ENOLOGY / Review article

Relationships between wine quality and substances exceeding the odour threshold produced by Hanseniaspora

Andrea Caridi, Marco Poiana
Received : 16 July 2025; Accepted : 20 April 2026; Published : 1 June 2026
DOI: https://doi.org/10.20870/oeno-one.2026.60.2.9494

Abstract

Several recent studies propose the use of yeasts from the Hanseniaspora genus in winemaking. In this paper, the negative and positive wine components due to the activity of these yeasts are listed and critically analysed. Only the substances produced in amounts exceeding their odour threshold are considered. This review highlights the relationships between the use of Hanseniaspora strains in winemaking and the quality of the wines so obtained. First of all, the acetic acid concentration in grape wines is very important; legal limits for its content in wine vary by country and wine type. A moderate production of ethyl acetate positively contributes to the sensory quality of the wines. Isobutanol is related to an unpleasant aroma. Acetoin, 1-hexanol, decanoic acid, octanoic acid, and hexanoic acid: the less, the better. Isoamyl alcohol may positively contribute to the complexity of the aroma of wines. 2-Phenylethanol contributes to fine rose notes in wine aroma. 2-Phenylethyl acetate is a compound that imparts flowery, rosy, and honey-like fragrances with fruity undertones to wines. Isoamyl acetate is one of the esters most markedly contributing to the aroma profile of white wines. Ethyl hexanoate is responsible for tropical fruity attributes. Hexyl acetate, ethyl decanoate, ethyl butyrate, β-citronellol, linalool, ethyl octanoate, β-damascenone: the more the merrier.

Introduction

The single most important factor in winemaking is the sensory quality of the final product (Plata et al., 2003). Even if the grape wine flavour cannot be totally predicted, its volatile profile is dominated by the components exceeding their odour thresholds. This is true both for positive and negative effects, depending on the aromatic profile of each component. Consequently, it is important that the positive components are over their odour thresholds and that the negative components are under their odour thresholds. Recently, many strains of the genus Hanseniaspora were proposed as starter and/or adjunct cultures for grape wine production; however, the concentration of the crucial volatile components was not always reported. Furthermore, it is important to note that the pure cultures of Hanseniaspora are always unable to finish the fermentation, resulting in unfermented sugar in the grape wines. Considering this, several methodologies are listed below to study the fermentative activity of Hanseniaspora strains in grape juices.

  • Fermentations were carried out for a period of 12 days (Valera et al., 2021). Fermentation kinetics were controlled as CO2 liberated by daily weighing. After 12 days of fermentation, samples were analysed, and residual sugars were quantified.
  • Fermentations were carried out for a period of 14 days (Andorrà et  al., 2010). Sugar consumption was monitored daily; fermentations were considered to be finished when the level of reducing sugars was below 2 g L–1. However, the pure H. uvarum culture did not finish the fermentation (20 g L–1 of residual glucose).
  • Fermentations were carried out for a period of 18 days (Testa et al., 2021). The alcoholic fermentation was monitored considering the variation of reducing sugars and ethanol content.
  • Fermentations were carried out for a period of 20 days (del Fresno et al., 2025). Fermentation was monitored daily measuring the decrease in density, which indicated the correct progress of the alcoholic fermentation until its completion.
  • Fermentations were carried out for a maximum period of 30 days (Cordero-Bueso et al., 2013). When sugar content was lower than 2 g L–1 or after 30 days of fermentation.
  • Fermentations were carried out to dryness for a maximum period of 180 days (du Plessis et al., 2017). The final wine chemical analyses were carried out when the wines showed residual sugar < 4 g L–1.
  • Fermentations were carried out without indicating the days of fermentation (de Benedictis et al., 2011). The kinetics of the fermentations were monitored by evaluating daily the loss of weight due to the production of CO2; when the CO2 evolution stopped (i.e., at constant weight), the samples were directly analysed or stored at –20 °C.

A quick review about negative and positive components related to Hanseniaspora activity in grape wines will now be carried out, reporting, in decreasing order of odour threshold, the components exceeding their odour threshold. Only the values that were higher than those produced by the control have been reported, i.e., spontaneous fermentation or fermentation using selected strains of Saccharomyces cerevisiae.

1. Negative components/off-flavours

1.1. Acetic acid

Several authors have tested yeasts of the genus Hanseniaspora in winemaking, but they have not reported the concentration of acetic acid in the produced wines (Bagheri et al., 2018; Hu et al., 2018; Olivera et al., 2024). Since, in our opinion, this parameter is essential for the use of Hanseniaspora in winemaking, the results of these papers have not been taken into consideration in this article. The aromatic descriptor of this compound is “vinegar”, its odour threshold is 700 mg L–1, and the presence of acetic acid at levels above the perception threshold is undesirable (Carrau et al., 2023).

The following papers reported acetic acid contents, i.e., volatile acidity, exceeding the odour threshold, thereby negatively affecting the sensory quality of the wine; all the other papers described in this article obtained, using Hanseniaspora, wines with acetic acid content below 700 mg L–1.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with: a) 106 cells mL–1 of H. uvarum HuB10 contained 37,500 mg L–1; b) 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 1580 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 490 mg L–1.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. osmophila NRRL-Y1613 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 1230 mg L–1; b) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 1100 mg L–1; c) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 940 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 840 mg L–1.

