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Article

Viroid GYSVd1 Exhibited Different Regulations on the Qualities of Berries and Wines from 6 Grape Varieties

by
Menghuan Wu
1,2,
Shuo Liu
1,
Ping Wang
3,
Zhaotan Li
1,
Junbo Zhang
3,
Yejuan Du
1,* and
Shuhua Zhu
2,*
1
Key Laboratory of Oasis Agricultural Pest Management and Plant Protection Resources Utilization, College of Agriculture, Shihezi University, Shihezi 832003, China
2
College of Chemistry and Material Science, Shandong Agricultural Univerisity, Taian 271018, China
3
College of Food Science and Technology, Shihezi University, Shihezi 832000, China
*
Authors to whom correspondence should be addressed.
Horticulturae 2025, 11(4), 345; https://doi.org/10.3390/horticulturae11040345
Submission received: 22 January 2025 / Revised: 21 March 2025 / Accepted: 21 March 2025 / Published: 22 March 2025
(This article belongs to the Section Viticulture)
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Abstract

:
Grapes are one of the widely cultivated fruits, with high nutritional value and economic value. The widespread occurrence of grape virus diseases has seriously affected the development of the grape industry. The grapevine varieties “Merlot”, “Cabernet Sauvignon”, “Syrah”, “Chardonnay”, “Welsch Riesling ”, and “Riesling Weiss” were used as materials for screening grapevines carrying the viroid GYSVd1 by RT-PCR. Principal component analysis (PCA) was employed to systematically assess the physicochemical indexes of both grape berries and wine in order to determine the extent of the influence of GYSVd1 on the quality of grape berries and wine. The results demonstrated that GYSVd1 infection significantly compromised both berries and wine quality across the tested cultivars, albeit with distinct varietal susceptibility patterns. Regarding berries’ quality, the negative impact of GYSVd1 followed this decreasing order: Merlot > Chardonnay > Welsch Riesling > Syrah > Riesling Weiss > Cabernet Sauvignon. Similarly, for wine quality, the negative impact exhibited the following gradient: Welsch Riesling > Riesling Weiss > Chardonnay > Cabernet Sauvignon > Syrah > Merlot. There were significant differences in the amount of sugar, acid and phenolic substances between GYSVd1-infected and -uninfected grapevines, but no significant differences in berry weight, berry shape index, and alcohol content. GYSVd1 affected the quality of berries and wine mainly by regulating the contents of sugar, acid, and phenolic substances.

1. Introduction

As one of the most widely cultivated fruit crops, grape (Vitis vinifera L.) holds substantial economic importance globally, driven by its distinctive sensory characteristics, including its flavor and esthetic qualities [1]. Grape berries are rich in minerals, vitamins, phenolic substance, amino acids, and other nutrients, with unique flavors, and are widely loved by consumers [2]. Xinjiang is one of the main areas in which wine and grapes are produced. There, the grape industry has developed rapidly, the cultivation area has been expanding, and propagation via cuttings is widespread, leading to the long-term accumulation of viruses and the widespread occurrence of grape virus diseases [3]. There are 102 grapevine virus species worldwide [4], and 22 have been reported in China [5]. Virus diseases are widely distributed and affect grape yield and quality [6]. Grapevine leafroll-associated virus 3 (GLRaV-3) affects fruit ripening, sugar content, and phenolic substance content [7,8]. Grapevine red blotch virus (GRBV) reduces berries’ weight and phenolic content, delays ripening, and affects the sensory quality of the wine [9,10]. Grapevine fanleaf virus (GFLV) affects the soluble solids (TSSs) of the berries, as well as the berries’ color, firmness, etc. [11]. Previous studies have mainly focused on the effects of viruses on grape berries and wine quality, while the effects of viroids on grape quality are still unclear.
Viroids are a class of single-stranded, covalently closed circular RNA molecules that do not encode any proteins but are capable of self-replication within host plants [12,13]. These pathogens are characterized by their long incubation periods and significant harm, primarily spreading through agricultural practices such as pruning and grafting [14]. The transmission routes of grapevine yellow speckle viroid 1 (GYSVd1) include seed transmission and mechanical transmission [15]. If pruning tools or farming equipment are not thoroughly disinfected, healthy grapevines can become infected. Once a grapevine is infected with a viroid, it will carry the pathogen for its entire lifespan. Grapevine yellow speckle viroid 1 (GYSVd1) belongs to the potato spindle tuber viroidae family (Posplviroidae) and the rust-fruit virus genus (Apscaviroid) [15]. Previous studies showed that in the process of infecting grapes, viroids often existed in the form of a complex infection with other viruses. There is potential for synergistic or antagonistic interactions between viruses and viroids, which can have variable effects on grape quality. Furthermore, the specific impacts of GYSVd1 infection on grape berry and wine quality have not yet been reported.
The quality of grapes directly affects the economic benefits and determines consumers’ choice tendencies [16]. The primary quality attributes of grapes encompass the fruit shape index, flavor profile, aroma, sugar–acid balance, anthocyanin content, tannin levels, among others [17,18,19]. Each quality index is independent of the others but has a particular relationship. The type and amount of sugar and acid are the key factors affecting the quality of berries and wine. Sugars improve the color, flavor, and taste of wine, but they also increase the smoothness of the taste and the fullness of the wine body and are also the substrates of alcoholic fermentation [20]. Grapevine roditis leaf discoloration-associated virus (GRLDaV) reduced the sugar content of grapes, reducing the quality of the berries and the wine [21]. Acid substances increase the taste and flavor of wine and effectively inhibit the growth of microorganisms in wine [22]. GFLV decreased the pH of grape berries, possibly by increasing their malic acid or tartaric acid content [11]. Phenolic substances are a class of compounds with complex structures and wide varieties, divided into flavonoids and non-flavonoids, including common anthocyanin, tannin, flavonol, hydroxybenzoic and hydroxycinnamic acids [23]. Phenolic substances have antioxidant, antithrombotic, antibacterial, antiultraviolet, and other functions [24]. As one of the viroids susceptible to grapevines, hop stunt viroid (HSVd) reduced the content of hop phenols, especially flavan-3 alcohols, but the effect on the phenolic substances in wine grapes has not been reported [25].
There are few studies on the effects of grape viroids on berries and wine quality. This study compared the effects of viroid GYSVd1 on the berries and wine quality of Merlot, Cabernet Sauvignon, Syrah, Chardonnay, Welsch Riesling, and Riesling Weiss to explore the possible usage to overcome the impact of viroids on grape berries and wine.

