Agrimonia pilosa ledeb, a plant widely distributed in Asia and Europe, is rich in secondary metabolites. It is commonly processed into tea using tender shoots in spring or ustilized as an herb in autumn. In the present study, a total of twenty terpenes were identified in A. pilosa volatile compounds, with distinct accumulation patterns observed across different tissues of tender seedlings and mature plants. Several terpenes, including β-myrcene, β-ocimene, linalool and nerolidol were detected exclusively in leaves of tender seedlings. In mature plants, eight terpenes, including β-myrcene, β-farnesene and cedr-8-ene were emitted solely by flowers, while caryophyllene was detected only in leaves. Eleven terpene synthases (TPSs) belonging to TPS-a, b and g subfamilies, were dug out from the transcriptomes of A. pilosa flowers and leaves, designated as ApTPS1 through ApTPS11. Three TPS members—ApTPS1, ApTPS2, and ApTPS3—were identified as bifunctional synthases, capable of synthesizing both monoterpenes and sesquiterpenes in vitro. ApTPS5 and ApTPS7 were implicated in monoterpene and sesquiterpene biosynthesis, respectively. The expression levels of ApTPS1, ApTPS2, ApTPS5, ApTPS6 and ApTPS8 showed a strong correlation with the accumulation of terpenes, including cedr-8-ene, β-farnesene, α-pinene, β-pinene, caryophyllene and α-maaliene. These findings suggest that the variation in ApTPSs expression play a crucial role in mediating terpenes accumulation. Terpenes metabolic profiling exhibited a tissue-specific accumulation in Agrimonia pilosa. The terpenes in A. pilosa exhibited seasonal variation, with higher content and greater diversity observed in tender seedlings compared to mature plants. Five ApTPSs were proved to produce the major terpenes detected in the volatile compounds of A. pilosa.

Water lilies are among the most basal groups of angiosperms and retain many morphological and physiological traits of early angiosperms, making them invaluable for studying angiosperm evolution, particularly floral organ development. Here we present the most comprehensive phylogeny of the genus Nymphaea to date, alongside gap-free genome assemblies for three species (Nymphaea colorata, Nymphaea thermarum and Nymphaea caerulea). Our analyses resolve 2 major clades, day-flowering (section A) and night-flowering (section B), which diverged approximately 50 million years ago. Comparative genomics reveals an angiosperm-exclusive pectin lyase gene specifically expressed during pollen tube elongation. Regarding floral traits, we identify the transcription factor NcolMYB75-like as a master regulator of blue anthocyanin biosynthesis. Furthermore, the expansion and diversification of the O-methyltransferase gene family drive the synthesis of species-specific floral scent volatiles. These findings deepen our understanding of early angiosperm innovations and provide a genomic framework for plant breeding and ecological conservation.

Understanding complex taste perception requires dissecting multi-molecular interactions at the receptor level. This study integrates metabolomics, peptidomics, and in silico molecular docking to construct a multi-layered chemical model of umami in six premium green teas. Our analysis revealed distinct flavor profiles driven by divergent chemical strategies: an amino acid-dominant profile (e.g., high theanine at 21.82 mg/g in GT4) versus a synergistic nucleotide-enhanced profile (e.g., driven by adenosine-5'-monophosphate in GT3). Peptidomics identified 315 unique water-soluble peptides, establishing distinct flavor fingerprints contributing to umami and kokumi sensations. Notably, we characterized novel theanine-substituted peptides, and identified abundant glycosylated peptides (e.g., in GT6) as potential Maillard reaction precursors, hypothetically linking thermal processing to distinct roasted flavor phenotypes. Furthermore, synergistic molecular docking models of the T1R1 receptor visually elucidated the multi-ligand binding mechanisms involving peptides, amino acids (with theanine exhibiting the highest affinity at -3.57 kcal·mol-1), nucleotides, and synergistic co-ligands. This comprehensive framework bridges the gap between macroscopic chemical fingerprints and specific molecular receptor interactions, providing novel insights into tea flavor science.