Cordero-Bueso et al., 2013 – Grape wine produced at 20 °C from Malvar grape must filter-sterilised inoculated with 106 cells mL–1 of H. guilliermondii CLI 921 contained 900 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae) contained 140 mg L–1.

de Benedictis et al., 2011 – Grape wine produced at 25 °C from Negroamaro grape must clarified by centrifugation, sterilised by membrane filtration, and inoculated with 106 cells mL–1 of: a) H. uvarum 8797 contained 780 mg L–1; b) H. uvarum 8799 contained 740 mg L–1; c) H. uvarum 8801 contained 750 mg L–1; d) H. uvarum 8802 contained 730 mg L–1; e) H. uvarum 8807 contained 810 mg L–1; f) H. uvarum 8810 contained 880 mg L–1; g) H. uvarum 8811 contained 950 mg L–1; h) H. uvarum 8813 contained 940 mg L–1; i) H. uvarum CBS314 contained 1220 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CM) contained 550 mg L–1.

du Plessis et al., 2017 – Synthetic wine produced at 22 °C from chemically defined grape juice, filter-sterilised and inoculated with: a) H. uvarum H2 contained 720 mg L–1; b) H. uvarum H3 contained 840 mg L–1; c) H. uvarum H10 contained 880 mg L–1; the control (inoculated with S. cerevisiae S3) contained 290 mg L–1.

Filippousi et al., 2024 – Grape wine produced at 16 °C from Sideritis grape must unfiltered, added with potassium metabisulfite to get total sulfur dioxide of 30 ppm and inoculated with 107 cells mL–1 of H. opuntiae L1 and, after eight hours, with 105 cells mL–1 of S. cerevisiae W7 contained 770 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae W7) contained 640 mg L–1.

Valera et al., 2021 – Grape wine produced at 20 °C from Chardonnay grape must, treated with 200 mg L–1 dimethyldicarbonate and inoculated with 105 cells mL–1 of H. uvarum AWRI1280, contained 910 mg L–1; the control (inoculated with 105 cells mL–1 of S. cerevisiae ALG804) contained 450 mg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and 106 cells mL–1 of S. cerevisiae CVE-SC45 contained 1660 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 1640 mg L–1.

Final commentary: the acetic acid concentration in grape wines is very important. Legal limits for acetic acid content in wine vary by country and wine type. Sensory detection often starts around 0.6–0.9 g L–1.

1.2. Ethyl acetate

Ethyl acetate is the most abundant ester in wine (Plata et al., 2003); its odour threshold is 7.5 mg L–1 and it can be described as “apple, fruity” (Lai et al., 2023). At low levels, it confers aromatic complexity to the wine, giving it a “fruity” aroma (Godoy et al., 2020); instead, according to Viana et al. (2008), ethyl acetate in wine can impart spoilage character at levels of 150–200 mg L–1. Some strains of Hanseniaspora were reported to produce ethyl acetate over these levels, thereby negatively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 558.07 mg L–1; b) 107 cells mL–1 of H. meyeri NRRL-Y27513 and 106 cells of S. cerevisiae Uvaferm CEG contained 332.86 mg L–1; c) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 318.93 mg L–1; d) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 312.98 mg L–1; e) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 291.07 mg L–1; f) 107 cells mL–1 of H. osmophila NRRL-Y1613 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 151.6 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 115.8 mg L–1.

Cordero-Bueso et al., 2013 – Grape wine produced as previously reported contained 874.4 mg L–1; the control contained 39.5 mg L–1.

de Benedictis et al., 2011 – Grape wine produced at 25 °C from Negroamaro grape must clarified by centrifugation, sterilised by membrane filtration, and inoculated with 106 cells mL–1 of: a) H. uvarum 8801 contained 151.7 mg L–1; b) H. uvarum 8807 contained 150.6 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CM) contained 14.3 mg L–1.

Filippousi et al., 2024 – Grape wine produced as previously reported contained 585.87 mg L–1; the control contained 170.04 mg L–1.

Gallo et al., 2023 – Grape wine produced at 18–20 °C from Chardonnay grape must, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Hv205 and, in sequential inoculation at 30 % of the alcoholic fermentation, with 5 × 106 cells mL–1 of S. cerevisiae LVCB contained 184 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae LVCB) contained 142 mg L–1.

Li et al., 2022 – Grape wine produced from Sauvignon blanc grape must added with sulfur dioxide (30 mg L–1) in the form of potassium metabisulfite and 3-isobutyl-2-methoxypyrazine (52 ng L–1), and inoculated with 5 × 106 cells mL–1 of H. uvarum H11_G1_2, kept at 20 °C for 5 days, transferred to 16 °C and inoculated with S. cerevisiae Lalvin EC1118™, contained 248.15 mg L–1; the control (inoculated with S. cerevisiae Lalvin EC1118™) contained 74.99 mg L–1.

Final commentary: a moderate production of ethyl acetate positively contributes to the wines’ sensory quality.

1.3. Acetoin (i.e., 3-hydroxybutan-2-one)

Acetoin affects the wine bouquet; its odour threshold is 150 mg L–1 and it is described as being “creamy, butter, fat” (Carrau et al., 2023). One strain of Hanseniaspora was reported to produce acetoin exceeding the odour threshold, so negatively contributing to the wines’ sensory quality.

Cordero-Bueso et al., 2013 – Grape wine produced as previously reported contained 189.2 mg L–1; the control contained 6.9 mg L–1.

Final commentary: acetoin – the less, the better.

1.4. Isobutanol (i.e., 2-methyl-1-propanol or isobutyl alcohol)

At least four different descriptors, all with the odour threshold of 40 mg L–1, were reported: “fusel, alcohol” (Guth, 1997), “wine, solvent, bitter” (Testa et al., 2021), “fusel oil, chemical” (Carrau et al., 2023), and “solvent, chemical alcoholic, malt notes, wineosity notes” (Tarko & Duda, 2024). Some strains of Hanseniaspora were reported to produce isobutanol levels exceeding the odour threshold, thereby negatively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cell  ml–1 of S. cerevisiae QA23 contained 55.39 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 24.51 mg L–1.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 190.04 mg L–1; b) 107 cells mL–1 of H. meyeri NRRL-Y27513 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 161.05 mg L–1; c) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 150.83 mg L–1; d) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 149.54 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 141.8 mg L–1.

de Benedictis et al., 2011 – Grape wine produced at 25 °C from Negroamaro grape must clarified by centrifugation, sterilised by membrane filtration, and inoculated with 106 cells mL–1 of: a) H. uvarum 8799 contained 54.2 mg L–1; b) H. uvarum 8801 contained 53.9 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CM) contained 12.1 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with 107 cells mL–1 of H. vineae T02/5A contained 42.52 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 34.04 mg L–1.

