Deciphering the underlying core microorganisms and the marker compounds of Liupao tea during the pile-fermentation process.

BACKGROUND: Pile fermentation is one of the key steps in developing the Liupao tea (LBT) quality and unique characteristics. The complex biochemical profile of LBT results from microorganisms present during the pile-fermentation process. However, the critical underlying microorganisms and the marker compounds still need to be determined. RESULTS: Staphylococcus, Brevibacterium, Kocuria, Aspergillus, and Blastobotrys were the common dominant microorganisms at the end of the pile fermentation of LBT. Staphylococcus, Aspergillus, Blastobotrys, and nine other genera carried by raw tea are the core microorganisms in the LBT during pile fermentation. A total of 29 critical compounds contributed to the metabolic changes caused by the processing of LBT. Of these, gallic acid, adenine, hypoxanthine, uridine, betaine, 3,4-dihydroxybenzaldehyde, and α-linolenic acid could be characterized as potential marker compounds. Correlation analysis showed that the core microorganisms, including Sphingomonas, Staphylococcus, Kocuria, Aureobasidium, Blastobotrys, Debaryomyce, and Trichomonascus, were closely related to major chemical components and differential compounds. Moreover, the mutually promoting Staphylococcus, Kocuria, Blastobotrys, and Trichomonascus were correlated with the enrichment of marker compounds. Integrated molecular networking and metabolic pathways revealed relevant compounds and enzymes that possibly affect the enrichment of marker compounds. CONCLUSION: This study analyzed the LBT fermentation samples by omics analysis to reveal the stable microbial community structure, critical microorganisms, and markers compounds affecting the quality of LBT, which contributes to a better understanding of pile fermentation of LBT and the fermentation theory of dark tea. © 2023 Society of Chemical Industry.

Development and evaluation of a qPCR detection method for citrinin in Liupao tea.

Penicillium is universal in dark tea, and Penicillium citrinum can produce a kidney toxin called citrinin (CIT). Determining CIT is difficult because of the complexity of the dark tea substrate and the diversity of CIT-producing fungi. Therefore, this study established a real-time PCR (qPCR) detection method for CIT-related synthetic genes (ctnD, orf1, ctnA, pksCT, orf5, orf7, and ctnG) in Liupao tea and determined the content of CIT in samples at different production stages and the toxin-producing abilities of fungi (Aspergillus oryzae, etc.) in Liupao tea. CIT was found in all samples during the pile-fermentation process of Liupao tea, and CIT was detected in two samples during the aging process. The established method demonstrated good sensitivity and specificity in detecting CIT-related synthetic genes. The reaction efficiency was within the preferred range of 100 ± 10%. CIT was not detected or was below the detection limit when the Ct value of one or more related synthetic genes was greater than 33.5. Therefore, the established qPCR method can effectively predict the production of CIT in Liupao tea, and it is applicable to the judgment of whether fungi produce CIT.

Regulation of fungal community and the quality formation and safety control of Pu-erh tea.

Pu-erh tea belongs to dark tea among six major teas in China. As an important kind of post-fermented tea with complex microbial composition, Pu-erh tea is highly praised by many consumers owing to its unique and rich flavor and taste. In recent years, Pu-erh tea has exhibited various physiological activities to prevent and treat metabolic diseases. This review focuses on the fungi in Pu-erh tea and introduces the sources, types, and functions of fungi in Pu-erh tea, as well as the influence on the quality of Pu-erh tea and potential safety risks. During the process of fermentation and aging of Pu-erh tea, fungi contribute to complex chemical changes in bioactive components of tea. Therefore, we examine the important role that fungi play in the quality formation of Pu-erh tea. The associations among the microbial composition, chemicals excreted, and potential food hazards are discussed during the pile-fermentation of Pu-erh tea. The quality of Pu-erh tea has exhibited profound changes during the process of pile-fermentation, including color, aroma, taste, and the bottom of the leaves, which are inseparable from the fungus in the pile-fermentation of Pu-erh tea. Specifically, the application prospects of various detection methods of mycotoxins in assessing the safety of Pu-erh tea are proposed. This review aims to fully understand the importance of fungi in the production of Pu-erh tea and further provides new insights into subtly regulating the piling process to improve the nutritional properties and guarantee the safety of Pu-erh tea.

Untargeted Metabolomics Combined with Bioassay Reveals the Change in Critical Bioactive Compounds during the Processing of Qingzhuan Tea.

