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Tea and coffee are caffeinated beverages commonly consumed around the world in daily life. Tea from Camellia sinensis is widely available and is a good source of caffeine and other bioactive compounds (e.g., polyphenols and carotenoids). Other tea-like beverages, such as those from the genus Ilex, the large-leaved Kudingcha (Ilex latifolia Thunb and Ilex kudingcha C.J. Tseng), Yerba Mate (Ilex paraguariensis A. St.-Hil), Yaupon Holly (Ilex vomitoria), and Guayusa (Ilex guayusa Loes) are also traditional drinks, with lesser overall usage, but have attracted much recent attention and have been subjected to further study. This review summarizes the distribution, composition, and health benefits of caffeinated beverages from the genus Ilex. Plants of this genus mainly contain polyphenols and alkaloids, and show diverse health benefits, which, as well as supporting their further popularization as beverages, may also lead to potential applications in the pharmaceutical or nutraceutical industries.

Questions We addressed the following questions: (1) did defoliation and die-off of the dominant P inus sylvestris, induced by an extreme drought episode, favour emergence of other tree species; (2) did the defoliated canopies of P . sylvestris resulting from drought promote radial growth among other pre-existing tree species seedlings under them? Location P . sylvestris forest in C entral P yrenees ( NE S pain) affected by a severe drought in 2004-2005. Methods Despite increased focus on climate-related forest die-off, studies of the effects on regeneration processes following extreme drought remain scarce. We analysed whether an episode of drought-induced mortality on the dominant P . sylvestris L . may act as a driver of vegetation shift. Seedlings of Q uercus humilis M ill. and Q . ilex L . from 27 plots were sampled under P . sylvestris canopies with <50% and greater than or equal to 50% defoliation (standing dead trees included) to determine age distribution and radial growth using a retrospective, dendrochronological approach. Results Drought-induced canopy losses appear not to be compensated by regeneration of P . sylvestris. Recruitment of below-canopy tree species (specifically Q . humilis and Q . ilex) that could potentially become dominant was high in the entire studied area. However, the spatial patterns of Q uercus spp. regeneration following the 2004-2005 drought were complex. While the emergence of new Q uercus spp. seedlings was reduced under open, drought-induced canopies, growth of seedlings already established was favoured in open-canopy conditions. Conclusions Although the effects of extreme drought events may disfavour the establishment of new recruits, enhanced growth responses of a pre-established seedling bank could still contribute to accelerate forest dynamics under drier conditions. Because of the predicted increases in intensity and frequency of extreme droughts, monitoring studies are key to elucidate whether the initial patterns observed will be maintained in the long term, eventually leading to a vegetation shift. Despite increased focus on climate-related forest dieoff, studies of the effects on regeneration processes following extreme drought remain scarce. Drought-induced canopy losses in a P inus sylvestris L . forest appear not to be compensated by its own regeneration. In contrast, enhanced growth responses of a pre-established seedling bank of Q uercus spp. could still contribute to accelerate forest dynamics under drier conditions.

Chocolate is a globally appreciated product and industry trends point to the search for innovations and demand for healthier products. Similarly, many studies have shown the nutritional benefits of yerba mate (Ilex paraguariensis), including the antioxidant activity attributed to its high content of phenolic compounds, making it a potential raw material for new products development. The aim of this study was the application of yerba mate extract in the white chocolate development for an innovative product that meets the healthiness of consumer needs. Formulations were prepared with 1%, 3% and 5% of yerba mate extract, determined by central composite design 22 for the yerba mate extract and vanillin. All samples showed microbiological standard as required by Brazilian law and were sensory analyzed in order to check which samples were most accepted by consumers. Formulations with 1% and 3% of yerba mate extract were the most accepted for all attributes. Phenolic compounds levels varied from 137.61 to 198.42 mg/100 g EAG and the antioxidant activity varied from 83.39 to 179.73 EC50 (mg/mg). Results showed that the developed product had a good sensory acceptability and that the yerba mate extract added phenolic compounds with antioxidant action to the white chocolate.