Domizio et al., 2011 – Grape wine produced by inoculating commercial white grape juice with 107 cells mL–1 of H. osmophila Ha32 and 107 cells of S. cerevisiae EC1118 contained 83.15 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae EC1118) contained 69.55 mg L–1 (trial 1).

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and 106 cells mL–1 of S. cerevisiae RV002 contained 130.47 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae RV002) contained 121.644 mg L–1.

Pietrafesa et al., 2020 – Grape wine produced at 26 °C from Merlot grape must sterilised by filtration and inoculated with 107 cells mL–1 of H. uvarum H2 and 103 cells mL–1 of S. cerevisiae S3 contained 47.96 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae S3) contained 44.58 mg L–1.

Testa et al., 2020 – Grape wine produced at 18 °C from Fiano grape must cold clarified at 0 °C for 48 h, added with potassium metabisulfite (50 mg L–1), inoculated with 106 cells mL–1 of H. guilliermondii BF1 and, after two days, with 106 cells mL–1 of S. cerevisiae 404 contained 61.2 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae 404) contained 43.3 mg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and 106 cells mL–1 of S. cerevisiae CVE-SC45 contained 83.821 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 55.944 mg L–1.

Final commentary: isobutanol has an unpleasant aroma (Mestre et al., 2019).

1.5. 1-Hexanol

At least two descriptors were reported: “green, grass” (Guth, 1997) and “herbaceous, grass, woody” (Peinado et al., 2004); the odour threshold is 1.1 mg L–1 (Peinado et al., 2004). Some strains of Hanseniaspora were reported to produce 1-hexanol exceeding the odour threshold, so negatively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 1431.11 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 1411.9 mg L–1.

del Fresno et al., 2021a – Grape wine produced in stainless steel barrels at 18 °C using a mixture of 50 % Tempranillo and 50 % Albillo Mayor grapes; the grape must, containing 15.05 mg L–1 of total sulfur dioxide, was inoculated with 107 cells mL–1 of H. vineae T02/5A and contained 4.17 mg L–1; the control (inoculated with S. cerevisiae Fermivin 3C) contained 4.46 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with: a) 107 cells mL–1 of H. vineae T02/5A contained 4.06 mg L–1 of hexanol, b) 107 cells mL–1 of H. opuntiae A56 contained 3.89 mg L–1 of hexanol; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 4.42 mg L–1 of hexanol.

Filippousi et al., 2024 – Grape wine produced as previously reported contained 66.84 mg L–1; the control contained 49.9 mg L–1.

Luan et al., 2018 – Grape wine produced at 25 °C from Cabernet-Sauvignon must added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. opuntiae CVE-Ho11 and 105 cells mL–1 of S. cerevisiae BDX contained 2.21 mg L–1; the control (inoculated with 105 cells mL–1 of S. cerevisiae BDX) contained 1.82 mg L–1.

Yan et al., 2020 – Grape wine produced at 25 °C from French Colombard grape must pasteurised and inoculated with 107 cells mL–1 of H. vineae 71-97 and 106 cells mL–1 of S. cerevisiae var. bayanus EC1118 contained 1.19 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae var. bayanus EC1118) contained 0.77 mg L–1.

Final commentary:1-hexanol – the less, the better.

1.6. Decanoic acid (i.e., capric acid)

At least two descriptors were reported: “fatty, unpleasant” (Guth, 1997) and “rancid, fatty” (Carrau et al., 2023); its odour threshold is 1.0 mg L–1 (Carrau et al., 2023). At least three strains of Hanseniaspora were reported to produce decanoic acid exceeding the odour threshold, thereby negatively contributing to the sensory quality of the wine.

Gallo et al., 2024a – Grape wine produced using a mixture of 50 % Glera and 50 % Gewurztraminer grapes, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Fermivin VINEAE and, after 48 hours, with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB contained 1.955 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 0.726 mg L–1.

Guzzon et al., 2024 – Sparkling wine produced at 20 °C using, in the secondary bottle fermentation, a mixed culture at a ratio of 1:1 made with H. uvarum NCYC 2380 and S. cerevisiae DV10; sparkling wines after bottle-fermentation from: a) Lambrusco Sorbara grapes contained 10.114 mg L–1; the control (inoculated with S. cerevisiae DV10) contained 10.085 mg L–1; b) Lambrusco Marani grapes contained 8.074 mg L–1; the control (inoculated with S. cerevisiae DV10) contained 8.014 mg L–1.

Medina et al., 2013 – Grape wine produced at 20 °C inoculating Chardonnay must, added with 30 mg L–1 of sulfur dioxide, with 5 × 105 cells mL–1 of H. vineae 02/5AF and, after six days, 2 × 106 cells of S. cerevisiae ALG 804 contained 4.607 mg L–1; the control (inoculated with 2 × 106 cells mL–1 of S. cerevisiae ALG 804) contained 3.796 mg L–1.

Final commentary: decanoic acid – the less, the better.

1.7. Octanoic acid (i.e., caprylic acid)

Octanoic acid is related to an unpleasant aroma that is described as “rancid, harsh, cheese, fatty acid” (Guth, 1997) and “sweet, cheese” (Carrau et al., 2023); its odour threshold is 0.5 mg L–1 (Carrau et al., 2023). At least two strains of Hanseniaspora were reported to produce octanoic acid exceeding the odour threshold, thereby negatively affecting the sensory quality of the wine.

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and 106 cells mL–1 of S. cerevisiae RV002 contained 1.178 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae RV002) contained 0.919 mg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in pilot-scale vinification, inoculating 90 L of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 4.716 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 3.922 mg L–1.

Final commentary: octanoic acid – the less, the better.