Qingzhuan tea (QZT) is a typical Chinese dark tea that has a long-time manufacturing process. In the present study, liquid chromatography coupled with tandem mass spectrometry was used to study the chemical changes of tea samples during QZT processing. Untargeted metabolomics analysis revealed that the pile-fermentation and turnover (post-fermentation, FT) was the crucial stage in transforming the main compounds of QZT, whose contents of flavan-3-ols and flavonoids glycosides were decreased significantly. The bioactivities, including the antioxidant capacities and inhibitory effects on α-amylase and α-glucosidase, were also reduced after the FT process. It was suggested that although the QZT sensory properties improved following pile-fermentation and aging, the bioactivities remained restrained. Correlation analysis indicated that the main galloylated catechins and flavonoid glycosides were highly related to their antioxidant capacity and inhibitory effects on α-amylase and α-glucosidase.

Effects of Microbial Proteins on Qingzhuan Tea Sensory Quality during Pile Fermentation.

To determine the effects of microbial proteins on Qingzhuan tea sensory quality during tea pile fermentation, tea leaf metabolomic and microorganism proteomic analyses were performed. In total, 1835 differential metabolites and 443 differentially expressed proteins of the microorganisms were identified. Correlation analysis between metabolomics and proteomics data revealed that the levels of microbial proteins EG II and CBH I cellulase may play important roles in cell wall construction and permeability, which were crucial for the interaction between tea leaves and microorganisms. Microbial proteins heat shock proteins (HSP), alcohol dehydrogenase (ADH), aldehyde dehydrogenase (ALDH), and CuAO related to detoxification and stress responses showed a positive correlation with tea theanine, glutamine, γ-aminobutyric acid, glutamic acid, catechin, (-)-gallocatechin gallate, and (-)-catechin gallate, suggesting their effects on tea characteristic compound accumulation, thus affecting Qingzhuan tea sensory quality.

Degradation of Cell Wall Polysaccharides during Traditional and Tank Fermentation of Chinese Liupao Tea.

The increase of polysaccharides in the dark tea pile process is thought to be connected to the cell wall polysaccharides' breakdown. However, the relationship between tea polysaccharides (TPSs) and tea cell wall polysaccharides has not been further explored. In this study, the structural changes in the cell wall polysaccharides [e.g., cellulose, hemicellulose (HC), and pectin] in Liupao tea were characterized before and after traditional fermentation and tank fermentation. Additionally, the degradation mechanism of tea cell wall polysaccharides during fermentation was assessed. The results showed that cellulose crystallinity decreased by 11.9-49.6% after fermentation. The molar ratio of monosaccharides, such as arabinose, rhamnose, and glucose in HC, was significantly reduced, and the molecular weight decreased. The esterification degree and linearity of water-soluble pectin (WSP) were reduced. TPS content increases during pile fermentation, which may be due to HC degradation and the increase in WSP caused by cell wall structure damage. Microorganisms were shown to be closely associated with the degradation of cell wall polysaccharides during fermentation according to correlation analyses. Traditional fermentation had a greater effect on the cellulose structure, while tank fermentation had a more noticeable impact on HC and WSP.

Co-occurrence network and functional profiling of the bacterial community in the industrial pile fermentation of Qingzhuan tea: Understanding core functional bacteria.

The distinct quality of Qingzhuan tea is greatly influenced by the bacterial community but was poorly characterized. Therefore, this study investigated the Co-occurrence network and functional profiling of the bacterial community, with special attention paid to core functional bacteria in the industrial pile fermentation. Microbiomics analysis indicated that Klebsiella and Pantoea dominated raw tea leaves, and were rapidly replaced by Pseudomonas in pile fermentation, but substituted mainly by Burkholderia and Saccharopolyspora in final fermented tea. Bacterial taxa were grouped into 7 modules with the dominant in module I, III, and IV, which were involved in flavor formation and biocontrol production. Functional profiling revealed that "penicillin and cephalosporin biosynthesis" increased in pile fermentation. Twelve bacterial genera were identified as core functional bacteria, in which Klebsiella, Pantoea, and Pseudomonas also dominated the pile fermentation. This work would provide theoretical basis for its chemical biofortification and quality improvement by controlling bacterial communities.

Exploring core functional fungi driving the metabolic conversion in the industrial pile fermentation of Qingzhuan tea.

