the storage condition. In conclusion, all results supposed that orange oil can be a promising candidate for improving the juiciness and tenderness of meat analogs, and the quality of samples was maintained for at least 6 months under frozen storage. The findings of this study are relevant to the development and promotion of meat analog as an alternative to animal meat.
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This study investigated the tenderizing and flavor-enhancing effects of koji, a fermented grain cultured with a single microorganism, on chicken breasts during curing. Chicken breasts were cured with different ingredients, including 4% (w/w) curing agent (GC), 5% (w/w) Aspergillus oryzae with rice (FR), A. oryzae with soybean (FS), and Bacillus subtilis with soybean (BS) for 4 h at 4 °C prior to cooking. After the superheated steam procedure, all samples were cooked in a convection oven, and their physicochemical properties were analyzed. Koji-treated samples exhibited significantly higher expressible moisture due to the degradation of the protein matrix (p < 0.05). Texture profile analysis showed that the tenderness of koji-treated samples was significantly higher than that of GC (p < 0.05). Furthermore, koji-treated samples were regarded as tenderer, and they were preferred over GC (p < 0.05) in the sensory evaluation. Principal attributes analysis revealed that the overall preference for koji-treated samples was highly correlated with umami, juiciness, and tenderness (p < 0.05). Overall, this study provides insights into applying koji as a potential curing treatment to improve the eating quality of chicken breasts. Koji can be used as a novel technology in the food industry to improve taste and tenderness simultaneously.
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The aim of this study is to formulate a stable water-in-oil-in-water (W1/O/W2) double emulsion using different types of oils and electrolytes. W1/O was formulated with different electrolyte solutions (W1) dispersed in various oils (O) using polyglycerol polyricinoleate as a stabilizer. External aqueous phase was Tween-80 (W2), and W1/O dispersed in W2 was used. The emulsion containing NaCl or MgCl2 exhibited high encapsulation efficiency (EE) and maintained particle size. Regarding the oil type, the emulsion with MCT oil showed a small droplet size and a high viscosity and EE, presenting a stable droplet distribution in optical observation. The stability of emulsion containing NaCl was maintained during the in vitro digestion experiments. MCT oil, NaCl and MgCl2 have the potential to produce stable double emulsions for storage stability and in vitro digestion studies. The findings would be useful for preparing stable double emulsions used in the food and cosmetic industries.
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Social interaction among conspecifics is essential for maintaining adaptive, cooperative, and social behaviors, along with survival among mammals. The 5-hydroxytryptamine (5-HT) neuronal system is an important neurotransmitter system for regulating social behaviors; however, the circadian role of 5-HT in social interaction behaviors is unclear. To investigate whether the circadian nuclear receptor REV-ERBα, a transcriptional repressor of the rate-limiting enzyme tryptophan hydroxylase 2 (Tph2) gene in 5-HT biosynthesis, may affect social interaction behaviors, we generated a conditional knockout (cKO) mouse by targeting Rev-Erbα in dorsal raphe (DR) 5-HT neurons (5-HTDR-specific REV-ERBα cKO) using the CRISPR/Cas9 gene editing system and assayed social behaviors, including social preference and social recognition, with a three-chamber social interaction test at two circadian time (CT) points, i.e., at dawn (CT00) and dusk (CT12). The genetic ablation of Rev-Erbα in DR 5-HTergic neurons caused impaired social interaction behaviors, particularly social preference but not social recognition, with no difference between the two CT points. This deficit of social preference induced by Rev-Erbα in 5-HTDR-specific mice is functionally associated with real-time elevated neuron activity and 5-HT levels at dusk, as determined by fiber-photometry imaging sensors. Moreover, optogenetic inhibition of DR to nucleus accumbens (NAc) 5-HTergic circuit restored the impairment of social preference in 5-HTDR-specific REV-ERBα cKO mice. These results suggest the significance of the circadian regulation of 5-HT levels by REV-ERBα in regulating social interaction behaviors.