1.8. Hexanoic acid (i.e., caproic acid)

At least three different descriptors were reported: “cheese, rancid, fatty” (Guth, 1997), “sweet, acid rancid” (Carrau et al., 2023), and “cheese, fatty, baked potato” (Lai et al., 2023); its odour threshold is 0.42 mg L–1 (Carrau et al., 2023). At least three strains of Hanseniaspora were reported to produce hexanoic acid exceeding the odour threshold, thereby negatively affecting the sensory quality of the wine.

Domizio et al., 2011 – Grape wine produced by inoculating commercial white grape juice with 107 cells mL–1 of H. osmophila Ha32 and 105 cells of S. cerevisiae EC1118 contained 0.72 mg L–1; the control (inoculated with 105 cells mL–1 of S. cerevisiae EC1118) contained 0.70 mg L–1.

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and 106 cells mL–1 of S. cerevisiae RV002 contained 1.709 mg L–1 – the control (inoculated with 106 cells mL–1 of S. cerevisiae RV002) contained 1.323 mg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in pilot-scale vinification, inoculating 90 L of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 2.366 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 2.159 mg L–1.

Final commentary: hexanoic acid – the less, the better.

2. Positive components

2.1. Isoamyl alcohol (i.e., 3-methyl-1-butanol)

At least three groups of descriptors were reported, all with an odour threshold of 30 mg L–1: “banana, cheese, sweet” (Guth, 1997; Sanoppa et al., 2019), “alcoholic, fermented, whiskey” (Bartowsky & Pretorius, 2009; Lu et al., 2015), and “banana, fruity, sweet” (Chen et al., 2015). Some strains of Hanseniaspora were reported to produce isoamyl alcohol exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 202.95 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 167.52 mg L–1.

del Fresno et al., 2021a – Grape wine produced in oak barrels at 18 °C using a mixture of 50 % Tempranillo and 50 % Albillo Mayor grapes; the grape must, containing 15.05 mg L–1 of total sulfur dioxide, was inoculated with 107 cells mL–1 of H. vineae T02/5A and contained 108.64 mg L–1; the control (inoculated with S. cerevisiae Fermivin 3C) contained 125.4 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with: a) 107 cells mL–1 of H. vineae T02/5A contained 133.92 mg L–1, b) 107 cells mL–1 of H. opuntiae A56 contained 119.16 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 137.81 mg L–1.

Domizio et al., 2011 – Grape wine produced inoculating commercial white grape juice with: a) 107 cells mL–1 of H. osmophila Ha32 and 107 cells of S. cerevisiae EC1118 contained 127.95 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae EC1118) contained 113.21 mg L–1 (trial 1); b) 107 cells mL–1 of H. osmophila Ha32 and 105 cells of S. cerevisiae EC1118 contained 114.28 mg L–1; the control (inoculated with 105 cells mL–1 of S. cerevisiae EC1118) contained 108.42 mg L–1 (trial 2).

Luan et al., 2018 – Grape wine produced as previously reported contained 249.74 mg L–1; the control contained 207.50 mg L–1.

Pietrafesa et al., 2020 – Grape wine produced as previously reported contained 240.74 mg L–1; the control contained 222.62 mg L–1.

Testa et al., 2020 – Grape wine produced as previously reported contained 72.1 mg L–1; the 3-methyl-1-butanol was not detected in the control.

Yan et al., 2020 – Grape wine produced as previously reported contained 105.81 mg L–1; the control contained 64.34 mg L–1.

Final commentary: isoamyl alcohol may contribute positively to the complexity of the wine aromas.

2.2. 2-Phenylethanol (i.e., 2-Phenyl ethyl alcohol)

At least four different descriptors were reported: “flowery, pollen, perfumed” (Guth, 1997), “floral, roses” (Tristezza et al., 2016), “honey, rose, spicy” (Carrau et al., 2023), and “floral, roses” (Lai et al., 2023); its odour threshold is 14 mg L–1 (Carrau et al., 2023). Some strains of Hanseniaspora were reported to produce 2-phenylethanol at levels exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 42.72 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 30.63 mg L–1.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. occidentalis GYBC-211 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 59.4 mg L–1; b) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 48.52 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 44.34 mg L–1.

Capozzi et al., 2019 – Grape wine produced at 25 °C, inoculating Negroamaro must, added with 100 mg L–1 potassium metabisulfite, with 106 cells mL–1 of H. uvarum ITEM 8795 and 104 cells of S. cerevisiae ITEM 17292, contained 17.96 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae ITEM 17292) contained 14.91 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with: a) 107 cells mL–1 of H. vineae T02/5A contained 25.02 mg L–1, b) 107 cells mL–1 of H. opuntiae A56 contained 21.45 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 9.66 mg L–1.

Gallo et al., 2024a – Grape wine produced using a mixture of 50 % Glera and 50 % Gewurztraminer grapes, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Fermivin VINEAE and, after 200 hours, with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB contained 14.03 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 13.62 mg L–1.

Guzzon et al., 2024 – Sparkling wine produced at 20 °C using, in the secondary bottle fermentation, a mixed culture at a ratio of 1:1 made with H. uvarum NCYC 2380 and S. cerevisiae DV10; sparkling wines after bottle-fermentation from: a) Lambrusco Sorbara grapes contained 21.047 mg L–1; the control (inoculated with S. cerevisiae DV10) contained 19.891 mg L–1; b) Lambrusco Marani grapes contained 12.186 mg L–1; the control (inoculated with S. cerevisiae DV10) contained 10.199 mg L–1.

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and, after 24 hours, with 106 cells mL–1 of S. cerevisiae RV171 contained 45.48 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae RV171) contained 44.94 mg L–1.

Luan et al., 2018 – Grape wine produced as previously reported contained 61.3 mg L–1; the control contained 51.2 mg L–1.

Testa et al., 2020 – Grape wine produced as previously reported contained 160.1 mg L–1; the control contained 92 mg L–1.

Yan et al., 2020 – Grape wine produced as previously reported contained 26.72 mg L–1; the control contained 16.08 mg L–1.