2. Materials and Methods

2.1. Plant Materials

In the field, 50 grapevines (selected with a five-point sampling method) from each variety were subjected to virus detection analysis. Systematic detection was performed for 12 viruses and 7 viroids, which were common in Xinjiang region. The corresponding primer sequences used in this detection are provided in detail in the Supplemental Files (Table S2).
Merlot (Me), Cabernet Sauvignon (CS), Syrah (Sy), Chardonnay (Ch), Welsch Riesling (WR), and Riesling Weiss (RW) grapevines infected with (5 grapevines) or without viroid GYSVd1 (5 grapevines) were in the grapery of Zhangyu Winery, Shihezi, China. The growing environment, climate conditions, and cultivation measures of the 6 grapevine varieties were the same, and the trees were 10 years old. After maturing, berries (40 kg) with GYSVd1 (GYSVd1-Me, GYSVd1-CS, GYSVd1-Sy, GYSVd1-Ch, GYSVd1-WR, GYSVd1-RW) and without GYSVd1 (UF-Me, UF-CS, UF-Sy, UF-Ch, UF-WR, UF-RW, UF: free from virus and viroid infections) were collected, respectively, from different orientations (top, middle, and bottom) of the 6 varieties of grapevines, and 30 kg of berries were used for physicochemical index detection, while another 10 kg were used for winemaking. The wine was made as described by Zhang et al. [26]; the details of the winemaking procedures can be found in the Supplemental Files.

2.2. Viroid GYSVd1 Identification

The infection status of the grapevines in the field was ascertained by testing for GYSVd1 using the RT-PCR technique. The RNA from leaves and berries were extracted with a kit from Aidlab Biotechnologies Co., Ltd, Suzhou, China, which was stored in a −80 °C refrigerator after extraction. The leaf total RNA was used as the template, and the EasyScript First-Strand cDNA Synthesis SuperMix kit (Beijing TransGen Biotech Co., Ltd, Beijing, China) was used to add reaction components to efficiently synthesize the first-strand cDNA with a reverse transcription system of 20 µL (Table S1). GYSVd1 viroid accumulation in berries was quantified with the q-PCR primers of the target gene and the internal reference gene EF1α in Table S3. All q-PCR reactions were set up with 5 biological replicates, and the relative accumulation was calculated by the 2−ΔΔCT method.

2.3. Determination of General Physical and Chemical Indexes

Berry weight: the grape berries (30 berries per grapevine) were randomly selected as samples and weighed on an electronic balance (TD2002A, TD-A, Tianjin, China).
Berry shape index: the grape berries (30 berries per grapevine) were randomly selected, and their longitudinal and transverse diameters were measured with digital vernier calipers (605B-01, HMCT, Ningbo, China).
The berry shape index was the ratio of longitudinal diameter to transverse diameter.
Wine alcoholic strength: the alcoholic strength was achieved by measuring the distillate’s density through a densimeter (DMA-5000, Anton-Paar, Graz, Austria).
Berry soluble solid (TSS): the grape berries (30 berries per grapevine) were randomly selected, ground, and filtered to produce the juice. To measure TSS, the grape juice (1.0 mL) was dripped into a digital refractometer (PAL-1, Atago, Tokyo, Japan).
Berry and wine pH: the grape berries were randomly selected, ground, and filtered to produce the juice. The grape juice or wine (20 mL) was measured with a calibrated pH meter (S400-K, Mettler Toledo, Shanghai, China).

2.4. Determination of Total Phenolic Compounds

The content of total phenolic compounds was determined by Folin’s phenol method, referring to the method of Dravie et al. [27]. The berries (1.0 g) were homogenized with 1.5 mL ethanol of 60% (v/v) and oscillated extraction at 60 °C for 2 h. The homogenate was centrifuged for 10 min (25 °C, 12,000 rpm), and then the supernatant was collected. The wine samples can be directly used for the detection of indicators. The distilled water (5.0 mL) and Folinshoka reagent (200 μL) (F9252, Merck, Darmstadt, Germany) were added to the supernatant or wine sample (100 μL), which was shaken well. After standing for 2 min, 10% (v/v) NaCO3 solution (2.0 mL) was added, mixed, and left in darkness for 1 h. The absorbance was measured at 765 nm. The results were quantified and expressed as gallic acid equivalents (GAEs), with concentrations reported in mg g−1 for berry samples and mg mL−1 for wine samples.

2.5. Determination of Anthocyanin

The anthocyanin content was quantified using pH differential spectrophotometry at wavelengths of 510 nm and 700 nm, following the methodology described by Shi et al. [28]. The berries powder (1.0 g) was extracted with 10 mL 0.2% (v/v) HCl, and ultrasonic extraction was performed for 30 min (40 kHz, 150 W). The supernatant or wine sample (500 μL) was added with potassium chloride buffer (pH 1.0) and sodium acetate buffer (pH 4.5), and the absorbance was measured at 510 nm and 700 nm, respectively. The anthocyanin concentration was calculated as follows:
C = ([(A510 − A700) pH1.0 − (A510 − A700) pH4.5] × MW × L) ÷ ε MW and ε are the molecular weight (449 gmol−1) and molar absorptivity (29,600 Lmol−1 cm−1) of centaurin-3-glucoside, respectively, and L is the dilution ratio. The results were quantified and expressed as cyanidin-3-glucoside equivalents (C3GEs), with concentrations reported in mg g−1 for berry samples and mg mL−1 for wine samples.

2.6. Determination of Tannin

The tannin content was determined by the Folin–Dennis method. The extraction method of tannin was the same as that of total phenol. The supernatant or wine sample (100 μL) was mixed distilled water (7.5 mL), Folin–Dennis reagent (0.5 mL) (47742-F, Merck, Germany), and 20% (v/v) NaCO3 solution (1.0 mL). The absorbance was measured at 760 nm after a 30 min light-shielding reaction [29]. The results were quantified and expressed as catechin equivalents (CEs), with concentrations reported in mg g−1 for berry samples and mg mL−1 for wine samples.

2.7. Determination of Soluble Sugar

The soluble sugar content was measured as described by Xiao et al. [30]. The berries (1.0 g) were ground with distilled water, bathed in water at 80 °C for 30 min, cooled to room temperature, and centrifuged for 10 min (4 °C, 12,000 rpm) to collect the supernatant. The supernatant or wine sample (1.0 mL) was taken in a 15 mL test tube, and DNS (2.0 mL) was added and boiled for 5 min. The absorbance was measured at 540 nm.

2.8. Data Analysis

There were 5 analytical replicates per treatment. The physicochemical indexes of berries and wine were analyzed using principal component analysis (PCA). Data were analyzed using a two-sample/group t-test. Different letters indicated significant differences when the p-value was < 0.05.

