非酿酒酵母属酵母的葡萄酒发酵应用研究进展

doi:10.3864/j.issn.0578-1752.2023.03.011

中国农业科学 ›› 2023, Vol. 56 ›› Issue (3): 529-548.doi: 10.3864/j.issn.0578-1752.2023.03.011

非酿酒酵母属酵母的葡萄酒发酵应用研究进展

王春晓¹(), 俞俊竹¹, 周文亚¹, 许引虎²

¹贵州大学酿酒与食品工程学院/贵州省发酵工程与生物制药重点实验室，贵阳 550025
²国家酵母技术研究推广中心，湖北宜昌 443003

收稿日期:2022-04-30 接受日期:2022-07-22 出版日期:2023-02-01 发布日期:2023-02-14
通信作者: 王春晓，Tel：18111908483；E-mail：cxwang@gzu.edu.cn
联系方式: 王春晓，Tel：18111908483；E-mail：cxwang@gzu.edu.cn
基金资助:
国家自然科学基金青年科学基金(32160544);国家自然科学基金青年科学基金(31801523);贵州省科技计划(黔科合支撑[2022]重点006号);贵州省教育厅青年科技人才成长项目(黔教合KY字[2018]120);贵州大学引进人才科研项目(贵大人基合字[2017]44号);贵州大学引进人才科研项目(贵大培育[2020]50号);贵州省发酵工程与白酒酿造人才基地(黔人领发[2018]3号);遵义市创新人才团队培养项目（遵义科人才[2020]9(遵义科人才[2020]9)

Research Progress on the Application of Non-Saccharomyces During Wine Fermentation

WANG ChunXiao¹(), YU JunZhu¹, ZHOU WenYa¹, XU YinHu²

¹School of Liquor and Food Engineering, Guizhou University/Key Laboratory of Fermentation Engineering and Biological Pharmacy of Guizhou Province, Guiyang 550025
²National Center for Yeast Technology Research and Promotion, Yichang 443003, Hubei

Received:2022-04-30 Accepted:2022-07-22 Published:2023-02-01 Online:2023-02-14

摘要/Abstract

摘要：

葡萄皮上天然存在着非酿酒酵母属酵母（non-Saccharomyces），主要在葡萄酒浸渍和发酵初期发挥作用，近年来非酿酒酵母属酵母在葡萄酒发酵中的应用受到越来越多的关注。相对于酿酒酵母，非酿酒酵母属酵母在酒精发酵中具有较弱的发酵力，可将还原糖转化为乙醇及其他代谢副产物，是生产复杂风味和低酒度葡萄酒的潜在优良酵母。不同非酿酒酵母属酵母菌种在葡萄酒发酵应用中具有不同的代谢特征，选择具有一定特征的优良非酿酒酵母属酵母应用于发酵中，可以提高葡萄酒的特色化品质。本研究在总结商业化非酿酒酵母属酵母的种类、酿造特点和应用方式的基础上，重点综述了不同非酿酒酵母属酵母对葡萄酒颜色、香气、口感和安全健康4个方面的积极作用、代谢机理和研究热点：具有高产酸、多糖、胞外丙酮酸及低吸附性等特性的非酿酒酵母属酵母可通过不同代谢机理促进葡萄酒颜色的稳定；不同非酿酒酵母属酵母通过低产乙醇、乙醛、降低挥发性酚类，高产乙酸乙酯、乙酸酯类化合物、乙酯类化合物、高级醇、与萜烯或硫醇释放相关的酶类等途径促进葡萄酒果味香气的提升，增加香气复杂性；非酿酒酵母属酵母通过高产甘油、多糖和乳酸，降解苹果酸等方式调节葡萄酒的口感特征；非酿酒酵母属酵母作为生物防治剂可以降低葡萄酒酿造中二氧化硫的用量，通过代谢降解作用减少有毒化合物，提升葡萄酒的安全质量。本文进一步解析了非酿酒酵母属酵母的基因组和微卫星位点分析研究现状，探讨了目前非酿酒酵母属酵母葡萄酒发酵应用研究的主要接种策略，提出了未来研究仍需关注的热点方向，为非酿酒酵母属酵母在葡萄酒酒精发酵中的应用研究提供理论参考。

关键词: 混合发酵, 颜色稳定性, 香气复杂性, 基因型, 商业化

Abstract:

Non-Saccharomyces yeast are naturally present on grape skin, which mainly play their roles in the maceration and initial stage of fermentation during winemaking process. More and more concerns have been recently focused on the application of non-Saccharomyces yeast in wine fermentation. Non-Saccharomyces yeast often have weaker fermentation ability than Saccharomyces cerevisiae and can convert the reducing sugar into ethanol and other fermentation by-products. Therefore, non-Saccharomyces are the potential excellent yeast for producing wine with complex flavor traits and lower ethanol degree. Different non-Saccharomyces yeast species have different metabolic characteristics in wine fermentation applications. The selection of non-Saccharomyces yeasts with certain characteristics for application in fermentation can improve the specialized quality of wine. This review was based on the initial summary on the species, winemaking traits, and application ways of commercial non-Saccharomyces yeast, and emphasized on the positive role of non-Saccharomyces in wine color, aroma, taste, safety and health, their metabolic mechanisms and corresponding research hotspot. The positive role of non-Saccharomyces in wine color included the promotion of high acid production, high polysaccharide production, high extracellular pyruvate production and low sorption properties. Different non-Saccharomyces yeast could promote the fruity aroma enhancement in wine by low production of ethanol, acetaldehyde, and volatile phenols, and high production of ethyl acetate, acetic acid ester, ethyl ester, higher alcohols, and enzymes related with the release of terpene and mercaptan, and therefore improve the aroma complexity of wine. Non-Saccharomyces yeast could regulate wine mouthfeel traits by high production of glycerol, polysaccharide, and lactic acid, and degradation of malic acid. In addition, non-Saccharomyces yeast could improve the safety quality of wine by decreasing the SO₂ dosage as biological control agent, and reducing the content of toxic compounds through metabolic degradation during alcoholic fermentation. This review further analyzed the current research status on genome and microsatellite loci analysis. The main inoculation strategies were discussed on application of non-Saccharomyces yeast during wine fermentation, and six aspects were posed for further research concern. This review provided theoretical references for the research on application of the non-Saccharomyces yeasts in wine alcoholic fermentation.

Key words: mixed fermentation, color stability, aroma complexity, genotype, commercialization

王春晓, 俞俊竹, 周文亚, 许引虎. 非酿酒酵母属酵母的葡萄酒发酵应用研究进展[J]. 中国农业科学, 2023, 56(3): 529-548.

WANG ChunXiao, YU JunZhu, ZHOU WenYa, XU YinHu. Research Progress on the Application of Non-Saccharomyces During Wine Fermentation[J]. Scientia Agricultura Sinica, 2023, 56(3): 529-548.

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图/表 6

表1

已经商业化应用于葡萄酒酿造领域的非酿酒酵母属酵母"