Intercropping is a traditional and sustainable agricultural practice that enhances soil fertility, reduces erosion, and promotes plant growth and metabolism. Despite widespread adoption in tea plantations, intercropping effects on yield, quality, and soil are highly context-dependent, varying with species choice, climate, and site-specific conditions. Using 157 paired observations from 41 articles, we performed a meta-analysis to quantify how intercropping affects tea yield, quality, and soil, and to determine the primary modulating factors. Across all included reports, intercropping significantly increased tea yield by 11.9%. It also elevated the levels of free amino acids (AAs) by 19.08% and reduced the levels of tea polyphenols (TPs) by 7.16%, leading to a notable decrease in the TP/AA ratio, which is associated with tea quality. Additionally, intercropping increased soil nutrient levels and soil pH. Initial organic matter and pH had a significant influence on tea yield and quality outcomes, while the type of plants used for intercropping and climatic factors such as mean annual temperature (MAT) and mean annual precipitation (MAP) were closely associated with improvements in soil quality. Overall, intercropping can improve tea yield and quality while enhancing soil conditions in tea plantations, although the magnitude of these benefits depends on companion crop selection, soil status, and climate.

The origin and diversification of angiosperms, particularly their ancestral ecological niche, remain an enduring mystery. While competing hypotheses exist, the genomic mechanisms driving repeated transitions to aquatic life are poorly understood. We assembled chromosome-scale genomes for three early-diverging water lilies-Cabomba caroliniana, Barclaya longifolia, and Brasenia schreberi-integrating them with other Nymphaeales genomes for a comparative analysis. Our findings reveal that aquatic adaptation was shaped by ancient whole-genome duplications and functional remodeling of key gene modules. This includes the expansion of photosynthesis and stress-related families, the contraction of nucleotide-binding leucine-rich repeat (NLR) immune receptors, and metabolic reprogramming of terpene-squalene and other stress-associated secondary pathways. These modifications enhanced energy capture, immunity modulation, and resource efficiency in aquatic environments. Collectively, our results underscore the importance of secondary aquatic adaptation in early water lilies, demonstrating that their distinctive aquatic traits can be parsimoniously explained by evolutionary changes associated with adaptation to submerged habitats.

Longjing tea is well-known for its exceptional umami/mellow flavor, but the complexity of interactions among chemical compositions has impeded in-depth understanding. This study comprehensively compared the taste contributors in 'Longjing 43' (LJ43) and 'Qunti' (QT). Peptidomics revealed 865 identified water-soluble peptides in QT, and 497 in LJ43, with 44 umami peptides predicted. Potential umami peptide-T1R1/T1R3 complexes were further modeled using AlphaFold 3. LJ43 had higher levels of theanine (>18 mg/g), and glutamine (similar to 4 mg/g) compared to QT. Similar amounts of flavan-3-ols (>150 mg/g), particularly EGCG (>60 mg/g) and ECG (>30 mg/g) were determined in LJ43 and QT. Moreover, Longjing teas' characterized flavor contributions by macromolecules (peptides), and small molecules (amino acids, catechins, saccharides, and Maillard reaction products) were integrated. These insights will greatly expand tea flavor chemistry and provide promising approaches promotion premium Longjing teas.

DS severely impacts tea plant growth and yield. Foliar application of Methyl jasmonate (MJ) -loaded Chitosan nanoparticles (CNPs) (MJ-CNPs) significantly boost tea DS resistance, necessitating further mechanism exploration. This study comprehensively investigated alterations at the phenotypic, biochemical, and genetic levels. Relative water content, total pigment content, and soluble proteins decreased, while soluble sugar increased significantly under DS and with apparent recovery after MJ-CNPs treatment. Catechins exhibited a significant decrease under DS, especially EGCG (24.6 to 13.4 mg•g-1), but were absolutely mitigated by treatment. Antioxidant capacities (DPPH, FRAP, ABTS, and SOA) showed a further improvement by MJ-CNPs treatment. Endogenous ABA and SA increased under DS, further elevated by MJ-CNPs. CsNCED, CsPYL8, CsPP2C, CsMYB, and CsABF genes involved in ABA-dependent pathways were confirmed with promoted expressions by foliar pre-treatment. The CsJAZ and CsMYC2 gene family which involved in the jasmonic acid (JA) pathway, displayed varied expression patterns. The integration of metabolite and gene expression levels provided a comprehensive illustration of tea DS tolerance mechanism and offered promising promotion strategies through foliar application of MJ-CNPs.