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Ritmo Circadiano , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares , Conducta Social , Animales , Ratones , Ritmo Circadiano/genética , Núcleo Dorsal del Rafe/metabolismo , Mamíferos/metabolismo , Ratones Noqueados , Neuronas/metabolismo , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares/genética , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares/metabolismo , Serotonina , Interacción SocialRESUMEN
Fruits and vegetables are a vital source of redox-active phytochemicals in the diet. Traditional green leafy vegetables (GLVs) are a rich source of carotenoids, dietary fiber, minerals, phenols, vitamins, and tocopherols and are commonly consumed in rural areas worldwide. In traditional Korean medicine, many GLVs are used to treat various ailments. However, data on the carotenoid and tocopherol content of many traditional GLVs consumed in the Republic of Korea are insufficient. The current work aims to compare the carotenoid and tocopherol profiles of 18 traditional GLVs by utilizing a single ion monitoring LC-MS approach to identify the potential GLVs for commercial cultivation and healthy diet formulations. Among the traditional GLVs investigated, (all-E)-lutein was the most abundant carotenoid, ranging from 44.4% in Glehnia littoralis to 52.1% in Heracleum moellendorffii. It was followed by (all-E)-violaxanthin and (all-E)-ß-carotene. The highest contents of (all-E)-violaxanthin (75.6 µg/g FW), 9-Z-neoxanthin (48.4 µg/g FW), (all-E)-luteoxanthin (10.8 µg/g FW), (all-E)-lutein (174.1 µg/g FW), total xanthophylls (310.5 µg/g FW), (all-E)-ß-carotene (69.6 µg/g FW), and total carotenoids (380.1 µg/g FW) were recorded in Pimpinella brachycarpa. Surprisingly, Taraxacum mongolicum also showed the highest contents of (all-E)-violaxanthin, (all-E)-lutein, and total carotenoids, which were statistically non-significant (p > 0.05, Tukey HSD) with P. brachycarpa. The highest concentration of (all-E)-zeaxanthin (14.4 µg/g FW) was recorded in Solidago virga-aurea. Among the studied herbs, 13.9 (H. moellendorffii)-133.6 µg/g FW (Toona sinensis) of α-tocopherol was recorded. Overall, the results suggest that P. brachycarpa and T. mongolicum are rich sources of carotenoids. On the other hand, T. sinensis is a rich source of α-tocopherol. These GLVs can be utilized in the diet to enhance the intake of health-beneficial carotenoids and α-tocopherol.
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Freezing raw laver is unsuitable for the laver industry due to process characteristics and economic problems. Therefore, this study attempted to investigate supercooled storage to extend the storage period without freezing, rather than refrigeration. To compare and analyze the storage ability of supercooling, the experiment was performed under refrigeration (5 °C), constant supercooling (CS, -2 °C), stepwise supercooling (SS, -2 °C), and freezing (-18 °C) conditions for 15 days, and the physicochemical changes according to the treatment and period were investigated. All SS samples, which were designed for stable supercooling, were kept in a supercooled state for 15 days. Two samples among the twelve total subjected to CS were frozen. At 9 days, the drip losses of the CS and SS samples were 6.32% and 6.48%, respectively, which was two times lower than that of refrigeration and three times lower than that of the frozen samples. The VBN of the refrigerated samples was 108.33 mg/100 g at 6 days, which exceeded the decomposition criterion. Simultaneously, the VBN of the other treatments was under the decomposition criterion of 30 mg/100 g. However, the VBN of both supercooling samples at 15 days increased to higher than the decomposition criterion. Regarding appearance, the refrigerated samples showed tissue destruction at 9 days, but tissue destruction of the CS and CC samples was observed at 15 days, and tissue destruction of the frozen samples was not observed until 15 days. Consequently, supercooling did not maintain quality for longer periods than freezing, but it did extend the shelf life more than refrigeration, and effectively preserved the quality for a short period.