Background Nitrogen (N) and phosphorus (P) are essential macronutrients that drive plant growth and photosynthesis; their deficiencies alters both plant physiology and metabolism. However, the molecular mechanisms by which Ilex chinensis responds to N and P starvation remain largely unknown. Results We subjected two-year-old I. chinensis seedlings to 10 weeks of low N (LN) and low P (LP) stress and profiled the leaf transcriptome. Both stresses restricted shoot elongation but stimulated lateral root proliferation, with the strongest phenotype under LN 2 and LP 2 regimes. Relative to the control, LN 2 group exhibited 2.1- to 3.9-fold increases in nitrate reductase (NR), glutamine synthetase (GS), superoxide dismutase (SOD), peroxidase (POD), and malondialdehyde (MDA) ( P  < 0.01). LP 2 group displayed 1.7- to 2.4-fold higher acid phosphatase (ACP), SOD, POD activity, anthocyanin content, and MDA ( P  < 0.01). Transcriptomic analysis revealed that pathways enriched under N and P deficiency were responsive to plant growth, root development, and N and P uptake. Conclusions Our data reveal the integrated physiological and transcriptional adjustments that allow I. chinensis to cope with N and P deficiency stress, and identifies potential target genes for improving nutrient use efficiency. These findings provide new insights into the physiological and molecular responses of I. chinensis to N and P deficiency stress and offer valuable information for optimizing its cultivation under nutrient-limited conditions.

Yerba mate (Ilex paraguariensis A. St.-Hil.) is an important subtropical tree crop cultivated on 326,000 ha in Argentina, Brazil and Paraguay, with a total yield production of more than 1,000,000 t. Yerba mate presents a strong limitation regarding sequence information. The NCBI GenBank lacks an EST database of yerba mate and depicts only 80 DNA sequences, mostly uncharacterized. In this scenario, in order to elucidate the yerba mate gene landscape by means of NGS, we explored and discovered a vast collection of I. paraguariensis transcripts. Total RNA from I. paraguariensis was sequenced by Illumina HiSeq-2000 obtaining 72,031,388 pair-end 100 bp sequences. High quality reads were de novo assembled into 44,907 transcripts encompassing 40 million bases with an estimated coverage of 180X. Multiple sequence analysis allowed us to predict that yerba mate contains ~32,355 genes and 12,551 gene variants or isoforms. We identified and categorized members of more than 100 metabolic pathways. Overall, we have identified ~1,000 putative transcription factors, genes involved in heat and oxidative stress, pathogen response, as well as disease resistance and hormone response. We have also identified, based in sequence homology searches, novel transcripts related to osmotic, drought, salinity and cold stress, senescence and early flowering. We have also pinpointed several members of the gene silencing pathway, and characterized the silencing effector Argonaute1. We predicted a diverse supply of putative microRNA precursors involved in developmental processes. We present here the first draft of the transcribed genomes of the yerba mate chloroplast and mitochondrion. The putative sequence and predicted structure of the caffeine synthase of yerba mate is presented. Moreover, we provide a collection of over 10,800 SSR accessible to the scientific community interested in yerba mate genetic improvement. This contribution broadly expands the limited knowledge of yerba mate genes, and is presented as the first genomic resource of this important crop.

Kuding tea (KT) is a bitter-tasting herbal tea that has been commonly used in traditional Chinese medicine (TCM). The large-leaved Ku-Ding-Cha (Aquifoliaceae) is composed of its representative species Ilex latifolia Thunb and Ilex kudingcha C.J. Tseng. Because of its potential lipid-lowering, body weight-reducing and blood-glucose-lowering properties, KT has increasingly been recognised for its importance over the past several decades. KT is no longer used only as a beverage, and various extraction methods have been applied to obtain aqueous and ethanolic KT extracts (KTE) or their fractions, which could potentially be used as dietary supplements. The major bioactive components of KT are triterpene saponins and polyphenols, but the composition of KT differs substantially between and among the different KT species. This in turn might affect the physiological effects of KT. KT exhibits antiobesity properties, possibly partly by affecting the intestinal microbiota. In addition, KT may mediate putative antioxidative, anti-inflammatory and anticancer activities. However, there is evidence that high KTE supplementation can adversely affect liver metabolism. The physiological relevance of KT in humans remains rather unclear since the potential health benefits of KT and its constituents reviewed here are mainly derived on the basis of in vitro and animal studies.

Taking the extraction amount of rutin as the index, the extraction process of rutin from Ilex asprella (Hook. et Arn.) Champ. ex Benth. using the ultrasonic-assisted deep eutectic solvent method was optimized through response surface method. The effects of various factors, including the types of deep eutectic solvent, molar ratio of the solvent, water content, liquid-solid ratio, the temperature, power and time of ultrasound on the extraction amount of rutin were investigated. In accordance with the result of univariate experiments, the extraction process was optimized through employing the Box-Behnken response surface design method. A three-factor, three-level experimental model was established with the liquid-solid ratio, water content and ultrasonic time as variables. The findings indicated that the optimal technological conditions were as stated below: the molar ratio of lactic acid to choline chloride at 1:1, the ultrasonic extraction temperature at 40 °C, the ultrasonic extraction time at 31 min, the water content at 28%, and the liquid-solid ratio at 20:1 mL/g. Under optimal conditions, the validation experimental result revealed that the extraction amount of rutin was 86.553 ± 1.35 µg/g, with an absolute error of less than 0.6% in comparison with the predicted value of the model. The use of deep eutectic solvents for the extraction of rutin and the application of response surface method for optimizing the extraction process of rutin from Ilex asprella (Hook. et Arn.) Champ. ex Benth. were demonstrated. The present study provided a reference for the comprehensive utilization of Ilex asprella (Hook. et Arn.) Champ. ex Benth. .