3. Results

3.1. Detection of GYSVd1 of Six Grape Varieties

There were 300 grapevines of six wine grape varieties tested for GYSVd1. The results showed that the viroid GYSVd1 could infect all six grape varieties, and the infection rate was high: 12.00~40.00%. These results indicated that GYSVd1 had strong infection ability and a wide infection range in the field (Figure 1).

3.2. Viroid Accumulation of GYSVd1 in 6 Grape Varieties

In Merlot and Riesling Weiss, the accumulation of GYSVd1 was significantly higher in infected grapevines compared to uninfected grapevines (p < 0.05), with levels reaching 35.30 and 5.55 times higher, respectively (Figure 2A,F). No significant difference was observed in the accumulation of GYSVd1 in Cabernet Sauvignon, Syrah, Chardonnay, and Welsch Riesling compared to grapevines without GYSVd1 (Figure 2B–E).

3.3. GYSVd1 Modulated the Changes in General Physicochemical Indexes of Grape Berries and Wine

For Merlot and Welsch Riesling berries, the pH of berries carrying GYSVd1 was significantly lower than that of the berries without GYSVd1 (p < 0.05), by 93.83% and 98.85%, respectively. There were no significant differences in the pH of the berries of the other four varieties with infected berries compared with the berries without GYSVd1. In Merlot, Cabernet Sauvignon, and Welsch Riesling, the TSSs of berries carrying GYSVd1 decreased to only 95.33%, 88.99%, and 80.31% of that of the berries without GYSVd1, respectively. In Syrah and Chardonnay, the TSSs of berries carrying GYSVd1 were significantly higher than those of the berries without GYSVd1 (p < 0.05), by 1.07 and 1.12 times. However, no significant differences were observed in berry weight and berry shape index between infected and uninfected berries across the six grape varieties examined (Table 1).
For wine made from Merlot, Cabernet Sauvignon, and Syrah, the pH of wine made from berries infected with GYSVd1 was significantly higher than that made from the berries without GYSVd1 (p < 0.05), by 1.04, 1.02, and 1.02 times, respectively. In terms of alcoholic strength, there was no significant difference between infected and uninfected berries in the six grape varieties (Table 1).

3.4. GYSVd1 Modulated the Total Phenolic Compounds Content of Grape Berries and Wine

For Chardonnay, Welsch Riesling, and Riesling Weiss berries, the total phenolic compounds content of berries carrying GYSVd1 was significantly higher than that of the berries without GYSVd1 (p < 0.05), by 1.10, 1.18, and 1.19 times, respectively (Figure 3D–F). However, the total phenolic compounds content of berries carrying GYSVd1 in Merlot was significantly lower than that of the berries without GYSVd1 (p < 0.05), which was 83.06% (Figure 3A). In Cabernet Sauvignon and Syrah, there were no significant differences in total phenolic compounds content between the infected berries and the uninfected berries (Figure 3B,C). For wine made from Merlot and Cabernet Sauvignon berries, the total phenolic compounds content of wines made from berries infected with GYSVd1 was significantly lower than that of the wines made from berries without GYSVd1 (p < 0.05), by 84.06% and 87.45%, respectively (Figure 3G,H). In Syrah and Welsch Riesling, the total phenolic compounds content of wines made from berries infected with GYSVd1 was significantly higher than that of wines made from berries without GYSVd1 (p < 0.05), by 1.06 and 1.11 times, respectively (Figure 3I,K). In Chardonnay and Riesling Weiss, no significant differences existed in the total phenolic compounds content of the wines made from berries infected with GYSVd1 compared with the wines made from the berries without GYSVd1 (Figure 3J,L).

3.5. GYSVd1-Modulated Anthocyanin Content of Grape Berries and Wine

For berries of Cabernet Sauvignon, the anthocyanin content of berries carrying GYSVd1 was 1.48 times higher than that of the berries without GYSVd1 (p < 0.05) (Figure 4B). However, in Merlot and Syrah, the anthocyanin content of berries carrying GYSVd1 had no significant effect compared with the berries without GYSVd1 (Figure 4A,C). For wine made from Merlot berries, the anthocyanin content of wine made from berries infected with GYSVd1 was 1.64 times higher than that of the wine made from berries without GYSVd1 (p < 0.05) (Figure 4D). However, in Cabernet Sauvignon, the anthocyanin content of the wine made from berries infected with GYSVd1 was significantly (51.69%) lower than that of the wine made from the berries without GYSVd1 (p < 0.05) (Figure 4E).

3.6. GYSVd1-Modulated Tannin Content of Grape Berries and Wine

For Syrah, Chardonnay, and Riesling Weiss berries, the tannin content of berries carrying GYSVd1 was significantly higher than that of the berries without GYSVd1 (p < 0.05), by 1.08, 1.11, and 1.28 times, respectively (Figure 5C,D,F). In Merlot, Cabernet Sauvignon, and Welsch Riesling, the tannin content of berries carrying GYSVd1 was not significantly different from that of the berries without GYSVd1 (Figure 5A,B,E). For wine made from Merlot and Riesling Weiss, the tannin content of wines made from berries infected with GYSVd1 significantly decreased to only 90.27% and 90.49% that of the wines made from berries without GYSVd1 (p < 0.05), respectively (Figure 5G,L). In Cabernet Sauvignon, Syrah, Chardonnay, and Welsch Riesling, there were no significant differences in tannin contents between the wines produced by berries with GYSVd1 and those produced by the berries without GYSVd1 (Figure 5H,I,K).

3.7. GYSVd1-Modulated Soluble Sugar Content of Grape Berries and Wine

For Chardonnay and Welsch Riesling berries, the soluble sugar content of berries carrying GYSVd1 was significantly lower than that of the berries without GYSVd1 (p < 0.05), which contained 74.18% and 78.95%, respectively (Figure 6D,E). However, in Merlot, Cabernet Sauvignon, Syrah, and Riesling Weiss, the soluble sugar content of berries carrying GYSVd1 had no significant effect on that of the berries without GYSVd1 (Figure 6A–C,F). For wine made from Welsch Riesling and Riesling Weiss berries, the soluble sugar content of wines made from berries infested with GYSVd1 was significantly lower than that of wines made from berries without GYSVd1 (p < 0.05), for which the soluble sugar content was 84.55% and 84.96%, respectively (Figure 6K,L). However, in Merlot, Cabernet Sauvignon, Syrah, and Chardonnay, there were no significant differences in the amount of soluble sugar in the wines made from the berries with GYSVd1 compared with the wines made from the berries without GYSVd1 (Figure 6G–J).