菌种名称 Species	葡萄酒酿造应用特点 Winemaking application characteristics	产品名称^a Product	生产商 Producer	菌剂加工类型 Yeast processing type
威克克鲁维酵母 Kluyveromyces wickerhamii	产真菌毒素抑制败坏菌生长 Production of a mycotoxin inhibiting the growth of spoilage microorganisms	ENARTIS FERM BRETT OUT K^[13]	意大利Enartis Italy Enartis	活性干酵母 Active dry yeast
耐热克鲁维酵母 Lachancea thermotolerans	产乳酸^[14-15] Production of lactic acid	CVE-7	中国安琪Angel China Angel	活性干酵母 Active dry yeast
	产乳酸增加酸度，低产乙醇 Production of lactic acid to increase acidity and low production of ethanol	LEVULIA Alcomeno	意大利AEB Italy AEB	活性干酵母 Active dry yeast
	高产乳酸增酸 High production of lactic acid to increase acidity	LAKTIA	葡萄牙Proenol Portugal Proenol	活性干酵母 Active dry yeast
	增加酸度 Increase acidity	Concerto^[16]	丹麦CHR Hansen Denmark CHR Hansen	活性干酵母 Active dry yeast
	增加酸度和清新感 Increase acidity and freshness	LEVEL2 LAKTIA	加拿大Lallemand Canada Lallemand	活性干酵母 Active dry yeast
核果梅奇酵母 Metschnikowia fructicola	葡萄采收期或冷浸渍期间抑制有害微生物生长 Inhibit the growth of harmful microorganisms during grape harvest or cold maceration	GAÏA	加拿大Lallemand Canada Lallemand	活性干酵母 Active dry yeast
美极梅奇酵母 Metschnikowia pulcherrima	增加品种香气，与酿酒酵母顺序接种使用 Increase the aroma of grapevarieties and inoculate with Saccharomyces cerevisiae in sequence	LEVULIA PULCHERRIMA	意大利AEB Italy AEB	活性干酵母 Active dry yeast
	葡萄采摘后立即使用，用于生物防治 Use immediately after grape picking for biocontrol purposes	PRIMAFLORA VB BIO	意大利AEB Italy AEB	活性干酵母 Active dry yeast
	控制葡萄表面的野生菌，减少二氧化硫使用 Control wild bacteria on the surface of grapes and reduce the use of sulfur dioxide	OenofermMProtect	德国Erbslöeh Germany Erbslöeh	活性干酵母 Active dry yeast
	生物防治，减少二氧化硫使用 Biological control and reduction of sulfur dioxide use	LEVEL² GUARDIA	葡萄牙Proenol Portugal Proenol	活性干酵母 Active dry yeast
	增加品种香气 Increase the aroma of grape varieties	FLAVIA	葡萄牙Proenol Portugal Proenol	活性干酵母 Active dry yeast
	高消耗溶解氧防止氧化，二氧化硫替代品 High consumption of dissolved oxygen to prevent oxidation, sulfur dioxide substitute	LEVEL² INITIA	葡萄牙Proenol Portugal Proenol	活性干酵母 Active dry yeast
	低温浸渍阶段生物防治 Biocontrol at low temperature maceration stages	ZYMAFLORE KHIO^MP	法国Laffort France Laffort	活性干酵母 Active dry yeast
	增加品种特性 Increase grape variety characteristics	LEVEL2 FLAVIA	加拿大Lallemand Canada Lallemand	活性干酵母 Active dry yeast
美极梅奇酵母+戴尔有孢圆酵母 Metschnikowia pulcherrima + Torulaspora delbrueckii	生物防治，减少二氧化硫使用 Biological control and reduction of sulfur dioxide use	ZYMAFLORE ÉGIDE^TDMP	法国Laffort France Laffort	活性干酵母 Active dry yeast
克鲁维毕赤酵母 Pichia kluyveri	增加果味 Increase fruitiness	Footzen^[16]	丹麦CHR Hansen Denmark CHR Hansen	活性干酵母 Active dry yeast
克鲁维毕赤酵母+瑟氏哈萨克斯坦酵母 P. kluyveri+Kazachstania servazzii	产生大量香气，生物防治用作二氧化硫替代品 Production of high aroma, and biocontrol is used as a sulphur dioxide substitute	Thrillyeast	意大利BioEnologia Italy BioEnologia	黏稠奶油状的液体酵母 Thick and creamy liquid yeast
酵母属+美极梅齐酵母 Saccharomyces + Metschnikowia pulcherrima	葡萄采摘后立即使用，用于生物防治 Used immediately after grape picking for biocontrol purposes	PRIMAFLORA VR BIO	意大利AEB Italy AEB	活性干酵母 Active dry yeast
酵母属+戴尔有孢圆酵母 Saccharomyces + Torulaspora delbrueckii	增加香气复杂性 Increase aroma complexity	Oenoferm wild & pure F3	德国Erbslöeh Germany Erbslöeh	活性干酵母 Active dry yeast
酿酒酵母+戴尔有孢圆酵母+耐热克鲁维酵母 S. cerevisiae+T. delbrueckii + L. thermotolerans	增加果香 Increase fruity aroma	Melody^[16]	丹麦CHR Hansen Denmark CHR Hansen	活性干酵母 Active dry yeast
粟酒裂殖酵母 Schizosaccharomyces pombe	降解苹果酸、葡萄糖酸和赭曲霉毒素，高产甘油、多糖和乙醇，低产乙酸，不产组胺，提升颜色 Degradation of malic acid, gluconic acid and ochratoxin, high production of glycerol, polysaccharides and ethanol, low production of acetic acid, no histamine, color enhancement	Atecrem 12H	意大利BioEnologia Italy BioEnologia	黏稠奶油状的液体酵母 Thick and creamy liquid yeast
粟酒裂殖酵母 Schizosaccharomyces pombe	降低苹果酸 Lower malic acid	PROMALIC	葡萄牙Proenol Portugal Proenol	胶囊封装酵母 Capsule encapsulated yeast
星形假丝酵母 Starmerella bacillaris	高产甘油，低产乙醇，需无二氧化硫酿造 High production of glycerine, low production of ethanol, no sulphur dioxide required for brewing	Atecrem 11H	意大利BioEnologia Italy BioEnologia	黏稠奶油状的液体酵母 Thick and creamy liquid yeast
戴尔有孢圆酵母 Torulaspora delbrueckii	增加香气复杂性，释放多糖增加口感圆润度 Increases aromatic complexity and releases polysaccharides to increase roundness on the palate	NSD	中国安琪Angel China Angel	活性干酵母 Active dry yeast
	增加果香和花香，低产挥发酸，增加复杂性 Increase fruit and floral aromas, low yield of volatile acids, increases complexity	EnartisFerm Q Tau	意大利Enartis Italy Enartis	活性干酵母 Active dry yeast
	增加香气复杂性 Increase aroma complexity	BIODIVA	葡萄牙Proenol Portugal Proenol	活性干酵母 Active dry yeast
	增加花香和品种香气，释放大量甘露糖蛋白增加口感圆润度 Increase floral and varietal aromas and release a large amount of mannose protein to increase roundness on the palate	VINIFERM NSTD	西班牙Agrovin Spain Agrovin	活性干酵母 Active dry yeast
	与酿酒酵母结合使用，增加口感长度和香气强度，低产挥发酸 Combination with Saccharomyces cerevisiae for increasing length and aroma intensity, and low volatile acid production	ZYMAFLORE ALPHA	法国Laffort France Laffort	活性干酵母 Active dry yeast
	增加品种特性和发酵酯类特性 Increase in grape variety characteristics and fermentation ester characteristics	LEVEL2 BIODIVA	加拿大Lallemand Canada Lallemand	活性干酵母 Active dry yeast
	增加香气 Increase aroma	Prelude^[16]	丹麦CHR Hansen Denmark CHR Hansen	活性干酵母 Active dry yeast
异常威克汉姆酵母 Wickerhamomyces anomalus	产真菌毒素抑制败坏菌生长 Production of a mycotoxin inhibiting the growth of spoilage microorganisms	ENARTIS FERM BRETT OUT W^[13]	意大利Enartis Italy Enartis	活性干酵母 Active dry yeast