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This study identified the effect of the type and concentration of vegetable oil on the quality of meat analogs and analyzed the differences in their physiochemical characteristics. Various vegetable oils, such as castor oil, orange oil, palm oil, shortening, and margarine, were added to meat analogs. The meat analog was prepared by adding 10, 20, 30, 40, and 50 g of each vegetable oil based on 100 g of textured vegetable protein. The cooking loss, water content, liquid-holding capacity, texture, and antioxidant content of the meat analogs were assessed, and a sensory evaluation was performed. The meat analog with orange oil had a higher water content than the others, regardless of the amount of added oil, and it had a relatively high liquid-holding capacity. The DPPH(2,2-diphenyl-1-picrylhydrazyl) radical scavenging activity of the meat analog with orange oil was higher than that of the others. The sensory evaluation also showed a decrease in soy odor and an increase in juiciness. Therefore, adding orange oil improves the preference, juiciness, soy odor, and quality of meat analogs. Our results demonstrate that orange oil has positive effects on the productivity of meat analogs and can help to improve meat analog consumption.
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This study determined the effects of physicochemical and microbial properties of emulsion as a fat replacer in meat analogs during freezing storage. Meat analogs were prepared with different fat replacers: vegetable oil (O) for control, oil in water emulsion (E), and non-emulsified oil in water emulsion (EC) for emulsion control. After that, meat analogs were stored for 0.5, one, three, and six months at −18 °C and −60 °C. The results showed that the drip loss of all samples was not significantly different (p > 0.05). However, the liquid holding capacity of EC and E was significantly higher than that of O (p < 0.05). Additionally, the microstructures of meat analogs of E and EC were smaller with denser pore sizes than O. This explains the significantly lower hardness of E and EC compared to O (p < 0.05). Overall, E showed superior physiochemical and sensory quality. During the storage, the stability of chemical properties, such as volatile basic nitrogen and thiobarbituric acid reactive substances, showed no significant changes (p > 0.05). Moreover, the microbial studies (total viable counts and Escherichia coli count) suggested that meat analogs did not deteriorate during the preparation and storage. Thus, this study suggests that emulsion-type fat replacers influence meat analogs' physicochemical and sensorial properties. However, these properties are not influenced by the storage temperature and duration.
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Supercooling has the advantage of maintaining the freshness of foods without a phase transition. However, it is hard to sustain the supercooled state. Static temperature control, one of the various supercooling technologies, is used for stable supercooling storage. In this experiment, the effect of following external factors in maintaining the supercooled state of foods was investigated. Three main parameters had an effect on the supercooled state of food: (1) properly setting the lower-temperature limit of the supercooling algorithm, (2) slow cooling to the target temperature, and (3) minimizing temperature fluctuation. Accordingly, the following stepwise cooling algorithm for pork loin was designed: a lower-temperature limit of −3.0 °C and a storage period = 36 h followed by a lower-temperature limit of −3.5 °C for 24 h. The samples conserved at −3.0 °C displayed a 100% supercooled state. Physicochemical properties including drip loss, cooking loss, texture, color, total volatile basic nitrogen (TVBN), and total aerobic count (TAC) of pork loin were analyzed. The drip loss values of the supercooled meat samples were lower than those of the superchilled ones. Furthermore, TVBN and TAC of the treated samples were not significantly different from those of the fresh samples (p > 0.05). In conclusion, supercooling storage extended the freshness and quality of pork loin better than refrigerated storage.
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Due to growing interest in health and sustainability, the demand for replacing animal-based ingredients with more sustainable alternatives has increased. Many studies have been conducted on plant-based meat, but only a few have investigated the effect of adding a suitable binder to plant-based meat to enhance meat texture. Thus, this study investigated the effects of the addition of transglutaminase (TG) and glucono-δ-lactone (GdL) on the physicochemical, textural, and sensory characteristics of plant-based ground meat products. The addition of a high quantity of GdL(G10T0) had an effect on the decrease in lightness (L* 58.98) and the increase in redness (a* 3.62). TG and GdL also decreased in terms of cooking loss (CL) and water holding capacity (WHC) of PBMPs. G5T5 showed the lowest CL (3.8%), while G3T7 showed the lowest WHC (86.02%). The mechanical properties also confirmed that G3T7-added patties have significantly high hardness (25.49 N), springiness (3.7 mm), gumminess (15.99 N), and chewiness (57.76 mJ). The improved textural properties can compensate for the chewability of PBMPs. Although the overall preference for improved hardness was not high compared to the control in the sensory test, these results provide a new direction for improving the textural properties of plant-based meat by using binders and forming fibrous structures.