Zhang et al., 2022 – Grape wine produced at 16–17 °C from Petit Manseng must added with sulfur dioxide (60 mg L–1) and inoculated with H. vineae CVE-HV6 and, after four days, with S. cerevisiae D254 contained 28.05 mg L–1; the control (inoculated with S. cerevisiae D254) contained 19.08 mg L–1.

Final commentary: 2-phenylethanol contributes fine rose notes to the wine aroma (Tristezza et al., 2016).

2.3. Hexyl acetate

The aromatic descriptors of this compound are “apple, cherry, pear, floral”; its odour threshold is 0.67 mg L–1 (Peinado et al., 2004). At least two strains of Hanseniaspora were reported to produce hexyl acetate exceeding the odour threshold, so positively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 23.22 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 6.81 mg L–1.

Lai et al., 2023 – Grape wine produced at 25 °C from Kyoho must filtered and inoculated with 5 × 106 cells mL–1 of H. uvarum Pi235 and 5 × 106 cells mL–1 of S. cerevisiae Gr112 contained 8.34 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Gr112) contained 2.13 mg L–1.

Final commentary: hexyl acetate – the more, the merrier.

2.4. 2-Phenylethyl acetate (i.e., 2-phenethyl acetate)

The aromatic descriptor of this compound is “fruity, honey, floral”; the odour threshold is 0.25 mg L–1 (Carrau et al., 2023). Some strains of Hanseniaspora were reported to produce 2-phenylethyl acetate at levels exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 4.23 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 2.99 mg L–1.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. osmophila NRRL-Y1613 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 1.24 mg L–1; b) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 1.18 mg L–1; c) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.68 mg L–1; d) 107 cells mL–1 of H. meyeri NRRL-Y27513 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.49 mg L–1; e) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.47 mg L–1; f) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL 1 of S. cerevisiae Uvaferm CEG contained 0.27 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 0.203 mg L–1.

del Fresno et al., 2021a – Grape wine produced in stainless steel barrels at 18 °C using a mixture of 50 % Tempranillo and 50 % Albillo Mayor grapes; the grape must, containing 15.05 mg L–1 of total sulfur dioxide, was inoculated with 107 cells mL–1 of H. vineae T02/5A and contained 15.38 mg L–1; the control (inoculated with S. cerevisiae Fermivin 3C) contained 9.28 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with: a) 107 cells mL–1 of H. vineae T02/5A contained 8.91 mg L–1, b) 107 cells mL–1 of H. opuntiae A56 contained 7.78 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 7.44 mg L–1.

Escott et al., 2021 – Grape wine produced from Tempranillo grapes inoculated with 1.3 × 108 cells mL–1 of H. vineae HV and, after eight days of fermentation, with 6 × 107 cells mL–1 of S. cerevisiae 7VA contained 6.6 mg L–1; the control (inoculated with 108 cells mL–1 S. cerevisiae 7VA) contained 5.1 mg L–1.

Filippousi et al., 2024 – Grape wine produced as previously reported contained 78.29 mg L–1; the control contained 29.71 mg L–1.

Gallo et al., 2023 – Grape wine produced as previously reported contained 3.19 mg L–1 – the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae LVCB) contained 0.19 mg L–1.

Gallo et al., 2024a – Grape wine produced using a mixture of 50 % Glera and 50 % Gewurztraminer grapes, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Fermivin VINEAE and, after 24 hours, with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB contained 4.64 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 0.385 mg L–1.

Gallo et al., 2024b – Grape wine produced from Glera grape must, added with sulfur dioxide (15 mg L–1) as potassium metabisulfite, inoculated with 5 × 106 cells mL–1 (98 % of H. vineae Fermivin VINEAE and 2 % of S. cerevisiae Fermivin LVCB) contained 7.6 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 0.12 mg L–1.

Lai et al., 2023 – Grape wine produced as previously reported contained 2.66 mg L–1; the control contained 1.71 mg L–1.

Medina et al., 2013 – Grape wine produced as previously contained 4.341 mg L–1; the control contained 0.425 mg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in pilot-scale vinification, inoculating 90 L of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 0.696 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 0.517 mg L–1.

Valera et al., 2021 – Grape wine produced at 20 °C from Chardonnay grape must, treated with 200 mg L–1 dimethyldicarbonate and inoculated with 105 cells mL–1 of H. vineae 025, contained 10.524 mg L–1; the control (inoculated with 105 cells mL–1 of S. cerevisiae ALG804) contained 0.116 mg L–1.

Yan et al., 2020 – Grape wine produced as previously reported contained 2.75 mg L–1; the control contained 0.29 mg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and, after two days, with 106 cells mL–1 of S. cerevisiae CVE-SC45 contained 8.342 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 2.591 mg L–1.

Zhang et al., 2022 – Grape wine produced as previously reported contained 1.55 mg L–1; the control contained 0.20 mg L 1.

Final commentary: 2-phenylethyl acetate is a compound responsible for imparting flowery, rosy, and honey-like fragrances with fruity undertones to wines (Filippousi et al., 2024).

2.5. Ethyl decanoate (i.e., ethyl caprate)

At least four different descriptors were reported: “fruity, fatty, pleasant” (Guth, 1997), “sweet, fruity” (Tristezza et al., 2016), “floral” (Mestre et al., 2019), and “fruity” (Tarko & Duda, 2024); its odour threshold is 0.2 mg L–1 (Ferreira et al., 2000). Some strains of Hanseniaspora were reported to produce ethyl decanoate exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.666 mg L–1; b) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.576 mg L–1; c) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.525 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 0.468 mg L–1.

Capozzi et al., 2019 – Grape wine produced at 25 °C, inoculating Negroamaro must, added with 100 mg L–1 potassium metabisulfite, with 106 cells mL–1 of H. uvarum ITEM 8795 and 104 cells of S. cerevisiae ITEM 17293 contained 1.88 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae ITEM 17293) contained 0.95 mg L–1.