3.8. Principal Component Analysis (PCA)

The total phenols were significantly positively correlated with anthocyanin, tannin, and fruit shape index, and berry pH was significantly negatively correlated with berry weight (Figure 7A). The wine TSS was significantly positively correlated with alcohol content, anthocyanin, and soluble sugar, and soluble sugar was also significantly positively correlated with total phenol, tannin, and anthocyanin (Figure 7D). According to the scree plots, two main components were extracted from berries and wine, respectively. The cumulative contribution rate of the two principal components to the comprehensive quality reached more than 80%, which proved that the two principal components could represent other quality indicators for analysis and evaluation (Figure 7B,E). The comprehensive score was calculated using the variance contribution rate corresponding to two principal components as the weight. As illustrated in the score and loading plots, GYSVd1 infection differentially affected grape berries and wine quality across various cultivars. The extent of berry quality reduction, quantified as the differential between GYSVd1-infected and uninfected berries’ scores, exhibited the following decreasing order: Merlot > Chardonnay > Welsch Riesling > Syrah > Riesling Weiss > Cabernet Sauvignon. Correspondingly, the wine quality decline, measured by the score difference between wines produced from infected versus healthy berries, exhibited the following decreasing order: Welsch Riesling > Riesling Weiss > Chardonnay > Cabernet Sauvignon > Syrah > Merlot (Figure 7C,F).

4. Discussion

Overall, the impact of GYSVd1 on grape quality varies significantly across different varieties, primarily influencing sugar content, acid, and phenolic compounds (Figure 8). Principal component analysis (PCA) revealed that Merlot berries experienced the most pronounced adverse effects from GYSVd1, and Welsch Riesling wine was, notably, the most detrimentally affected.
Recently, China has reported five kinds of viroids: Grapevine yellow speckle viroid 1 (GYSVd1), Grapevine yellow speckle viroid 2 (GYSVd2), Citrus exocortis viroid (CEVd), Hop stunt viroid (HSVd), and Australian grapevine viroid (AGVd). These viroids can not only cause infections individually but also coinfect with multiple viruses, leading to more complex disease manifestations. Viroids are infectious agents of plants, and despite their non-protein-coding RNA properties, they replicate autonomously in host cells [31]. Viroids are capable of both intercellular movement via cytodesmosis and long-distance invasion of the distal end of the host plant via phloem [32]. Their infection disrupts various cellular processes, including plant cell cycles, cell death pathways, macromolecule transport, cell signaling, protein turnover, transcriptional regulation, and defense mechanisms [9,33]. Viroids are mainly mechanically transmitted and usually have a pathogenic effect on infected plants. Some viroids are pathogenic, but others replicate only in the host without causing disease [34]. It was found that the infection range of viroid GYSVd1 was wide, and the infection rate was higher than those of other viruses in six grape varieties. It can be seen that although the viroid structure is simple, the infection ability is powerful. It may be related to the replication ability of viroid GYSVd1 in the host plant and its nucleotide mutation in adapting to the host’s new environment to enhance its infection and pathogenicity. In addition, q-PCR analysis revealed that GYSVd1 accumulation in infected Merlot and Riesling Weiss vines was significantly higher than in uninfected grapevines. In contrast, no significant difference in GYSVd1 accumulation was observed in other grape varieties compared to uninfected grapevine controls. This indicates that GYSVd1 exhibits distinct varietal specificity in its ability to infect different grape varieties. Notably, the Merlot showed higher GYSVd1 viroid accumulation, leading to a more pronounced negative impact on berry quality compared to other varieties. GYSVd1 likely disrupts the accumulation of key berry quality components by suppressing the accumulation of viroids involved in the biosynthesis of sugars and phenols, with the elevated accumulation of GYSVd1 in Merlot further exacerbating this inhibitory effect. Interestingly, although the berry quality of Merlot was significantly compromised by GYSVd1 infection, the quality of Merlot wine was less affected. This phenomenon may be attributed to the complex transformation of phenols, sugars, and other metabolites during the winemaking process. Further research is needed to elucidate the specific mechanisms underlying these observations.
The quality of grape berries and wines directly determines their economic value. The morphological characteristics of grape berries are an essential index for measuring quality; whether the berry is whole and tasty and whether the shape is beautiful is also a significant standard which consumers use to judge its quality [35]. Previous studies have found that infection with GLRaV-3 and GRBV can delay ripening, reduce berries’ weight [36], and reduce berries’ quality. However, this study found that viroid GYSVd1 had no significant effects on berries’ weight and shape index, indicating that different viruses might affect quality indexes differently. At the same time, the sugar and acid content and the TSSs are also critical indicators for measuring the quality of berries and wine, and TSS is an essential indicator of whether the berries are ripe [37]. GFLV infection significantly increased TSSs content [3]. There were some differences between GYSVd1 infection and the results of this experiment. The TSSs content in berries and wine from some grape varieties significantly increased, while those of some grape varieties significantly decreased. The reason might be that different grape varieties were affected differently by the viroid GYSVd1.
Sugar can be used not only as a primary energy material for plant growth and development but also as a raw material for synthesizing secondary metabolites to improve fruit quality [38]. Previous studies have demonstrated that viral infections significantly impact sugar metabolism in grapevines. For instance, coinfection with GLRaV-3, grapevine fleck virus (GFKV), and grapevine rupestris stem pitting-associated virus (GRSPaV) leads to a notable reduction in berries’ sugar content by suppressing the expression of genes involved in sugar synthesis [3]. Similarly, infection with GRBV alone has been shown to significantly decrease the sugar content in berries [39]. These research findings are consistent with the phenomenon observed in this study, where GYSVd1 infection led to a significant reduction in sugar content. Specifically, GYSVd1 infection markedly decreased the sugar levels in Chardonnay berries, Welsch Riesling berries and wine, as well as Riesling Weiss wine. These results suggest that GYSVd1 negatively regulates pathways associated with sugar synthesis and metabolism. However, the specific changes in gene expression responsible for the reduction in grape sugar content require further investigation to elucidate the underlying mechanisms. Acids can affect the color and taste of grape juice and processed products such as wine [40]. Tartaric acid is the primary acid in grape berries and wine, determining the fermented product’s flavor properties, taste, and aging potential [41]. In this study, viroid GYSVd1 had different effects on pH in different grape varieties; significantly reduced the pH of Merlot, Welsch Riesling berries, and Chardonnay wine; and significantly increased the pH of Merlot, Cabernet Sauvignon, and Syrah wine.
Phenolic substances in fruits are a class of essential nutrients that neutralize free radicals in the body, thereby reducing the oxidative stress-induced cell damage, and play an important role in protecting health and preventing diseases. The active components of phenolic substances are mainly phenolic compounds, including tannins, anthocyanins, and flavonoids. Anthocyanins are water-soluble natural pigments found in the peel and are one of the flavonoid compounds [42]. Previous studies have demonstrated that viral infections significantly impact the accumulation of phenolic compounds in grapes. For example, GLRaV-3 not only reduces the phenolic content in berries but also significantly decreases anthocyanin levels by suppressing the expression of genes involved in anthocyanin biosynthesis [7,8]. Similarly, GRBV infection has been shown to significantly decrease anthocyanin content [38]. These findings highlight the negative effects of viral infections on grape berry color and quality. In contrast to previous studies, the present research reveals that the viroid GYSVd1 exerts varying effects on phenolic content across different grape varieties. Specifically, GYSVd1 infection led to a significant increase in phenolic content in Cabernet Sauvignon, Riesling Weiss, Syrah, Chardonnay, and Welsch Riesling berries, whereas a notable decrease was observed in Merlot berries. These results underscore the variety-specific responses of grapevines to GYSVd1 infection. Such variability may be attributed to differences in genetic background, metabolic regulatory networks, and physiological adaptability to viroid infection among grape varieties. Future research should focus on elucidating the molecular regulatory mechanisms underlying these variety-specific responses to GYSVd1 infection, thereby providing deeper insights into the intrinsic factors driving these differences. In addition, the effects of viroid GYSVd1 on phenolic substances in berries and wine of the same grape variety were not precisely the same. It can be seen that the wine’s final quality was closely related to the quality of the berries, and the winemaking process also played an important role.