Ilex species are important sources of high-quality raw plant materials for the production of drugs and functional foods. The precise identification of different species within the Ilex genus would greatly facilitate authentication and certification as well as forest resource monitoring in plantations. Combining DNA barcoding with high-resolution melting (HRM) analysis represents a robust strategy for species discrimination, as demonstrated in recent DNA barcoding studies. Here, using concatenated and aligned complete chloroplast genomes of different Ilex species, we conducted a sliding window analysis to identify regions of high nucleotide diversity (Pi). We optimized and validated the utility of PCR-based HRM coupled with microsatellite markers to discriminate among the four Ilex species, Ilex integra Thunb., Ilex rotunda Thunb., Ilex cornuta Lindl. and Paxton, and Ilex x wandoensis C.F. Mill and M. Kim, from wild populations in southwestern Korea. The marker trnSUGA-psbZ produced clear melting patterns and distinct melting curve profiles for the four Ilex species using HRM analysis. We applied this protocol to commercially available Ilex accessions and consistently identified the correct species for all 15 accessions tested. Therefore, combining DNA barcoding with HRM analysis is a powerful method for identifying different species within the same genus, which could be used for quality control of raw materials in the functional food/medicinal plant industry.

The cut branches of Ilex verticillata are highly ornamental and have high economic value. Since its introduction to China, it has received widespread attention. In the context of climate change today, ensuring its promotion and sustainable production in China is of great significance. In this study we evaluated the habitat suitability of the species using MaxEnt, combined with climate and soil variables, to assess the impact of climate change on its potential suitable habitat. We used 121 I . verticillata occurrence data and validated the model prediction using extensive field testing (12 test sites located in areas from 23.19° N to 42.91° N and 76.17° E to 125.14° E). The habitat suitability model (AUC = 0.854) performed well. Among them, three precipitation variables and one temperature variable were the main factors determining the distribution of I . verticillata in China. Field trial tests and model predictions of the suitability of I . verticillata were consistent, indicating that our model predictions are biologically meaningful and economically valuable. Under the Shared Socioeconomic Pathways (SSPs) climate change scenario, the high and medium suitable habitats for this species will be reduced in the future climate. This study helps to better understand the impact of climate change on I . verticillata and provides suggestions for the introduction and cultivation areas and protection of this species in China.

It is not clear that the genes involved with flavonoids synthesis, regulation and transport in Ilex asprella. Transcriptome analysis of leaf, stem and root has uncovered 28,478 differentially expressed genes (DEGs) that are involved in various biological processes. Among these, the expression of 31 candidate synthetase genes, 19 transcription factors, and 5 transporters associated with flavonoid biosynthesis varies across tissues, encompassing seven complete biosynthetic pathways (stilbene, aurone, flavone, isoflavone, flavonol, phlobaphene, and anthocyanin) and one partial pathway (proanthocyanidin). Tissue-specific expression patterns suggest that the stilbenes, aurones, flavones and anthocyanin branches are more prominent in roots, as indicated by key genes such as STS(Ilex_044726) , CH4 ʹ GT(Ilex_047989) , FNS(Ilex_043640) and UFGT(Ilex_014720) . In leaves, the phlobaphenes and flavonols branches are dominant, determined by CHI(Ilex_005941) , FNR(Ilex_039777) and FLS(Ilex_046424) . The isoflavone pathway appears to be more active in stems due to the presence of IFS(Ilex_029360) , mirroring the accumulation of the intermediate metabolite chalcone, which is regulated by CHS(Ilex_047537) . The absence of LAR genes implies that gallocatechin, and catechin liked proanthocyanidins cannot be synthesized in I. asprella . Meanwhile, the general phenylpropanoid pathway is more active in roots, stems than in leaves, as evidenced by the expression of PAL(Ilex_042231, Ilex_014816) , C4H(Ilex_017598) , and 4CL(Ilex_042033) . Flavanone, dihydroflavonol and leucoanthocyanidin, key intermediates, accumulate more rapidly in stem, stem and root, respectively, regulated by CHI(Ilex_005941) , F3H(Ilex_004635) and DFR(Ilex_004771) . Correlation and network analyses reveal that candidate regulators and transporters are closely associated with the synthesis genes. The study provides profound snoop into flavonoids metabolism in I. asprella and offers valuable refer for medicinal plant.