Gallo et al., 2024a – Grape wine produced using a mixture of 50 % Glera and 50 % Gewurztraminer grapes, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Fermivin VINEAE and, after 48 hours, with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB contained 0.511 mg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 0.16 mg L–1.

Li et al., 2022 – Grape wine produced as previously reported contained 0.570 mg L–1; the control contained 0.457 mg L–1.

Medina et al., 2013 – Grape wine produced as previously contained 0.685 mg L–1; the control contained 0.56 mg L–1.

Mestre et al., 2019 – Grape wine produced at 25 °C from Malbec grape must added with 50 ppm of free sulfur dioxide and inoculated with 5 × 106 cells mL–1 of H. uvarum BHu9 and, after two days, with 2 × 106 cells mL–1 of S. cerevisiae BSc114 contained 1.14 mg L–1; the control (inoculated with 2 × 106 cells mL–1 of S. cerevisiae BSc114) contained 0.59 mg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in pilot-scale vinification, inoculating 90 L of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 0.252 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 0.188 mg L–1.

Final commentary: ethyl decanoate – the more, the merrier.

2.6. Isoamyl acetate (i.e., 3-methylbutyl acetate)

The aromatic descriptor of this compound is “banana, fruity, sweet”, the odour threshold is 0.16 mg L–1 (Peinado et al., 2004). Some strains of Hanseniaspora were reported to produce isoamyl acetate exceeding the odour threshold, so positively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL—1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 0.7 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 0.25 mg L–1.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 3.788 mg L–1; b) 107 cells mL–1 of H. meyeri NRRL-Y27513 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 2.731 mg L–1; c) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 2.523 mg L–1; d) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 2.12 mg L–1 – the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 1.089 mg L–1.

Capozzi et al., 2019 – Grape wine produced at 25 °C inoculating Negroamaro must, added with 100 mg L–1 potassium metabisulfite, with 106 cells mL–1 of H. uvarum ITEM 8795 and 104 cells of S. cerevisiae ITEM 17292 contained 4.11 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae ITEM 17292) contained 3.77 mg L–1; grape wine produced at 25 °C inoculating Negroamaro must, added with 100 mg L–1 potassium metabisulfite, with 106 cells mL–1 of H. uvarum ITEM 8795 and 104 cells of S. cerevisiae ITEM 17293 contained 3.11 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae ITEM 17293) contained 2.11 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with: a) 107 cells mL–1 of H. vineae T02/5A contained 3.33 mg L–1, b) 107 cells mL–1 of H. opuntiae A56 contained 16.10 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 2.70 mg L–1.

Domizio et al., 2011 – Grape wine produced by inoculating commercial white grape juice with 107 cells mL–1 of H. osmophila Ha32 and 105 cells of S. cerevisiae EC1118 contained 0.28 mg L–1; the control (inoculated with 105 cells mL–1 of S. cerevisiae EC1118) contained 0.21 mg L–1.

Escott et al., 2021 – Grape wine produced as previously reported contained 2.3 mg L–1; the control contained 2 mg L–1.

Gallo et al., 2024b – Grape wine produced as previously reported contained 1.02 mg L–1; the control contained 0.72 mg L–1.

Lai et al., 2023 – Grape wine produced as previously reported contained 0.73 mg L–1; the control contained 0.27 mg L–1.

Li et al., 2022 – Grape wine produced as previously reported contained 7.922 mg L–1; the control contained 4.695 mg L–1.

Li et al., 2024 – Grape wine produced at 14–16 °C using Sauvignon blanc grape must, added with 1 mg L–1 of H2SO3 (calculated as 60 mg L–1 of sulfur dioxide), inoculated with 107 cells mL–1 of H. uvarum HU4487 and 106 cells mL–1 of S. cerevisiae VL3 contained 1.503 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae VL3) contained 1.361 mg L–1.

Medina et al., 2013 – Grape wine produced as previously contained 0.948 mg L–1; the control contained 0.871 mg L–1.

Mestre et al., 2019 – Grape wine produced as previously reported contained 3.21 mg L–1; the control contained 2.89 mg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in industrial vinification, inoculating 7 tons of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 2.597 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 0.312 mg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and 106 cells mL 1 of S. cerevisiae CVE-SC45 contained 1.129 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 0.964 mg L–1.

Zhang et al., 2022 – Grape wine produced as previously reported contained 2.044 mg L–1; the control contained 1.709 mg L–1.

Final commentary: isoamyl acetate is one of the esters most markedly contributing to the aroma profile of white wines (Plata et al., 2003).

2.7. Ethyl butyrate (i.e., ethyl butanoate)

For ethyl butyrate, at least two different descriptors, both with the odour threshold of 0.02 mg L–1, were reported: “fruity” (Guth, 1997; Tristezza et al., 2016) and “strawberry, apple” (Peng et al., 2013; Li et al., 2020). Some strains of Hanseniaspora were reported to produce ethyl butyrate exceeding the odour threshold, so positively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.312 mg L–1; b) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.272 mg L–1; c) 107 cells mL–1 of H. meyeri NRRL-Y27513 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.248 mg L–1; d) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.241 mg L–1; e) 107 cells mL–1 of H. osmophila NRRL-Y1613 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.227 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 0.222 mg L–1.

del Fresno et al., 2025 – Grape wine produced at 18 °C from Albillo Mayor grape must, added with sulfur dioxide at a dose of 4 g hL–1 in the form of potassium metabisulfite, and inoculated with 107 cells mL–1 of H. vineae T02/5A contained 4.03 mg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae CTPL14) contained 1.73 mg L–1.

Filippousi et al., 2024 – Grape wine produced as previously reported contained 22.84 mg L–1; the control contained 19.15 mg L–1.

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and 106 cells mL–1 of S. cerevisiae RV002 contained 0.233 mg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae RV002) contained 0.204 mg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in pilot-scale vinification inoculating 90 L of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920 contained 0.425 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 0.425 mg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 0.386 mg L–1.