5. Conclusions

Regarding berry quality, the negative impact of GYSVd1 followed this decreasing order: Merlot > Chardonnay > Welsch Riesling > Syrah > Riesling Weiss > Cabernet Sauvignon. Similarly, for wine quality, the negative impact exhibited the following gradient: Welsch Riesling > Riesling Weiss > Chardonnay > Cabernet Sauvignon > Syrah > Merlot. The amount of phenolic substances in Cabernet Sauvignon, Riesling Weiss, Syrah, Chardonnay, and Welsch Riesling berries increased significantly, while the amount of phenolic substances in Merlot berries decreased significantly. The amount of sugar, acid, and phenolic substances decreased in Chardonnay, Welsch Riesling, Riesling Weiss wine, while the amount of acid and phenolic substances increased significantly in Merlot, Chardonnay, and Syrah wine. GYSVd1 affected the quality of berries and wine mainly by regulating the content of sugar acid and phenolic substances.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/horticulturae11040345/s1, Table S1: cDNA component; Table S2: Primers for detection of viruses and viroids; Table S3. Primers for fluorescent quantitative PCR

Author Contributions

M.W.: Investigation, Formal analysis, Writing—original draft. S.L.: Investigation, Formal analysis. P.W.: Methodology. Z.L.: Investigation. J.Z.: Investigation. Y.D.: Conceptualization, Supervision. S.Z.: Writing—review and editing, Supervision. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the National Natural Science Foundation of China (31860496, 32071808), Shandong Province Key R&D Program (Action Plan for Revitalizing Science and Technology Innovation and Revitalization of Rural Areas) (2022TZXD0023, 2023TZXD033).

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.

Acknowledgments

This work was significantly supported by academic assistance from the De partment of Shihezi University and Shandong Agricultural University, for which the authors are thankful. Without their help and efforts, this task could not have been finished.