The distinct sensory quality of Qingzhuan tea is mainly formed in pile fermentation by a group of functional microorganisms but the core functional ones was poorly characterized. Therefore, this study investigated the dynamic changes in the fungal community and metabolic profile by integrating microbiomics and metabolomics, and explored the core functional fungi driving the metabolic conversion in the industrial pile fermentation of Qingzhuan tea. Indicated by microbiomics analysis, Aspergillus dominated the entire pile-fermentation process, while Thermoascus, Rasamsonia, and Cylindrium successively abounded in the different stages of the pile fermentation. A total of 50 differentially changed metabolites were identified, with the hydrolysis of galloyl/polymeric catechins, biosynthesis of theabrownins, oxidation of catechins, N-ethyl-2-pyrrolidinone substitution of catechins, and deglycosylation of flavonoid glucosides. Nine fungal genera were identified as core functional fungi, in which Aspergillus linked to the hydrolysis of polymeric catechins and insoluble polysaccharides as well as biosynthesis of theabrownins, while Thermoascus participated in the biosynthesis of theabrownins, deglycosylation of flavonoid glucosides, and N-ethyl-2-pyrrolidinone substitution of catechins. These findings would advance our understanding of the quality formation of Qingzhuan tea and provide a benchmark for precise inoculation for its quality improvement.

Insights into the airborne microorganisms in a Sichuan south-road dark tea pile fermentation plant during production.

Introduction: Sichuan south-road dark tea (SSDT) is generally produced through a series of processes, including fixing, rolling, pile fermentation, and drying, with microbial action during pile fermentation playing a crucial role in determining tea quality. The air within the SSDT pile fermentation plant (SSDTPP) is considered an important source of these microbes, but research in this area has been limited. Methods: In this study, air samples from SSDTPP were collected on the 1st (SSDT1), 12th (SSDT2), and 24th (SSDT3) days of pile fermentation and comprehensively analyzed by high-throughput sequencing. Results and discussion: The results revealed the presence of 2 and 24 phyla, 9 and 49 classes, 18 and 88 orders, 28 and 153 families, 38 and 253 genera, and 47 and 90 species of fungi and bacteria, respectively, across all samples. SSDT1 and SSDT2 individually had the highest fungal and bacterial diversity, while Aspergillus was the dominant genus throughout the pile fermentation with an abundance of 34.6%, 91.17%, and 67.86% in SSDT1, SSDT2, and SSDT3, respectively. Microbial populations in SSDT1 were predominantly involved in xenobiotic biodegradation and metabolism, amino acid metabolism, the biosynthesis of other secondary metabolites, etc. However, SSDT2 exhibited a higher prevalence of human disease-related functions. SSDT3 primarily focused on the metabolism of other amino acids and carbohydrate metabolism. Additionally, 104 genera and 22 species coexisted in both SSDTPP air and piled SSDT, suggesting that frequent microbial exchange may occur between them. These findings pave the way for microbial traceability during SSDT production and provide a foundation for further functional microbial research.

A preliminary metagenomic study of puer tea during pile fermentation.

BACKGROUND: Up to now, there has been no report on the taxonomic and functional analysis of the microbial community in fermenting puer tea by pyrosequencing. In this study, metagenomic pyrosequencing was first used in fermenting puer tea to delineate a relatively comprehensive overview of the microbial taxonomy while also preliminarily characterising the functional ontologies of microbial genes present in puer tea pile fermentation. RESULTS: A total of 251 738 pyrosequencing reads (9197 contigs and 145 402 singletons) were generated by pyrosequencing. Taxonomic analysis revealed three dominant bacterial phyla, Actinobacteria (30.08%), Proteobacteria (24.47%) and Firmicutes (20.23%), and one dominant eukaryotic phylum, Ascomycota (15.21%) [corrected]. A total of 58 664 hits were categorised into 28 functional subsystems based on the SEED database. Moreover, two categories, 'metabolism of terpenoids and polyketides' and 'biosynthesis of other secondary metabolites', were selectively analysed and 69 enzyme genes were presented in 16 pathways. CONCLUSION: The dominant microbes of puer tea fermentation were bacteria in the present study, and yeasts rather than moulds accounted for the overwhelming majority of Eukaryota. The analysis of functional genes and metabolic pathways will be helpful for further study of the mechanism of puer tea fermentation at molecular level.