Yellow-leaf tea cultivar ‘Huangjinya’ is valued for distinctive appearance and umami flavor, yet the mechanisms underlying pigment accumulation and flavor formation responding to cultivation and processing remain unclear. Targeted metabolomic quantified 16 carotenoids and 18 flavonoids, revealing significantly higher concentrations in high-altitude site fresh leaves (551.88 ± 7.09 μg·g−1 and 213.91 ± 3.78 mg·g−1, respectively), approximately 2.8-fold and 1.2-fold higher than in low-altitude site samples. These observed regional differences likely reflect environmental differences between the high- and low-altitude cultivation sites. Flavonoids remained relatively stable during green tea (GT) processing but declined markedly during black tea (BT) production. Carotenoids decreased by 27.91% during green tea (GT) processing but increased by 43.64% in black tea (BT) when using low-altitude site fresh leaves; in contrast, high-altitude site leaves showed a 4.66% increase in GT and a sharp 65% decrease in BT. Thirty-eight primary metabolites showed significant changes responding to altitudes and processing, especially amino acids and oligosaccharides. These findings clarify how altitude and processing affect flavor-related metabolism in ‘Huangjinya’, offering a chemical foundation for improving tea quality.

Drought stress (DS) significantly hampers the growth and productivity of tea plants, necessitating effective strategies to enhance their resilience. This study comprehensively investigated the mechanisms of carbonyl volatiles-methyl jasmonate (MeJA) and cis-3-hexenyl acetate (cis-3-HAC) and signal peptide CLAVATA3/EMBRYO-SURROUNDING REGION-RELATED 25 (CLE25) promotion DS resistance using integrative metabolomics and proteomics strategy. Total pigment content decreased, while soluble sugar and proteins increased significantly under DS and further increased after foliar inducement of CLE25, MeJA, and cis-3-HAC. Gallated catechins and amino acids exhibited apparent decreased under DS, especially EGCG (24.6-13.4 mg g-1) and theanine (10.66-3.78 mg g-1), but significantly mitigated by CLE25 inducement. Antioxidant enzymes activity, such as catalase (CAT), jumped from 23.1 to 48.2 and further boosted to 118.8 Ug-1min-1 FW with CLE25. Proteomic analysis revealed massive increased in stress tolerance proteins, particularly dehydrins and heat shock proteins, rising by >50.0 % with CLE25 inducement and the expression levels of peroxidase (POX), superoxide dismutase (SOD), α-galactosidase (α-GAL), carboxypeptidases (CPs), and transaldolase (TAL) exhibited higher after inducement. Furthermore, stress signaling-related proteins were in-depth explored, especially thioredoxin proteins; sucrose non-fermenting 1-related protein kinase 2 (SnRK2) was novelly verified in activating abscisic acid (ABA) responding. Differences among drought resistance mechanisms after carbonyl volatile and CLE25 treatments were comprehensively studied. The integration of metabolite and protein levels provided a comprehensive illustration of tea DS tolerance mechanisms and offered promising promotion strategies through foliar application of MeJA, cis-3-HAC, and CLE25.

Tea pruning litter is frequently added to soil to improve soil fertility and the C pool in tea plantations. However, the effects of different methods of incorporating tea pruning litter, such as direct return to the field, processing to organic fertilizer, and biochar, on soil organic carbon mineralization and its underlying mechanisms, especially microbial mechanisms, are still poorly understood. Therefore, we conducted an incubation experiment to explore the efficacy of tea pruning litter and its derivatives, biochar, and organic fertilizer, on the mineralization and chemical composition of soil organic C (SOC), labile organic C, functional gene abundance, microbial community composition, and activities of enzymes associated with C cycling in tea plantation soil. The results indicated that cumulative soil CO2 emissions were ranked as pruned litter > organic fertilizer > biochar > control soil. The cumulative soil CO2 emissions was decreased significantly by decreasing of the content of O-alkyl C and microbial biomass carbon, beta-glucosidase/cellobiohydrolase activities, and the abundance of GH48 and cbhI (P < 0.05). This indicates that lower CO2 emissions following organic fertilizer and biochar addition (cf. pruned litter) were associated with a decrease in O-alkyl C content, beta-glucosidase and cellobiohydrolase activities, as well as the abundance of GH48 and cbhI. More interestingly, changes in the microbial community structure, especially in some key species, suchas Acidobacteria, Actinobacteria, Sordariomycetes, and Mortierellomycetes, can significantly affect the rate of SOC mineralization. Our study demonstrates that applying organic fertilizer and biochar derived from pruned litter significantly mitigated SOC mineralization compared to pruned litter application alone, and both have great potential for maintaining soil C stock in tea plantation soil.