Conflicts of Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

References

  1. Yue, X.; Ju, Y.; Zhang, T.; Yu, R.; Xu, H.; Zhang, Z. Application of salicylic acid to cv. Muscat Hamburg grapes for quality improvement: Effects on typical volatile aroma compounds and anthocyanin composition of grapes and wines. LWT 2025, 182, 1067–1080. [Google Scholar] [CrossRef]
  2. Karimi, R.; Koulivand, M.; Ollat, N. Soluble sugars, phenolic acids and antioxidant capacity of grape berries as affected by iron and nitrogen. Acta Physiol. Plant. 2019, 41, 974–985. [Google Scholar] [CrossRef]
  3. Ren, F.; Qiao, S.; Fan, X.; Hu, G.; Zhou, Z.; Dong, Y. Impacts of three virus combinations on grape composition and fruit quality of Vitis vinifera ‘Kyoho’. Hortic. Plant J. 2024, 45, 98–109. [Google Scholar] [CrossRef]
  4. Fuchs, M.; Rwahnih, M.A.; Blouin, A.G.; Burger, J.; Chooi, K.M.; Constable, F.; Várallyay, É. A list of eclectic viruses, virus-like diseases and viroids of grapevines that should not be considered for regulatory oversight: A global plea from virologists. J. Plant Pathol. 2025, 25, 876–882. [Google Scholar] [CrossRef]
  5. Umer, M.; Liu, J.; You, H.; Xu, C.; Dong, K.; Luo, N.; Kong, L.; Li, X.; Hong, N.; Wang, G.; et al. Genomic, Morphological and Biological Traits of the Viruses Infecting Major Fruit Trees. Viruses 2019, 11, 515. [Google Scholar] [CrossRef]
  6. Chiaki, Y.; Ito, T.; Sato, A.; Sugiura, H.; Nishimura, R. Dwarfing caused by viral pathogens and leaf malformations in ‘Shine Muscat’ grapevine. J. Gen. Plant Pathol. 2019, 86, 34–38. [Google Scholar] [CrossRef]
  7. Lee, J.; Rennaker, C.D.; Thompson, B.D.; Dahan, J.; Karasev, A.V. Idaho ‘Cabernet Sauvignon’ grape composition altered by grapevine leafroll-associated virus 3. NFS J. 2023, 31, 1–6. [Google Scholar]
  8. Vega, A.; Gutiérrez, R.A.; Peña-Neira, A.; Cramer, G.R.; Arce-Johnson, P. Compatible GLRaV-3 viral infections affect berry ripening decreasing sugar accumulation and anthocyanin biosynthesis in Vitis vinifera. Plant Mol. Biol. 2011, 77, 261–274. [Google Scholar]
  9. Blanco-Ulate, B.; Hopfer, H.; Figueroa-Balderas, R.; Ye, Z.; Rivero, R.M.; Albacete, A.; Pérez-Alfocea, F.; Koyama, R.; Anderson, M.M.; Smith, R.J.; et al. Red blotch disease alters grape berry development and metabolism by interfering with the transcriptional and hormonal regulation of ripening. J. Exp. Bot. 2017, 68, 1225–1238. [Google Scholar]
  10. Pereira, G.E.; Padhi EM, T.; Sudarshana, M.R.; Fialho, F.B.; Medina-Plaza, C.; Girardello, R.C.; Tseng, D.; Bruce, R.C.; Erdmann, J.N.; Slupsky, C.M.; et al. Impact of grapevine red blotch disease on primary and secondary metabolites in ‘Cabernet Sauvignon’ grape tissues. Food Chem. 2021, 342, 1278–1284. [Google Scholar]
  11. Martínez, L.; Miranda, C.; Royo, J.B.; Urrestarazu, J.; Martínez de Toda, F.; Balda, P.; Santesteban, L.G. Direct and indirect effects of three virus infections on yield and berry composition in grapevine (Vitis vinifera L.) cv. ‘Tempranillo’. Sci. Hortic. 2016, 212, 20–28. [Google Scholar] [CrossRef]
  12. Matoušek, J.; Siglová, K.; Jakše, J.; Radišek, S.; Brass JR, J.; Tsushima, T.; Guček, T.; Duraisamy, G.S.; Sano, T.; Steger, G. Propagation and some physiological effects of Citrus bark cracking viroid and Apple fruit crinkle viroid in multiple infected hop (Humulus lupulus L.). J. Plant Physiol. 2017, 213, 166–177. [Google Scholar] [CrossRef] [PubMed]
  13. Palukaitis, P. What has been happening with viroids? Virus Genes 2014, 49, 175–184. [Google Scholar] [CrossRef] [PubMed]
  14. Yan, J.; Tang, J.; Perez-Egusquiza, Z.; Thompson, J.R. Development of TaqMan RT-qPCR for the detection of regulated citrus viruses and viroids in Aotearoa New Zealand. J. Virol. Methods 2024, 327, 56–62. [Google Scholar] [CrossRef]
  15. Salman, T.M.; Habili, N.; Shi, B. Effect of temperature on symptom expression and sequence polymorphism of grapevine yellow speckle viroid 1 in grapevine. Virus Res. 2014, 189, 243–247. [Google Scholar] [CrossRef]
  16. Zhou, Y.; Cheng, Y.; Zhong, R.; Tang, J.; Pervaiz, T.; Zhou, S.; Liu, J.; Wang, B.; Jia, H. Brassinolide and gibberellin promote grape fruit development and quality. Sci. Hortic. 2024, 338, 76–84. [Google Scholar] [CrossRef]
  17. Algul, B.E.; Al Shoffe, Y.; Park, D.; Cheng, L.; Watkins, C.B. Preharvest 1-methylcyclopropene and aminoethoxyvinylglycine treatment effects on ‘NY2’ (RubyFrost®) apple fruit quality and postharvest watercore dissipation at different temperatures. Postharvest Biol. Technol. 2025, 220, 2674–2689. [Google Scholar] [CrossRef]
  18. Owoyemi, A.; Balaklav, M.; Kochanek, B.; Porat, R.; Koenigstein, N.; Salzer, Y.; Lichter, A. Deviations from optimal storage temperature and its impact on postharvest quality of table grape cv. Scarlotta Seedless. Postharvest Biol. Technol. 2024, 215, 3401–3412. [Google Scholar] [CrossRef]
  19. Palma-López, J.; Sánchez-Rodríguez, A.R.; del Campillo, M.C.; León-Gutiérrez, J.M.; Ramírez-Pérez, P. Influence of soil properties on grape and must quality in the Montilla − Moriles protected designation of origin (southern Spain). Catena 2024, 241, 67–79. [Google Scholar] [CrossRef]
  20. Yang, Y.; Huang, Z.; Wu, Y.; Wu, W.; Lyu, L.; Li, W. Combined transcriptomic and metabolomic analysis of the effect of different nitrogen application levels on soluble sugar and organic acid accumulation in blackberry fruit. Sci. Hortic. 2024, 338, 3485–3495. [Google Scholar] [CrossRef]
  21. Maliogka, V.I.; Olmos, A.; Pappi, P.G.; Lotos, L.; Efthimiou, K.; Grammatikaki, G.; Candresse, T.; Katis, N.I.; Avgelis, A.D. A novel grapevine badnavirus is associated with the Roditis leaf discoloration disease. Virus Res. 2015, 203, 47–55. [Google Scholar] [PubMed]
  22. Su, J.; Li, M.; Yang, H.; Shu, H.; Yu, K.; Cao, H.; Xu, G.; Wang, M.; Zhu, Y.; Zhu, Y.; et al. Enrichment of grape berries and tomato fruit with health-promoting tartaric acid by expression of the Vitis vinifera transketolase VvTK2 gene. Int. J. Biol. Macromol. 2024, 257, 734–745. [Google Scholar] [CrossRef] [PubMed]