Effects of pile-fermentation on the aroma quality of dark tea from a single large-leaf tea variety by GC × GC-QTOFMS and electronic nose.

Aroma is one of the significant quality factors of dark tea (DT). However, for a single large-leaf tea variety, there are few studies analyzing the effect of pile-fermentation on the aroma quality of DT. The GC × GC-QTOFMS, electronic nose (E-nose) and GC-olfactometry (GC-O) techniques were employed to analysis the difference of tea products before and after pile-fermentation. A total of 149 volatile metabolites (VMs) were identified, with 92 VMs exhibiting differential characteristics. Among these, 31 VMs with OAV > 1.0 were found to be correlated with E-nose results (|r| > 0.8). Additionally, GC-O analysis validated seven major differential metabolites. Notably, naphthalene, 2-methylnaphthalene, and dibenzofuran were found to enhance the woody aroma, while (Z)-4-heptenal, 2-nonenal and 1-hexanol were associated with an increase in mushroom, fatty and sweet odors, respectively. Moreover, 1-octen-3-ol was linked to reducing pungent fishy smell. These findings could provide a certain theoretical basis for understanding the influence of pile-fermentation on the aroma quality of dark tea.

Changes in sensory characteristics, chemical composition and microbial succession during fermentation of ancient plants Pu-erh tea.

"Ancient tea plants" are defined as tea trees > 100 years old, or with a trunk diameter > 25 cm; their leaves are manufactured to high - quality, valuable ancient plants pu-erh tea (APPT). In this study, a fermentation of APPT were developed, and outstanding sweetness of APPT infusion was observed. During fermentation, the content of soluble sugars, theabrownins (p < 0.05), as well as 41 metabolites were increased [Variable importance in projection (VIP) > 1.0; p < 0.05 and Fold-change (FC) FC > 2]; While relative levels of 72 metabolites were decreased (VIP > 1.0, p < 0.05 and FC < 0.5. Staphylococcus, Achromobacter, Sphingomonas, Thermomyces, Rasamsonia, Blastobotrys, Aspergillus and Cladosporium were identified as dominant genera, and their relative levels were correlated with contents of characteristic components (p < 0.05). Together, changes in sensory characteristics, chemical composition and microbial succession during APPT fermentation were investigated, and advanced the formation mechanism of its unique quality.

Microbial Diversity and Characteristic Quality Formation of Qingzhuan Tea as Revealed by Metagenomic and Metabolomic Analysis during Pile Fermentation.

In order to analyze the changes in the microbial community structure during the pile fermentation of Qingzhuan tea and their correlation with the formation of quality compounds in Qingzhuan tea, this study carried out metagenomic and metabolomic analyses of tea samples during the fermentation process of Qingzhuan tea. The changes in the expression and abundance of microorganisms during the pile fermentation were investigated through metagenomic assays. During the processing of Qingzhuan tea, there is a transition from a bacterial dominated ecosystem to an ecosystem enriched with fungi. The correlation analyses of metagenomics and metabolomics showed that amino acids and polyphenol metabolites with relatively simple structures exhibited a significant negative correlation with target microorganisms, while the structurally complicated B-ring dihydroxy puerin, B-ring trihydroxy galloyl puerlin, and other compounds showed a significant positive correlation with target microorganisms. Aspergillus niger, Aspergillus glaucus, Penicillium in the Aspergillaceae family, and Talaromyces and Rasamsonia emersonii in Trichocomaceae were the key microorganisms involved in the formation of the characteristic qualities of Qingzhuan tea.

Effects of pile fermentation on the physicochemical, functional, and biological properties of tea polysaccharides.

This study investigated the influence of pile fermentation on the physicochemical, functional, and biological properties of tea polysaccharides (TPS). Results indicated that the extraction yield, uronic acid content, and polyphenol content of TPS greatly increased from 1.8, 13.1 and 6.3 % to 4.1, 27.9, and 7.8 %, respectively, but the molecular weight markedly decreased from 153.7 to 76.0 kDa after pile fermentation. Additionally, the interfacial, emulsion formation, and emulsion stabilization properties of TPS were significantly improved after pile fermentation. For instance, 1.0 wt% TPS isolated from dark tea (D-TPS) can fabricate 8.0 wt% MCT oil-in-water nanoemulsion (d32 ≈ 159 nm) with potent storage stability. Moreover, the antioxidant and α-glucosidase inhibitory activities of D-TPS was higher than that of TPS isolated from sun-dried raw tea (R-TPS). Overall, this study indicated that pile fermentation markedly affected the physicochemical and structural characteristics of TPS, thereby improving their functional and biological properties.