Albino tea was known for umami and exhibited varying flavors responding to cultivation and processing, necessitating further exploration. In this study, peptidomics and targeted metabolomics unraveled crucial taste related compounds in low-altitude (L) and high-altitude (H) samples. >300 peptides were newly identified via de novo sequencing, with 24 umami and 19 bitter peptides predicted with potential active cores. Total amino acids content was higher in L (31.62 mg/g) than in H (28.35 mg/g), with theanine and glutamic acid accounting for 54.29 % and 2.02 %, compared to 54.95 % and 7.47 % in H, respectively. Lutein and β-carotene were more abundant at L (> 400 μg/g) than at H (∼200 μg/g), and significantly increased during processing. Epigallocatechin gallate reached 65.18 mg/g in L that higher than H (58.66 mg/g). These findings expanded the understanding of flavor peptides in albino tea and will underscore the importance of altitude-specific strategies achieving premium quality.

Variations in cultivars and cultivation altitudes have significant impacts on tea flavour compounds however lack of comprehensive understanding. This study provided insights into differential accumulation of crucial flavour compounds in response to cultivars, cultivation altitudes, and processing. Twelve flavonoids (262.4 ∼ 275.4 mg•g-1) and 20 amino acids (AAs) (56.5 ∼ 64.8 mg•g-1) were comparative analyzed in 'Longjing 43' and 'Qunti' fresh leaves harvested at low (80 m, LA) and high (500 m, HA) altitudes. Additionally, an in-depth correlation unravelling of 31 alkaloids, 25 fatty acids, 31 saccharides, 8 organic acids, and 7 vitamins and flavonoids/AAs during green tea (GT) and black tea (BT) processing was performed. Enhenced flavonoid accumulation alongside higher AAs and saccharides in HA GT promoted a sweet/mellow flavour. Abundant flavonoids, AAs, and saccharides derivates in LA BT gave rise to a sweet aftertaste. The study presents an integrated illustration of major flavour compounds' differential accumulation patterns and their interrelations, providing new insights into the influence of cultivation conditions on tea flavour.

Soil carbon (C) and nitrogen (N) are vital for enhancing tea production and ensuring the sustainability of tea plantation ecosystems. However, research on the dynamics of soil C and N pools and their associated microbial mechanisms in tea plantations with varying cultivation durations is scarce. We compared soil samples from a forest and two tea plantations—young established (YTP) and century-old (OTP)—to assess changes in soil C and N concentrations and the impact of fungal community structure on these elements. Soil organic carbon (SOC) and total nitrogen (TN) were markedly higher in OTP than in the YTP and forest (65.9% and 30.1%, respectively, relative to YTP). Eurotiomycetes in the YTP group accounted for a relatively higher proportion at 51.6%, surpassing its presence in both the forest (14.3%) and OTP (4.78%) groups and it can be the main microbial factor affecting the C cycle in tea plantation soils and facilitating SOC mineralization. Enhancing planting years or changing land use patterns improves fertilizer and biomass sedimentation and increases the relative abundance of Eurotiomycetes in the soil and the C sink potential of tea plantations. This study provides valuable insights into the role of soil C and N dynamics and fungal communities in tea plantation ecosystems, highlighting the importance of managing these factors for sustainable tea production.

Intelligently processed black teas (BT) possess premium quality but there is a lack in comprehensive understanding of flavor formation mechanism. In this study, the accumulation of carotenoids, flavonoids, and Maillard products and (non)enzymatic degradations and conjugations to characterized flavors were comprehensively studied. Significant decrease was observed that flava-3-ols were heavily oxidised from > 240 mgg(-1) in fresh leaves (FL) to < 30 mgg(-1), while other 21 flavonoids decreased by < 30% in BT, accompanied by a sweet aftertaste. Carotenes and xanthophylls, significantly accumulated during withering compared to FL (from 641 +/- 39.7 mu gg(-1) to 728 +/- 44.9 mu gg(-1)) but decreased in BT. Strong correlations were confirmed between the 218 primary metabolites, carotenoids, and flavonoids, and their contributions to BT sweet tastes were elucidated. Furthermore, 45 floral/sweet volatiles with VIP > 1 originating from carotenoids, lipids, and amino acids were screened. An integrated illustration of pigments thermal- and enzymatic-dominated contributions to BT flavour was comprehensively conducted.