Zhang et al., 2022 – Grape wine produced as previously reported contained 0.125 mg L–1; the control contained 0.119 mg L–1.

Final commentary: ethyl butyrate – the more, the merrier.

2.8. β-Citronellol

del Fresno et al. (2021b) highlight the high impact of H. vineae on the formation of fruity and floral terpenes as β-citronellol. At least two different descriptors were reported: “green lemon” (Guth, 1997) and “citric” (del Fresno et al., 2021b); its odour threshold is 18 µg L–1 (Čuš & Jenko, 2013). At least three strains of Hanseniaspora were reported to produce β-citronellol exceeding the odour threshold, so positively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 38 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 27 µg L–1.

del Fresno et al., 2021b – Grape wine produced at 18 °C from Albillo Mayor grape must, inoculated with 107 cells mL–1 of H. vineae followed sequentially by 107 cells mL–1 of S. cerevisiae Fermivin 3C, contained 103 µg L–1; the control (inoculated with 107 cells mL–1 of S. cerevisiae Fermivin 3C) contained 24 µg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and, after two days, with 106 cells mL–1 of S. cerevisiae CVE-SC45 contained 25.6 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 20.2 µg L–1.

Final commentary: β-citronellol – the more, the merrier.

2.9. Linalool

At least four different descriptors were reported: “citrus, floral, sweet, grape-like” (Peinado et al., 2004), “floral, lemon” (del Fresno et al., 2021b), “Muscat, flowery, fruit” (Testa et al., 2021), and “floral, citrus” (Lai et al., 2023); its odour threshold is 15 µg L–1 (Peinado et al., 2004). Some strains of Hanseniaspora were reported to produce linalool exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 73.91 µg L–1; b) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 72.93 µg L–1; c) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 72.5 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 72.43 µg L–1.

del Fresno et al., 2021b – Grape wine produced as previously reported contained 70 µg L–1; the control contained 18 µg L–1.

Gallo et al., 2024a – Grape wine produced using a mixture of 50 % Glera and 50 % Gewurztraminer grapes, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Fermivin VINEAE and, after 100 hours, with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB contained 104 µg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 99 µg L–1.

Testa et al., 2020 – Grape wine produced as previously reported contained 112 µg L–1; the control contained 82 µg L–1.

Testa et al., 2021 – Grape wine produced at 23-24 °C from Aglianico grape must added with potassium metabisulfite (70 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum AS27 contained 69.3 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae FE) contained 23.2 µg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and, after four days, with 106 cells mL–1 of S. cerevisiae CVE-SC45 contained 47.2 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 42 µg L–1.

Final commentary: linalool – the more, the merrier.

2.10. Ethyl hexanoate (i.e., ethyl caproate)

At least three very similar descriptors were reported: “fruity, apple” (Mestre et al., 2019), “pineapple, fruity” (Lai et al., 2023), and “fruity/anise” (Tarko & Duda, 2024); its odour threshold is 14 µg L–1 (Ferreira et al., 2000). Some strains of Hanseniaspora were reported to produce ethyl hexanoate exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Andorrà et al., 2010 – Grape wine produced at 20 °C from Macabeo must sterilised by the addition of 250 mg L–1 of dimethyldicarbonate and inoculated with 9 × 105 cells mL–1 of H. uvarum HuB10 and 105 cells mL–1 of S. cerevisiae QA23 contained 220 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae QA23) contained 30 µg L–1.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 92 µg L–1; b) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 77 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 76 µg L–1.

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and 106 cells mL–1 of S. cerevisiae RV002 contained 396 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae RV002) contained 296 mg L–1.

Li et al., 2022 – Grape wine produced as previously reported contained 968 µg L–1; the control contained 942 µg L–1.

Mestre et al., 2019 – Grape wine produced as previously reported contained 600 µg L–1; the control contained 200 µg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in pilot-scale vinification, inoculating 90 L of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 561 µg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 510 µg L–1.

Final commentary: ethyl hexanoate is responsible for tropical fruity attributes.

2.11. Ethyl octanoate (i.e., ethyl caprylate)

At least two very similar descriptors were reported: “pineapple, pear” (Mestre et al., 2019) and “fruity/fresh” (Tarko & Duda, 2024); its odour threshold is 5 µg L–1 (Ferreira et al., 2000). Some strains of Hanseniaspora were reported to produce ethyl octanoate exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 541 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 494 µg L–1.

Lai et al., 2023 – Grape wine produced as previously reported contained 1860 µg L–1; the control contained 1580 µg L–1.

Li et al., 2020 – Grape wine produced at 22 °C from Granoir grape must, added with sulfur dioxide (60 mg L–1) and inoculated with 106 cells mL–1 of H. uvarum Yun268 and 106 cells mL–1 of S. cerevisiae RV002 contained 380 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae RV002) contained 261 µg L–1.

Li et al., 2024 – Grape wine produced as previously reported contained 59.2 µg L–1; the control contained 57.8 µg L–1.

Mestre et al., 2019 – Grape wine produced as previously reported contained 872 µg L–1; the control contained 369 µg L–1.

Tristezza et al., 2016 – Grape wine produced at 20 °C in industrial vinification, inoculating 7 tons of Negroamaro must, added with 20 g hL–1 of potassium metabisulfite, with 107 cells mL–1 of H. uvarum ITEM8795 and 105 cells mL–1 of S. cerevisiae ITEM6920, contained 661 µg L–1; the control (inoculated with S. cerevisiae ITEM6920) contained 476 µg L–1.

Final commentary: ethyl octanoate – the more, the merrier.

2.12. β-Damascenone

For β-damascenone, at least two different descriptors have been reported: “honey” (Tarko & Duda, 2024) and “sweet, exotic flowers, stewed” (Pineau et al., 2007); its odour threshold is 0.05 µg L–1 (Guth, 1997; Langen et al., 2016). Some strains of Hanseniaspora have been reported to produce β-damascenone exceeding the odour threshold, thereby positively contributing to the sensory quality of the wine.