表1

图1

图2

图3

与酵母菌发酵途径关联的香气物质形成机理[9,17,19,28,38⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓⇓-54] ①高产乙醛：日本裂殖酵母、粟酒裂殖酵母、星形假丝酵母；低产乙醛、乙酸或3-羟基丁烷-2-酮：耐热克鲁维酵母、L. fermentati、美极梅奇酵母、特立科拉毕赤酵母、库德毕赤酵母、星形假丝酵母、戴尔有孢圆酵母；②降低乙醇：耐热克鲁维酵母、美极梅奇酵母、星形假丝酵母、戴尔有孢圆酵母；③高产高级醇类化合物（Ehrlich途径、糖代谢合成途径）：葡萄酒有孢汉逊酵母、季也蒙毕赤酵母、美极梅奇酵母、发酵毕赤酵母、胶红酵母、日本裂殖酵母；低产高级醇类化合物：有孢汉逊酵母属、星形假丝酵母、粟酒裂殖酵母、接合酵母属；④高产乙酸酯类化合物：加利福尼亚假丝酵母、土星形塞伯林德纳氏酵母、季也蒙有孢汉逊酵母、耐渗透压有孢汉逊酵母、葡萄汁有孢汉逊酵母、有孢汉逊酵母属、好氧哈萨克斯坦酵母、瑟氏哈萨克斯坦酵母、美极梅奇酵母、克鲁维毕赤酵母、特立科拉毕赤酵母、库德毕赤酵母；葡萄酒有孢汉逊酵母通过分枝酸-预苯酸途径高产乙酸酯类化合物；⑤高产乙酸乙酯：季也蒙有孢汉逊酵母、葡萄酒有孢汉逊酵母、葡萄汁有孢汉逊酵母、特立科拉毕赤酵母、库德毕赤酵母、克鲁维毕赤酵母、日本裂殖酵母、拜耳接合酵母；⑥高产乙酯类化合物：有孢汉逊酵母属、葡萄汁有孢汉逊酵母、克鲁维毕赤酵母、库德毕赤酵母、特立科拉毕赤酵母、克拉通覆膜孢酵母、星形假丝酵母、戴尔有孢圆酵母、拜耳接合酵母；⑦高产酶类，释放萜烯类化合物或品种硫醇等：加利福尼亚假丝酵母、铁红假丝酵母、有孢汉逊酵母属、葡萄汁有孢汉逊酵母、耐热克鲁维酵母、美极梅奇酵母、克鲁维毕赤酵母、库德毕赤酵母、胶红酵母、星形假丝酵母、克拉通覆膜孢酵母、戴尔有孢圆酵母"

图3

图4

表2

非酿酒酵母属酵母的基因组相关信息"