Insect-induced plant volatile organic compounds (VOCs) may function as either direct defence molecules to deter insects or indirect defence signals to attract the natural enemies of the invading insects. Tea (Camellia sinensis L.), an important leaf-based beverage crop, is mainly infested by Ectropis obliqua which causes the most serious damage. Here, we report a mechanistic investigation of tea plant-derived VOCs in an indirect defence mechanism against E. obliqua. Parasitoid wasp Parapanteles hyposidrae, a natural enemy of E. obliqua, showed strong electrophysiological response and selection behaviour towards S-linalool and β-ocimene, two monoterpenes with elevated emission from E. obliqua-damaged tea plants. Larvae frass of E. obliqua, which also released S-linalool and β-ocimene, was found to attract both mated female or male Pa. hyposidrae according to gas chromatography-electroantennogram detection and Y-tube olfactometer assays. In a field setting, both S-linalool and β-ocimene were effective in recruiting both female and male Pa. hyposidrae wasps. To understand the molecular mechanism of monoterpenes-mediated indirect defence in tea plants, two novel monoterpene synthase genes, CsLIS and CsOCS-SCZ, involved in the biosynthesis of S-linalool or β-ocimene, respectively, were identified and biochemically characterised. When the expression of these two genes in tea plants was inhibited by antisense oligodeoxynucleotide, both volatile emission and attraction of wasps were reduced. Furthermore, gene expression analysis suggested that the expression of CsLIS and CsOCS-SCZ is regulated by the jasmonic acid signalling pathway in the tea plant.

Tropospheric ozone (O3) pollution can affect plant nutritional quality and secondary metabolites by altering plant biochemistry and physiology, which may lead to unpredictable effects on crop quality and resistance to pests and diseases. Here, we investigated the effects of O3 (ambient air, Am; ambient air +80 ppb of O3, EO3) on the quality compounds and chemical defenses of a widely cultivated tea variety in China (Camellia sinensis cv. ‘Baiye 1 Hao’) using open-top chamber (OTC). We found that elevated O3 increased the ratio of total polyphenols to free amino acids while decreasing the value of the catechin quality index, indicating a reduction in leaf quality for green tea. Specifically, elevated O3 reduced concentrations of amino acids and caffeine but shows no impact on the concentrations of total polyphenols in tea leaves. Within individual catechins, elevated O3 increased the concentrations of ester catechins but not non-ester catechins, resulting in a slight increase in total catechins. Moreover, elevated O3 increased the emission of biogenic volatile organic compounds involved in plant defense against herbivores and parasites, including green leaf volatiles, aromatics, and terpenes. Additionally, concentrations of main chemical defenses, represented as condensed tannins and lignin, in tea leaves also increased in response to elevated O3. In conclusion, our results suggest that elevated ground-level O3 may reduce the quality of tea leaves but could potentially enhance the resistance of tea plants to biotic stresses.

Wintersweet (Chimonanthus praecox), a classical winter-blooming magnoliids species with great aesthetic value highlighting the on the brilliant yellow flower and pleasant floral aroma, possesses diverse varieties mainly classified in three groups (Concolor, Patens, and Rubrum group) with the most striking variation in the tepal color. The recent release of the genome in two wintersweet varieties belonging to Patens and Rubrum group successfully elucidated the molecular mechanisms underlying some ornamental traits highlighting the flowering time, floral scent and tepal color. In this study, the chromosome-scale genome of C. praecox var. concolor was assembled to further verify the placement of magnoliids and exploit the genetic variation underscoring the diverse flowering traits among different wintersweet groups. Phylogenomic analyses placed magnoliids including C. praecox var. concolor as the sister to eudicots. The C. praecox var. concolor’s genome experienced two rounds of whole genome duplication (WGD). Multiple gene families involved in floral development including ANR1 and PI genes were retained and expanded during these WGD events. The expansion and expression pattern of SVP and FUL genes suggest their pivotal role in unique character of winter-flowering. With the pleasant fragrance as the aesthetic characteristics, linalool and benzyl acetate were identified as the top abundant constituents in the floral scent of Concolor wintersweet. Integrating metabolic, genomic, and transcriptomic analyses with enzyme assays led to the identification of CpTPS3 (CpLIS) and CpBAHD2 (CpBEAT), which are responsible for the biosynthesis of linalool and benzyl acetate, respectively, in the Concolor variety. Taken together, the assembled genome of C. praecox var. concolor enables the comparative analysis of the genetic basis underlying both conserved and variable floral traits and provides a useful foundation for variety improvements by molecular breeding in wintersweet.