  23. Liu, M.; Ji, H.; Jiang, Q.; Liu, T.; Cao, H.; Zhang, Z. Effects of full shading of clusters from véraison to ripeness on fruit quality and volatile compounds in Cabernet Sauvignon grapes. Food Chem. X 2024, 21, 1256–1263. [Google Scholar]
  24. Chen, M.; Zhang, S.; Ren, Y.; Le, Z.; Li, L.; Sun, B. Effects of Different Brewing Technologies on Polyphenols and Aroma Components of Black Chokeberry Wine. Foods 2023, 12, 868. [Google Scholar] [CrossRef]
  25. Sultanova, N.; Bayramova, N.; Aliyeva, D.; Rastgou, M.; Huseynova, I. Induced changes in metabolic constituents of grapevine (Vitis vinifera L.) leaves infected with grapevine leafroll-associated virus-3. Physiol. Mol. Plant Pathol. 2019, 106, 57–63. [Google Scholar]
  26. Zhang, B.; Zhang, C.; Chen, J.; Zhao, C.; Du, Y.; Yang, Y.; Xie, X.; He, L.; Liu, S.; Shi, K. On-vine drying (passérillage) improves the quality of “Hutai No. 8” table grape wine: Focusing on phenolics, aromas, color and sensory attributes. Food Chem. 2025, 463, 975–986. [Google Scholar] [CrossRef]
  27. Dravie, E.E.; Kortei, N.K.; Essuman, E.K.; Tettey, C.O.; Boakye, A.A.; Hunkpe, G. Antioxidant, phytochemical and physicochemical properties of sesame seed (Sesamum indicum L). Sci. Afr. 2020, 8, 451–463. [Google Scholar]
  28. Shi, M.; Zhang, Y.; Zhang, T.; Zhang, W.; Wang, S.; Wei, M.; Wang, S.; Zhao, L. The NAC activator, MdNAC77L, regulates anthocyanin accumulation in red flesh apple. Hortic. Plant J. 2024, 45, 1221–1231. [Google Scholar]
  29. Shang, Y.-F.; Cao, H.; Ma, Y.-L.; Zhang, C.; Ma, F.; Wang, C.-X.; Ni, X.-L.; Lee, W.-J.; Wei, Z.-J. Effect of lactic acid bacteria fermentation on tannins removal in Xuan Mugua fruits. Food Chem. 2019, 274, 118–122. [Google Scholar]
  30. Xiao, Q.; Ye, S.; Wang, H.; Xing, S.; Zhu, W.; Zhang, H.; Zhu, J.; Pu, C.; Zhao, D.; Zhou, Q.; et al. Soluble sugar, organic acid and phenolic composition and flavor evaluation of plum fruits. Food Chem. X 2024, 24, 232–239. [Google Scholar]
  31. Hadidi, A.; Randles, J.W. Viroids, and the Legacy of Ricardo Flores (1947–2020). Cells 2021, 10, 2570. [Google Scholar] [CrossRef] [PubMed]
  32. Lee, B.D.; Koonin, E.V. Viroids and Viroid-like Circular RNAs: Do They Descend from Primordial Replicators? Life 2022, 12, 103. [Google Scholar] [CrossRef] [PubMed]
  33. Pallas, V.; García, J.A. How do plant viruses induce disease? Interactions and interference with host components. J. Gen. Virol. 2011, 92, 2691–2705. [Google Scholar] [CrossRef] [PubMed]
  34. Owens, R.A.; Tech, K.B.; Shao, J.Y.; Sano, T.; Baker, C.J. Global Analysis of Tomato Gene Expression During Potato spindle tuber viroid Infection Reveals a Complex Array of Changes Affecting Hormone Signaling. e-Xtra 2011, 25, 583–595. [Google Scholar] [CrossRef]
  35. Leng, X.; Li, J.; Miao, W.; Liu, Y.; Haider, M.S.; Song, M.; Li, Q. Comparison of physicochemical characteristics, antioxidant and immunomodulatory activities of polysaccharides from wine grapes. Int. J. Biol. Macromol. 2023, 239, 589–595. [Google Scholar] [CrossRef]
  36. Montero, R.; Mundy, D.; Albright, A.; Grose, C.; Trought MC, T.; Cohen, D.; Chooi, K.M.; MacDiarmid, R.; Flexas, J.; Bota, J. Effects of Grapevine Leafroll associated Virus 3 (GLRaV-3) and duration of infection on fruit composition and wine chemical profile of Vitis vinifera L. cv. Sauvignon blanc. Food Chem. 2016, 197, 1177–1183. [Google Scholar]
  37. Saenphon, C.; Ditcharoen, S.; Malai, C.; Saengprachatanarug, K.; Wongpichet, S.; Sirisomboon, P.; Saechua, W.; Khurnpoon, L.; Phuphaphud, A.; Maraphum, K.; et al. Total soluble solids, dry matter content prediction and maturity stage classification of durian fruit using long-wavelength NIR reflectance. J. Food Compos. Anal. 2023, 124, 3421–3436. [Google Scholar]
  38. White, A.C.; Rogers, A.; Rees, M.; Osborne, C.P. How can we make plants grow faster? A source–sink perspective on growth rate. J. Exp. Bot. 2016, 67, 31–45. [Google Scholar] [CrossRef]
  39. Lee, J.; Rennaker, C.D.; Thompson, B.D.; Karasev, A.V. Influence of Grapevine red blotch virus (GRBV) on Idaho ‘Syrah’ grape composition. Sci. Hortic. 2021, 282, 548–557. [Google Scholar] [CrossRef]
  40. DeBolt, S.; Cook, D.R.; Ford, C.M. L-Tartaric acid synthesis from vitamin C in higher plants. PNAS 2006, 103, 5608–5613. [Google Scholar]
  41. Borsani, J.; Budde, C.O.; Porrini, L.; Lauxmann, M.A.; Lombardo, V.A.; Murray, R.; Andreo, C.S.; Drincovich, M.F.; Lara, M.V. Carbon metabolism of peach fruit after harvest: Changes in enzymes involved in organic acid and sugar level modifications. J. Exp. Bot. 2009, 60, 1823–1837. [Google Scholar]
  42. Wang, Y.; Zhang, Z.; Wang, X.; Yuan, X.; Wu, Q.; Chen, S.; Zou, Y.; Ma, F.; Li, C. Exogenous dopamine improves apple fruit quality via increasing flavonoids and soluble sugar contents. Sci. Hortic. 2021, 280, 346–361. [Google Scholar]
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Figure 1. The detection rate of GYSVd1 in 6 grape varieties.
Figure 1. The detection rate of GYSVd1 in 6 grape varieties.
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Figure 2. Accumulation of GYSVd1 in the berries of Merlot (A), Cabernet Sauvignon (B), Syrah (C), Chardonnay (D), Welsch Riesling (E) and Riesling Weiss (F). *** means p ≤ 0.001, * means p ≤ 0.05, and ns means p > 0.05.
Figure 2. Accumulation of GYSVd1 in the berries of Merlot (A), Cabernet Sauvignon (B), Syrah (C), Chardonnay (D), Welsch Riesling (E) and Riesling Weiss (F). *** means p ≤ 0.001, * means p ≤ 0.05, and ns means p > 0.05.
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Figure 3. Effect of GYSVd1 on the total phenolic compounds content in Merlot (A,G), Cabernet Sauvignon (B,H), Syrah (C,I), Chardonnay (D,J), Welsch Riesling (E,K), Riesling Weiss (F,L) berries, and wine. *** means p ≤ 0.001; * means p ≤ 0.05; ns means p > 0.05.