Badura et al., 2023 – Grape wine produced at 22 °C from pasteurised Gewürztraminer grape must inoculated with: a) 107 cells mL–1 of H. osmophila NRRL-Y1613 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.7 µg L–1; b) 107 cells mL–1 of H. opuntiae GYBC-284 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.69 µg L–1; c) 107 cells mL–1 of H. occidentalis GYBC-211 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.64 µg L–1; d) 107 cells mL–1 of H. nectarophila GYBC-283 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.61 µg L–1; e) 107 cells mL–1 of H. guilliermondii NRRL-Y1625 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.56 µg L–1; f) 107 cells mL–1 of H. uvarum DSM2768 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.55 µg L–1; g) 107 cells mL–1 of H. meyeri NRRL-Y27513 and 106 cells mL–1 of S. cerevisiae Uvaferm CEG contained 0.53 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae Uvaferm CEG) contained 0.46 µg L–1.

Gallo et al., 2024a – Grape wine produced using a mixture of 50 % Glera and 50 % Gewurztraminer grapes, added with sulfur dioxide (15 mg L–1) and inoculated with 5 × 106 cells mL–1 of H. vineae Fermivin VINEAE and, after 68 hours, with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB contained 2.13 µg L–1; the control (inoculated with 5 × 106 cells mL–1 of S. cerevisiae Fermivin LVCB) contained 1.43 µg L–1.

Yan et al., 2020 – Grape wine produced as previously reported contained 8.38 µg L–1; the control contained 8.11 µg L–1.

Zhang et al., 2018 – Icewine produced at 16 °C from Vidal blanc grape juice, added with sulfur dioxide (80 mg L–1) and inoculated with 107 cells mL–1 of H. vineae CVE-HV11 and, after two days, with 106 cells mL–1 of S. cerevisiae CVE-SC45 contained 101.7 µg L–1; the control (inoculated with 106 cells mL–1 of S. cerevisiae CVE-SC45) contained 48.1 µg L–1.

Zhang et al., 2022 – Grape wine produced as previously reported contained 11.51 µg L–1; the control contained 9.87 µg L–1.

Final commentary: β-damascenone – the more, the merrier.

Concluding remarks

Among the many recent reviews (Padilla et al., 2016; Varela, 2016; Martin et al., 2018; Morata et al., 2020; Tufariello et al., 2021; van Wyk et al., 2024; Wang et al., 2024) exploring the contribution of yeast to wine aroma, this minireview highlights the relationship between the application Hanseniaspora yeasts in winemaking and the quality of the wine by considering the odour thresholds of the produced compounds (Figure 1).

Figure 1. Graphical abstract image.

Table 1. Negative wine components reported in decreasing order of odour threshold.

Component

Aromatic descriptors

Odour threshold

Reference

Acetic acid

Vinegar

700 mg L–1

Carrau et al., 2023

Ethyl acetate

Spoilage character

150–200 mg L–1

Viana et al., 2008

Acetoin

Butter, creamy, fat

150 mg L–1

Carrau et al., 2023

Isobutanol

Bitter, chemical alcoholic, fusel oil, malt notes, nail polish, solvent, wineosity notes

40 mg L–1

Guth, 1997; Testa et al., 2021; Carrau et al., 2023; Tarko & Duda, 2024

1-Hexanol

Grass, green, herbaceous, woody

1.1 mg L–1

Peinado et al., 2004

Decanoic acid

Fatty, rancid, unpleasant

1 mg L–1

Carrau et al., 2023

Octanoic acid

Cheese, fatty acid, harsh, rancid, sweet

0.5 mg L–1

Carrau et al., 2023

Hexanoic acid

Acid rancid, baked potato, cheese, fatty, rancid, sweet

0.42 mg L–1

Carrau et al., 2023

Table 2. Positive wine components reported in decreasing order of odour threshold.

Component

Aromatic descriptors

Odour threshold

Reference

Isoamyl alcohol

Alcoholic, banana, cheese, fermented, fruity, sweet, whiskey

30 mg L–1

Guth, 1997; Bartowsky & Pretorius, 2009; Chen et al., 2015

2-Phenylethanol

Floral, flowery, honey, perfumed, pollen, roses, spicy

14 mg L–1

Carrau et al., 2023

Hexyl acetate

Apple, cherry, floral, pear

0.67 mg L–1

Peinado et al., 2004

2-Phenylethyl acetate

Floral, fruity, honey

0.25 mg L–1

Carrau et al., 2023

Ethyl decanoate

Fatty, floral, fruity, pleasant, sweet

0.2 mg L–1

Ferreira et al., 2000

Isoamyl acetate

Banana, fruity, sweet

0.16 mg L–1

Peinado et al., 2004

Ethyl butyrate

Apple, fruity, strawberry

0.02 mg L–1

Guth, 1997; Peng et al., 2013

β-Citronellol

Citric, green lemon

18 µg L–1

Čuš & Jenko, 2013

Linalool

Citrus, floral, flowery, fruit, grape-like, lemon, Muscat, sweet

15 µg L–1

Peinado et al., 2004

Ethyl hexanoate

Anise, apple, fruity, pineapple

14 µg L–1

Ferreira et al., 2000

Ethyl octanoate

Fruity/fresh, pear, pineapple

5 µg L–1

Ferreira et al., 2000

β-Damascenone

Exotic flowers, honey, stewed, sweet

0.05 µg L–1

Guth, 1997

Acknowledgements

The authors have nothing to report.

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Authors

Andrea Caridi

acaridi@unirc.it

Affiliation : Mediterranea University of Reggio Calabria, Department of Agriculture, Via Feo di Vito, 89122 Reggio Calabria, Italy

Country : Italy

Marco Poiana

Affiliation : Mediterranea University of Reggio Calabria, Department of Agriculture, Via Feo di Vito, 89122 Reggio Calabria, Italy

Country : Italy

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