Flavanols are known as the most important antioxidants in tea (Camellia sinensis), but their contribution to the antioxidant capacity across tea cultivars has not been quantified. This study explored whether the variations of antioxidant capacity across tea cultivars could be linked to variations in main flavanol concentrations using 20 widely planted Chinese tea cultivars. The results showed that concentrations of flavanols, both monomeric (total catechins; 3.77%–8.85% d.w.) and polymeric forms (condensed tannins; 9.48%–17.67% d.w.), varied largely across tea cultivars. The contribution of total catechins to the antioxidant capacity in tea (R2 = 0.54–0.55) was greater than that of condensed tannins (R2 = 0.35–0.36) and total phenolic concentrations (R2 = 0.33–0.36). Individual catechin components collectively explained 54.94%–56.03% of the variations in antioxidant capacity across tea cultivars. Epigallocatechin gallate (EGCG) was the leading catechin component that determined the antioxidant capacity in tea (p < 0.001), accounting for up to 57% of the differences in catechin composition between tea cultivars with high and low antioxidant capacities. These results suggested that flavanols were stronger predictors of the antioxidant capacity across tea cultivars compared to total phenolic concentrations, providing guidance for breeding tea cultivars with strong antioxidant capacities.

Although the application of biochar increases crop productivity and soil fertility, the effects of different biochar application methods on tea plant growth and soil nutrient status remain unclear. In this study, we conducted a root chamber experiment to assess the effects of various biochar application methods on root growth in tea plants and edaphic factors. Four treatments were included: T1) No biochar, T2) Homogenized biochar, T3) Localized biochar, and T4) Strip biochar. Biochar application generally resulted in higher macronutrient contents in tea plants, with 114-371%, 196-1167%, and 327-960% increases in nitrogen (N), phosphorus (P), and potassium (K), respectively. Biochar application also increased the soil pH by 18.7-22.5% compared to the control. T2 treatment increased the leaf and root biomasses by 80.9% and 262.2%, respectively, compared to the control, and the total carbon and soil nutrient [i.e., total N, inorganic N, available P, and available K, calcium (Ca), and magnesium (Mg)] contents were significantly higher than in the control. Variation in root growth was signifi-cantly explained by soil properties, including microbial biomass carbon, Ca, Mg, and total N (P < 0.05), with contributions of 47.9%, 15.6%, 17.6%, and 4.7%, respectively. These results suggest that biochar application in acidic soils would increase tea plant productivity and soil nutrient contents. Overall, homogenized biochar application led to optimal tea plant growth and soil fertility.

Strip/needle green teas (SGT/NGT) processed using innovative technologies are in high demand; however, mechanisms behind their color and flavor have not been comprehensively studied. We aimed to reveal the dynamics of major pigmented components (carotenoids, lipids, flavonoids, and Maillard products) and their contributions to the flavor of green teas. The total content of flavonoids in SGT and NGT were 255 ± 4.51 and 201 ± 3.91 mg·g-1, respectively; these values are slightly lower than that in fresh leaves (FLs), resulting in a fresh and sweet aftertaste. In average, carotene content in SGT/NGT (24.8 μg·g-1) was higher than in FL (17.4 μg·g-1), whilst xanthophyll content (603 μg·g-1) decreased to one-half of that in FL (310 μg·g-1). Among the 218 primary metabolites, glutamine, glutamic acid, and arginine were found to accumulate and were dominate contributors for the umami and sweet taste. Notably, more than 96 volatiles were screened and revealed their correlations with carotenoids, lipids, and amino acids. Overall, the synergism between pigments and their non-enzymatic derivates' contribution to GT characterized flavor was illustrated.