Figure 3. Effect of GYSVd1 on the total phenolic compounds content in Merlot (A,G), Cabernet Sauvignon (B,H), Syrah (C,I), Chardonnay (D,J), Welsch Riesling (E,K), Riesling Weiss (F,L) berries, and wine. *** means p ≤ 0.001; * means p ≤ 0.05; ns means p > 0.05.
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Figure 4. Effect of GYSVd1 on the anthocyanin content in Merlot (A,D), Cabernet Sauvignon (B,E), Syrah (C,F) berries, and wine. ** means p ≤ 0.01; * means p ≤ 0.05; ns means p > 0.05.
Figure 4. Effect of GYSVd1 on the anthocyanin content in Merlot (A,D), Cabernet Sauvignon (B,E), Syrah (C,F) berries, and wine. ** means p ≤ 0.01; * means p ≤ 0.05; ns means p > 0.05.
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Figure 5. Effect of GYSVd1 on the tannin content in Merlot (A,G), Cabernet Sauvignon (B,H), Syrah (C,I), Chardonnay (D,J), Welsch Riesling (E,K), Riesling Weiss (F,L) berries, and wine. *** means p ≤ 0.001; * means p ≤ 0.05; ns means p > 0.05.
Figure 5. Effect of GYSVd1 on the tannin content in Merlot (A,G), Cabernet Sauvignon (B,H), Syrah (C,I), Chardonnay (D,J), Welsch Riesling (E,K), Riesling Weiss (F,L) berries, and wine. *** means p ≤ 0.001; * means p ≤ 0.05; ns means p > 0.05.
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Figure 6. Effect of GYSVd1 on the soluble sugar content in Merlot (A,G), Cabernet Sauvignon (B,H), Syrah (C,I), Chardonnay (D,J), Welsch Riesling (E,K), Riesling Weiss (F,L) berries, and wine. *** means p ≤ 0.001; * means p ≤ 0.05, ns means p > 0.05.
Figure 6. Effect of GYSVd1 on the soluble sugar content in Merlot (A,G), Cabernet Sauvignon (B,H), Syrah (C,I), Chardonnay (D,J), Welsch Riesling (E,K), Riesling Weiss (F,L) berries, and wine. *** means p ≤ 0.001; * means p ≤ 0.05, ns means p > 0.05.
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Figure 7. Principal component analysis (PCA) of grape berries and wine quality indexes. (A,D): heatmap of sample correlations; the figures represent Pearson’s correlation coefficient (R2); (B,E): PCA scree plot; (C,F): PCA score plot and load plot. 1: UF-Me, 2: GYSVd1-Me, 3: UF-CS, 4: GYSVd1-CS, 5: UF-Sy, 6: GYSVd1-Sy, 7: UF-Ch, 8: GYSVd1-Ch, 9: UF-WR, 10: GYSVd1-WR, 11: UF-RW, 12: GYSVd1-RW. * means p ≤ 0.05.
Figure 7. Principal component analysis (PCA) of grape berries and wine quality indexes. (A,D): heatmap of sample correlations; the figures represent Pearson’s correlation coefficient (R2); (B,E): PCA scree plot; (C,F): PCA score plot and load plot. 1: UF-Me, 2: GYSVd1-Me, 3: UF-CS, 4: GYSVd1-CS, 5: UF-Sy, 6: GYSVd1-Sy, 7: UF-Ch, 8: GYSVd1-Ch, 9: UF-WR, 10: GYSVd1-WR, 11: UF-RW, 12: GYSVd1-RW. * means p ≤ 0.05.
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Figure 8. Effects of viroid GYSVd1 on grape berries and wine quality. Blue arrow: GYSVd1-infected berries and wine showed a significant decrease in a specific quality indicator compared to uninfected samples. Red arrow: GYSVd1-infected berries and wine showed a significant increase in a specific quality indicator compared to uninfected samples.
Figure 8. Effects of viroid GYSVd1 on grape berries and wine quality. Blue arrow: GYSVd1-infected berries and wine showed a significant decrease in a specific quality indicator compared to uninfected samples. Red arrow: GYSVd1-infected berries and wine showed a significant increase in a specific quality indicator compared to uninfected samples.
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Table 1. Determination of general physicochemical indexes of berries and wine of 6 wine grape varieties.
Table 1. Determination of general physicochemical indexes of berries and wine of 6 wine grape varieties.
Grape VarietyBerryWine
WeightShape IndexpHTSSpHAlcoholic Strength
MerlotUF-Me1.21 ± 0.331.04 ± 0.054.22 ± 0.0823.13 ± 0.483.80 ± 0.0510.00 ± 0.00
GYSVd11.09 ± 0.26 ns1.01 ± 0.03 ns3.96 ± 0.08 ***22.05 ± 0.92 *3.95 ± 0.07 ***11.04 ± 0.83 ns
Cabernet SauvignonUF-CS1.11 ± 0.310.99 ± 0.024.07 ± 0.0124.07 ± 0.133.96 ± 0.018.80 ± 0.10
GYSVd11.35 ± 0.20 ns1.01 ± 0.03 ns4.03 ± 0.13 ns21.42 ± 2.15 ***4.03 ± 0.06 **9.92 ± 0.50 ns
SyrahUF-Sy1.91 ± 0.260.97 ± 0.043.77 ± 0.0121.60 ± 0.263.70 ± 0.0111.57 ± 0.15
GYSVd11.66 ± 0.38 ns0.95 ± 0.04 ns3.76 ± 0.05 ns23.10 ± 1.07 ***3.77 ± 0.05 ***10.82 ± 0.67 ns
ChardonnayUF-Ch1.43 ± 0.191.00 ± 0.043.54 ± 0.0120.66 ± 0.343.92 ± 0.0111.03 ± 0.25
GYSVd11.44 ± 0.26 ns0.99 ± 0.02 ns3.88 ± 0.15 ns23.16 ± 0.05 ***3.90 ± 0.18 ns9.42 ± 1.17 ns
Welsch RieslingUF-WR1.05 ± 0.390.97 ± 0.054.35 ± 0.0325.90 ± 0.684.24 ± 0.029.97 ± 0.25
GYSVd11.16 ± 0.25 ns0.95 ± 0.04 ns4.30 ± 0.06 ***20.80 ± 0.79 ***4.27 ± 0.12 ns7.42 ± 0.64 ns
Riesling WeissUF-RW1.14 ± 0.310.98 ± 0.044.33 ± 0.0920.57 ± 0.303.95 ± 0.0111.57 ± 0.15
GYSVd11.22 ± 0.31 ns1.00 ± 0.03 ns4.21 ± 0.12 ns19.85 ± 1.70 ns3.93 ± 0.08 ns10.82 ± 0.67 ns
Note: TSS: total soluble solid. *** means p ≤ 0.001, ** means p ≤ 0.01, * means p ≤ 0.05, and ns means p > 0.05.
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Wu, M.; Liu, S.; Wang, P.; Li, Z.; Zhang, J.; Du, Y.; Zhu, S. Viroid GYSVd1 Exhibited Different Regulations on the Qualities of Berries and Wines from 6 Grape Varieties. Horticulturae 2025, 11, 345. https://doi.org/10.3390/horticulturae11040345

AMA Style

Wu M, Liu S, Wang P, Li Z, Zhang J, Du Y, Zhu S. Viroid GYSVd1 Exhibited Different Regulations on the Qualities of Berries and Wines from 6 Grape Varieties. Horticulturae. 2025; 11(4):345. https://doi.org/10.3390/horticulturae11040345

Chicago/Turabian Style

Wu, Menghuan, Shuo Liu, Ping Wang, Zhaotan Li, Junbo Zhang, Yejuan Du, and Shuhua Zhu. 2025. "Viroid GYSVd1 Exhibited Different Regulations on the Qualities of Berries and Wines from 6 Grape Varieties" Horticulturae 11, no. 4: 345. https://doi.org/10.3390/horticulturae11040345

APA Style

Wu, M., Liu, S., Wang, P., Li, Z., Zhang, J., Du, Y., & Zhu, S. (2025). Viroid GYSVd1 Exhibited Different Regulations on the Qualities of Berries and Wines from 6 Grape Varieties. Horticulturae, 11(4), 345. https://doi.org/10.3390/horticulturae11040345

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