Dynamic Changes in the Microbial Community and Metabolite Profile during the Pile Fermentation Process of Fuzhuan Brick Tea.

The pile fermentation process of Fuzhuan brick tea is unique in that it involves preheating without the use of starter cultures. The detailed metabolite changes and their drivers during this procedure are not known. Characterizing these unknown changes that occur in the metabolites and microbes during pile fermentation of Fuzhuan brick tea is important for industrial modernization of this traditional fermented food. Using microbial DNA amplicon sequencing, mass spectrometry-based untargeted metabolomics, and feature-based molecular networking, we herein reveal that significant changes in the microbial community occur before changes in the metabolite profile. These changes were characterized by a decrease in Klebsiella and Aspergillus, alongside an increase in Bacillus and Eurotium. The decrease in lysophosphatidylcholines, unsaturated fatty acids, and some astringent flavan-3-ols and bitter amino acids, as well as the increase in some less astringent flavan-3-ols and sweet or umami amino acids, contributed importantly to the overall changes observed in the metabolite profile. The majority of these changes was caused by bacterial metabolism and the corresponding heat generated by it.

Influence of thermophilic microorganism on non-volatile metabolites during high-temperature pile-fermentation of Chinese dark tea based on metabolomic analysis.

Pile-fermentation is a critical procedure for producing Chinese dark tea, during which thermophilic microorganisms would play an irreplaceable role. However, there have been little researches on the influences of thermophilic microorganism pile-fermentation (TMPF) in high-temperature of Chinese dark tea. Thus, we conducted high-performance liquid chromatography and nontargeted metabolomic to analyze the non-volatile metabolites of TMPF. Our results discovered that the amounts of ( -)-epigallocatechin gallate, ( -)-epigallocatechin, ( -)-epicatechin gallate, and ( -)-epicatechin were decreased significantly (p < 0.05) after TMPF. By using nontargeted metabolomic analysis, a total of 1733 ion features were detected. KEGG pathway enrichment analysis showed that TMPF had a significant impact on caffeine metabolism. Also, theophylline, 3-methylxanthine, and 1,3,7-trimethyluric acid were increased significantly after TMPF, which suggested that demethylation and oxidation reaction might be the main pathways of caffeine metabolism. This study provides a better understanding of the mechanism of TMPF during high-temperature for Chinese dark tea and lays a foundation for further research. Supplementary Information: The online version contains supplementary material available at 10.1007/s10068-022-01098-9.

Microbial Community Analysis in Sichuan South-road Dark Tea Piled Center at Pile-Fermentation Metaphase and Insight Into Organoleptic Quality Development Mediated by Aspergillus niger M10.

Microbes are critical in the Sichuan South-road Dark Tea (SSDT) organoleptic quality development during pile-fermentation. Piled tea center at fermenting metaphase is crucial for the conversion of its quality components. In this study, we investigated the microbial community of piled SSDT center below the stacked tea surface of 15 cm (SSDTB), 50 cm (SSDTX), and 85 cm (SSDTH) on the second turning time of pile-fermentation, respectively. Results showed that SSDTH and SSDTB had a higher similarity in the microbial community. Pantoea (36.8%), Klebsiella (67.7%), and Aspergillus (35.3%) were the most abundant in SSDTH, SSDTB, and SSDTX, respectively. We found 895 species were common among all samples, but 86, 293, and 36 species were unique to SSDTB, SSDTX, and SSDTH, respectively. Aspergillus niger showed high co-occurrence and was positively correlated with numerous microbes in SSDT samples, and Aspergillus niger M10 isolated from SSDTX was excellent at enhancing soluble sugar (SS), amino acids (AAs), theaflavin (TF), and thearubigins (TR) contents, while decreasing catechin (Cat), tea polyphenols (TPs)/AA, Caf/SS, Cat/SS, TPs/SS, and (TPs + Caf)/SS levels in AM10 post-fermentation, as compared with the control. Moreover, it also produced a noticeable difference in the CIELab parameters in dried, liquor, and infused tea colors between AM10 and control during fermentation. When it was further inoculated on differential mediums, we detected glycoside hydrolases, namely, ß-glucosidase, mannosidase, pectinase, cellulase, amylase, and α-galactosidase being secreted by Aspergillus niger M10. Taken together, SSDXT presented a more unique microbial community. Aspergillus niger M10 probably improved the sweet and mellow taste, and the yellow brightness and red color of SSDT during fermentation. It also provided new insights into the microbial profile and organoleptic quality development mechanism of SSDT during pile-fermentation.