菌种名称¹⁾ Species	基因组大小²⁾ Genome size			微卫星位点分析 Microsatellite locus analysis
加利福尼亚假丝酵母 C. californica	细胞核：12.32 Mb Nucleus: 12.32 Mb	2	未知 Unknown	无 None
光滑假丝酵母 C. glabrata	细胞核：12.14—14.56 Mb，13条染色体；线粒体：20 kb Nucleus: 12.14-14.56 Mb, 13 Chromosomes; Mitochondria: 20 kb	36	未知 Unknown	无 None
铁杉假丝酵母 C. railenensis	无 None	无 None	未知 Unknown	无 None
土星形塞伯林德纳氏酵母 C. saturnus	细胞核：13.43 Mb Nucleus: 13.43 Mb	1	未知 Unknown	无 None
汉氏德巴利氏酵母 D. hansenii	细胞核：11.46—18.56 Mb，7条染色体；线粒体：30 kb Nucleus: 11.46-18.56 Mb, 7 Chromosomes; Mitochondria: 30 kb	12	单倍体，1个子囊1个孢子，偶尔2个 Haploid, 1 ascus 1 spore, occasionally 2	无 None
季也蒙有孢汉逊酵母 H. guilliermondii	细胞核：9.04—9.14 Mb，8—9条染色体 Nucleus: 9.04-9.14 Mb, 8-9 Chromosomes	2	倍性未知，1个子囊4个孢子 Ploidy unknown, 1 ascus 4 spores	无 None
耐渗透压有孢汉逊酵母 H. osmophila	细胞核：11.46—12.16 Mb Nucleus: 11.46-12.16 Mb	2	倍性未知，1个子囊1—2个孢子 Ploidy unknown, 1 ascus 1-2 spores	无 None
葡萄汁有孢汉逊酵母 H. uvarum	细胞核：8.1—9.5 Mb，8—9条染色体；线粒体：11 kb Nucleus: 8.1-9.5 Mb, 8-9 Chromosomes; Mitochondria: 11 kb	10	倍性未知可能是二倍体，1个子囊1个孢子，罕见2个孢子 Unknown ploidy is likely diploid, 1 ascus 1 spore, rare 2 spores	10个微卫星位点^[83] Ten microsatellite loci^[83]
葡萄酒有孢汉逊酵母 H. vineae	细胞核：11.1—11.37 Mb，5条染色体；线粒体：28—45 kb Nucleus: 11.1-11.37 Mb, 5 Chromosomes; Mitochondria: 28-45 kb	3	倍性未知，1个子囊1个孢子，罕见2 个孢子 Ploidy unknown, 1 ascus 1 spore, rare 2 spores	无 None
好氧哈萨克斯坦酵母 K. aerobia	细胞核：12.09 Mb Nucleus: 12.09 Mb	1	倍性未知，1个子囊1个孢子 Ploidy unknown, 1 ascus 1 spore	无 None
瑟氏哈萨克斯坦酵母 K. servazzii	细胞核：11.77—12.84 Mb Nucleus: 11.77-12.84 Mb	4	倍性未知，1个子囊1个孢子，偶尔 2—4个孢子 Ploidy unknown, 1 ascus 1 spore, occasionally 2-4 spores	无 None
L. fermentati	细胞核：10.26 Mb，8条染色体 Nucleus: 10.26 Mb, 8 Chromosomes	2	倍性未知，1个子囊1—4个孢子 Ploidy unknown, 1 ascus 1-4 spores	无 None
耐热克鲁维酵母 L. thermotolerans	细胞核：10.24—10.39 Mb，8条染色体；线粒体：21.9—25.1 kb Nucleus: 10.24-10.39 Mb, 8 Chromosomes; Mitochondria: 21.9-25.1 kb	3	倍性有争议，单倍体或二倍体，1个子囊1—4个孢子 Ploidy is controversial, haploid or diploid, 1 ascus 1-4 spores	14个微卫星位点^[84] Fourteen microsatellite loci^[84]
核果梅奇酵母 M. fructicola	细胞核：26.13—26.18 Mb Nucleus: 26.13-26.18 Mb	2	倍性未知，1个子囊2个孢子 Ploidy unknown, 1 ascus 2 spores	无 None
美极梅齐酵母 M. pulcherrima	细胞核：21.77 Mb Nucleus: 21.77 Mb	1	二倍体，1个子囊1—2个孢子 Diploid, 1 ascus 1-2 spores	无 None
季也蒙毕赤酵母 M. guilliermondii	细胞核：10.59—10.71 Mb；线粒体：31 kb Nucleus: 10.59-10.71 Mb; Mitochondria: 31 kb	10	倍性未知，1个子囊1—4个孢子 Ploidy unknown, 1 ascus 1-4 spores	3个微卫星位点^[85] Three microsatellite loci^[85]