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Microplastics (MPs) released into soil environments, along with the existing pollutants in soil, may have adverse effects on plants. However, the chronic effects of MPs in soils contaminated with heavy metals on crop plants remain unidentified. This study aimed to investigate the chronic effects of MPs (polystyrene, 20 nm) on the reproductive and nutritional status of pea crop plant (Pisum sativum) grown in Cu- (40 mg/kg) and MP-contaminated soils (40, 20 mg/kg). The crop yield reduced in all groups, with an evident decrease in the complex exposure group (comprising MPs and Cu). Moreover, significant changes in plants were identified regarding the weight, color, amino acids, and protein content of peas. Nutrient content in beans increased by MP exposure in single and complex exposure groups. Cu accumulation did not differ in the presence and absence of MPs. Additionally, MPs that infiltrated into incomplete casparian strips during root formation translocated into aerial parts via the apoplast pathway along the cell walls of the vascular bundle. Therefore, long-term exposure to MPs in soil can significantly affect plants while collective application of Cu and MPs imposed severe damage. The changes in the crop quality and nutrient contents may in turn affect human health through the food chain.
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Pisum sativum , Contaminantes del Suelo , Cobre/metabolismo , Humanos , Microplásticos , Pisum sativum/metabolismo , Plantas/metabolismo , Plásticos/metabolismo , Suelo , Contaminantes del Suelo/análisisRESUMEN
Supercooling storage refers to lowering the product temperature below its freezing point without phase transition and has the potential to extend shelf life. Nevertheless, supercooled objects are in a thermodynamically unstable state, and nucleation can occur spontaneously. To achieve supercooling storage, slow cooling and insulation are essential. Hence, a stepwise algorithm for the supercooling storage of pork loins was designed and validated in this study. Pork loins were stored at 3°C, -18°C, and -3°C (freezing), and supercooled for 16 days. All samples remained in a supercooled state and were unfrozen at the end of storage. Supercooled pork loins were superior in terms of drip loss, cooking loss, and water-holding capacity compared to frozen samples. Additionally, supercooling treatment prevented discoloration, increase of volatile basic nitrogen, and microbial growth. Thus, supercooling of pork loin was achieved using a stepwise program and was effective to maintain meat quality.
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Parkinson's disease is a neurodegenerative disease characterized by progressive dopaminergic neuronal loss. Motor deficits experienced by patients with Parkinson's disease are well documented, but non-motor symptoms, including mood disorders associated with circadian disturbances, are also frequent features. One common phenomenon is "sundowning syndrome," which is characterized by the occurrence of neuropsychiatric symptoms at a specific time (dusk), causing severe quality of life challenges. This study aimed to elucidate the underlying mechanisms of sundowning syndrome in Parkinson's disease and their molecular links with the circadian clock. We demonstrated that 6-hydroxydopamine (6-OHDA)-lesioned mice, as Parkinson's disease mouse model, exhibit increased depression- and anxiety-like behaviors only at dawn (the equivalent of dusk in human). Administration of REV-ERBα antagonist, SR8278, exerted antidepressant and anxiolytic effects in a circadian time-dependent manner in 6-OHDA-lesioned mice and restored the circadian rhythm of mood-related behaviors. 6-OHDA-lesion altered DAergic-specific Rev-erbα and Nurr1 transcription, and atypical binding activities of REV-ERBα and NURR1, which are upstream nuclear receptors for the discrete tyrosine hydroxylase promoter region. SR8278 treatment restored the binding activities of REV-ERBα and NURR1 to the tyrosine hydroxylase promoter and the induction of enrichment of the R/N motif, recognized by REV-ERBα and NURR1, as revealed by ATAC-sequencing; therefore, tyrosine hydroxylase expression was elevated in the ventral tegmental area of 6-OHDA-injected mice, especially at dawn. These results indicate that REV-ERBα is a potential therapeutic target, and its antagonist, SR8278, is a potential drug for mood disorders related to circadian disturbances, namely sundowning syndrome, in Parkinson's disease.