Dynamic Analysis of Physicochemical Properties and Polysaccharide Composition during the Pile-Fermentation of Post-Fermented Tea.

Ultra-high performance liquid chromatography-quadrupole-time of flight tandem mass spectrometry (UHPLC-Q-TOF-MS/MS) was used to study the diversity of tea polysaccharides and the dynamic changes in the physicochemical indexes of tea samples. FT-IR spectra and the free radical scavenging ability of tea polysaccharides, during pile-fermentation of post-fermented tea, were analyzed. The results showed that 23 saccharide co mponents in tea polysaccharides were identified: these belonged to 11 monosaccharides, 5 oligosaccharides, and 6 derivatives of monosaccharides and oligosaccharides. The abundance of oligosaccharides decreased gradually, while monosaccharides, and derivatives of monosaccharides and oligosaccharides increased gradually with the development of pile-fermentation. According to the differences in polysaccharide composition and their abundance, the tea polysaccharide samples extracted from different pile-fermentation stages could be clearly classed into three groups, W-0, W-1~W-4 and W-5~C-1. The pile-fermentation process affected the yield, the content of each component, FT-IR spectra, and the DPPH free radical scavenging ability of tea polysaccharides. Correlation analysis showed that microorganisms were directly related to the changes in composition and the abundance of polysaccharides extracted from different pile-fermentation stages. The study will further help to reveal the function of tea polysaccharides and promote their practical application as a functional food.

Microbial Succession and Interactions During the Manufacture of Fu Brick Tea.

Fu Brick tea is a very popular post-fermented tea that is known for its "golden flower fungus," Aspergillus cristatus, which becomes the dominant microbe during the maturation process. This study used both culture-dependent methods and high-throughput sequencing to track microbial succession and interactions during the development of the golden flower fungus, a crucial component of the manufacturing process of Fu Brick tea. Among the bacterial communities, Klebsiella and Lactobacillus were consistently cultured from both fresh tea leaves and in post-fermentation Fu Brick tea. Methylobacterium, Pelomonas, and Sphingomonas were dominant genera in fresh tea leaves but declined once fermentation started, while Bacillus, Kluyvera, and Paenibacillus became dominant after piling fermentation. The abundance of A. cristatus increased during the manufacturing process, accounting for over 98% of all fungi present after the golden flower bloom in the Fu Brick tea product. Despite their consistent presence during culture work, network analysis showed Lactobacillus and Klebsiella to be negatively correlated with A. cristatus. Bacillus spp., as expected from culture work, positively correlated with the presence of golden flower fungus. This study provides complete insights about the succession of microbial communities and highlights the importance of co-occurrence microbes with A. cristatus during the manufacturing process of Fu Brick tea.

Dynamic changes in the aroma profile of Qingzhuan tea during its manufacture.

Changes in key odorants and aroma profiles of Qingzhuan tea (QZT) during its manufacture were determined using headspace solid-phase microextraction gas chromatography-mass spectrometry/olfactometry. An aroma profile was constructed to illustrate sensory changes during manufacture. The characteristic aroma of QZT was aged fragrance, which was mostly developed during pile fermentation and was enhanced during the aging and drying stages. Using volatile compounds found in the raw materials, sun-dried green tea and QZT finished product were compared by orthogonal partial least square-discriminant analysis. Among 108 detected volatiles, 19 were significantly upregulated and 15 were downregulated. (E)-ß-Ionone, (E,Z)-2,6-nonadienal, 1-octen-3-one, (E,E)-2,4-heptadienal, (E,E)-2,4-nonadienal, safranal, (E)-2-nonenal, α-ionone, and 1,2,3-trimethoxybenzene were found to be significant contributors to the aged QZT fragrance, reflecting their high odor-activity values and aroma intensities. Finally, the metabolic transformation of key aroma-active compounds was systematically analyzed. This study provided a theoretical basis for improving the processing and quality of QZT.