发酵毕赤酵母 P. fermentans	细胞核：10.57 Mb，可能2条染色体 Nucleus: 10.57 Mb, possibly 2 chromosomes	1	倍性未知，1个子囊2—4个孢子 Ploidy unknown, 1 ascus 2-4 spores	无 None
克鲁维毕赤酵母 P. kluyveri	细胞核：10.96—12.40 Mb；线粒体：43.1 kb Nucleus: 10.96-12.40 Mb; Mitochondria: 43.1 kb	3	二倍体，1个子囊4个孢子 Diploid, 1 ascus 4 spores	无 None
库德毕赤酵母 P. kudriavzevii	细胞核：7.25—12.94 Mb，5条染色体；线粒体：50 kb Nucleus: 7.25-12.94 Mb, 5 Chromosomes; Mitochondria: 50 kb	33	二倍体，1个子囊1—2个孢子 Diploid, 1 ascus 1-2 spores	无 None
特立科拉毕赤酵母 P. terricola	细胞核：11.01—14.04 Mb Nucleus: 11.01-14.04 Mb	6	倍性未知，1个子囊1—4个孢子 Ploidy unknown, 1 ascus 1-4 spores	无 None
胶红酵母 R. mucilaginosa	细胞核：19.04—20.22 Mb；线粒体：47 kb Nucleus: 19.04-20.22 Mb; Mitochondria: 47 kb	102	未知 Unknown	无 None
星形假丝酵母 S. bacillaris	细胞核：7.27—9.36 Mb，3条染色体；线粒体：23 kb Nucleus: 7.27-9.36 Mb, 3 Chromosomes; Mitochondria: 23 kb	5	倍性未知可能是单倍体，无证据表明具备形成孢子的能力 Ploidy unknown probably haploid, no evidence of ability to form spores	十个微卫星位点^[86] Ten microsatellite loci^[86]
克拉通覆膜孢酵母 S. crataegensis	细胞核：15.37 Mb Nucleus: 15.37 Mb	1	倍性未知可能是单倍体，偶尔形成子囊， 1个子囊2个孢子 Ploidy unknown probably haploid, occasionally forming ascus, 1 ascus 2 spores	无 None
日本裂殖酵母 S. japonicus	细胞核：11.54—11.73 Mb；线粒体：80 kb Nucleus: 11.54-11.73 Mb; Mitochondria: 80 kb	2	倍性未知，1个子囊6—8个孢子 Ploidy unknown, 1 ascus 6-8 spores	无 None
粟酒裂殖酵母 S. pombe	细胞核：12.59—13.15 Mb，3条染色体；线粒体：20 kb Nucleus: 12.59-13.15 Mb, 3 Chromosomes; Mitochondria: 20 kb	20	倍性未知，1个子囊2—4个孢子 Ploidy unknown, 1 ascus 2-4 spores	无 None
戴尔有孢圆酵母 T. delbrueckii	细胞核：9.00—11.53 Mb，8条染色体；线粒体：28—45 kb Nucleus: 9.00-11.53 Mb, 8 Chromosomes; Mitochondria: 28-45 kb	77	倍性未知可能是二倍体，1个子囊1个孢子，偶尔2—3个孢子 Ploidy unknown probably diploid, 1 ascus 1 spore, occasionally 2-3 spores	8个微卫星位点^[87] Eight microsatellite loci^[87]
异常威克汉姆酵母 W.anomalus	细胞核：12.72—26.55 Mb Nucleus: 12.72-26.55 Mb	10	单倍体和二倍体，1个子囊1-4个孢子 Haploid and diploid, 1 ascus 1-4 spores	无 None
拜耳接合酵母 Z. bailii	细胞核：10.27—21.14 Mb，5—13条染色体；线粒体：29.5 kb Nucleus: 10.27-21.14 Mb, 5-13 Chromosomes; Mitochondria: 29.5 kb	3	单倍体和二倍体，1个子囊1—4个孢子 Haploid and diploid, 1 ascus 1-4 spores	无 None

表2

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非酿酒酵母属酵母的葡萄酒发酵应用研究进展

Research Progress on the Application of Non-Saccharomyces During Wine Fermentation

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