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Enfermedades Neurodegenerativas , Enfermedad de Parkinson , Animales , Humanos , Isoquinolinas , Ratones , Trastornos del Humor/tratamiento farmacológico , Trastornos del Humor/etiología , Miembro 1 del Grupo D de la Subfamilia 1 de Receptores Nucleares/metabolismo , Oxidopamina/toxicidad , Enfermedad de Parkinson/patología , Calidad de Vida , Tiofenos , Tirosina 3-Monooxigenasa/metabolismoRESUMEN
Supercooling is the method of lowering the temperature of a foodstuff below its freezing point without phase transitions. This storage technique has a potential advantage for extending shelf life. Nevertheless, the supercooled state of food is thermodynamically unstable. To accomplish supercooling storage, slow cooling rate and minimized temperature fluctuation are necessary. Thus, a stepwise cooling algorithm was designed and applied in this study. Pork belly and chicken breast were stored at 3 °C, -18 °C (freezing), and supercooling treatment was applied to them for 12 days. All samples preserved their supercooled state and were unfrozen during the storage period. Overall, supercooled samples were advantageous in respect of drip loss compared to that of frozen samples, regardless of type of sample. Total volatile basic nitrogen, total aerobic account, and cooking loss of pork belly was higher than in the chicken breast due to the high fat retention in pork belly as compared to chicken breast, in particular, at refrigerated storage condition. Samples stored at supercooling treatment prevented increase in volatile basic nitrogen and microbial growth. Therefore, the supercooled state was successful when using stepwise algorithm, and it was effective at maintaining meat quality compared to freezing and refrigeration storage.
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Probiotic extracts have various positive attributes, such as antioxidant, tyrosinase inhibitory, and antimicrobial activity. Lactobacillus curvatus produces bacteriocin, which activates the lipid membrane structure and has potential as a natural preservative for cosmetic emulsions. In this study, L. curvatus extract was encapsulated in liposomes and formulated as an oil-in-water (O/W) emulsion. Radical scavenging activity, tyrosinase inhibition, and challenge tests were conducted to confirm the liposome activity and the activity of the applied lotion emulsion. The liposome-encapsulated extract had a relatively high absolute ζ-potential (52.53 > 35.43), indicating its stability, and 96% permeability, which indicates its potential as an active agent in lotion emulsions. Characterization of emulsions containing the liposomes also indicated a stable state. The liposome-encapsulated extract exhibited a higher radical scavenging activity than samples without the extract and non-encapsulated samples, and the functionality was preserved in the lotion emulsion. The tyrosinase inhibition activity of the lotion emulsion with the liposome-encapsulated extract was similar to that of the non-treated extract. Candida albicans and Aspergillus niger were also inhibited in the challenge test with the lotion emulsions during storage. Collectively, these findings indicate that the liposome-encapsulated extract and the lotion containing the encapsulated extract have potential applicability as natural preservatives.
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The genus Hosta, which has a native distribution in temperate East Asia and a number of species ranging from 23 to 40, represents a taxonomically important and ornamentally popular plant. Despite its taxonomic and horticultural importance, the genus Hosta has remained taxonomically challenging owing to insufficient diagnostic features, continuous morphological variation, and the process of hybridization and introgression, making species circumscription and phylogenetic inference difficult. In this study, we sequenced 11 accessions of Hosta plastomes, including members of three geographically defined subgenera, Hosta, Bryocles, and Giboshi, determined the characteristics of plastomes, and inferred their phylogenetic relationships. We found highly conserved plastomes among the three subgenera, identified several mutation hotspots that can be used as barcodes, and revealed the patterns of codon usage bias and RNA editing sites. Five positively selected plastome genes (rbcL, rpoB, rpoC2, rpl16, and rpl20) were identified. Phylogenetic analysis suggested (1) the earliest divergence of subg. Hosta, (2) non-monophyly of subg. Bryocles and its two sections (Lamellatae and Stoloniferae), (3) a sister relationship between H. sieboldiana (subg. Giboshi) and H. ventricosa (subg. Bryocles), and (4) reciprocally monophyletic and divergent lineages of H. capitata in Korea and Japan, requiring further studies of their taxonomic distinction.