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Bioethanol Production Based on Saccharomyces cerevisiae: Opportunities and Challenges

Tianjin Key Laboratory of Food Biotechnology, College of Biotechnology and Food Science, Tianjin University of Commerce, Tianjin 300134, China
Author to whom correspondence should be addressed.
Fermentation 2023, 9(8), 709;
Submission received: 29 June 2023 / Revised: 19 July 2023 / Accepted: 22 July 2023 / Published: 26 July 2023
(This article belongs to the Special Issue Yeast for the Production of Biochemicals and Biofuels)


The large consumption of non-renewable fossil fuels has brought about energy depletion and environmental pollution, spawning the production of renewable biofuels, an important alternative to alleviate the energy crisis effectively. As one of the ideal types of biofuel, bioethanol synthesis in Saccharomyces cerevisiae has attracted much attention. S. cerevisiae has been developed as essential chassis cells with high efficiency for bioethanol synthesis on account of many advantages. This study systematically summarized the preponderance of S. cerevisiae in biosynthesis. It objectively stated the research strategies of bioethanol synthesis based on S. cerevisiae and the existing bottleneck problems. This study further proposed reasonable prospects for bioethanol synthesis by S. cerevisiae, attempting to provide alternative research strategies.

1. Introduction

The increasing demand for fossil fuels caused by burgeoning anthropogenic activities and rapid economic growth provoked wicked environmental issues and resource depletion [1,2], which is a direct boost to reconstruct the energy structure, develop and industrialize renewable biofuels [3,4,5].
Biofuels produce in response to the proper time and conditions coping with world environmental concerns and the exhaustion of non-renewable fossil-based fuels [6,7,8]. Biofuel refers to the renewable and sustainable fuel obtained through the processing of biomass materials (crop straw, wood, wheat grass, etc.) that can replace traditional fossil fuels [9], among which bioethanol is particularly attractive, having the potential to accelerate sustainable use of resources and change the global economy toward a greener future [10,11,12].
Continuous biotechnology innovation strongly promotes the upgrading and mass production of biofuels represented by bioethanol. Bio-fermentation based on important model microorganisms is a technology with great development potential beyond all doubt for biofuel production at present and in the future [13,14].
With the rapid development of synthetic biology technology, based on the huge market demand for biofuels, Saccharomyces cerevisiae (S. cerevisiae) is increasingly used in the biosynthesis of biofuels due to their superiorities [15,16]. S. cerevisiae is a food-grade budding yeast eukaryote inextricably linked with human production and life [17], which can be effortlessly found in both natural habitats and various environments affected by human activities [18]. Since ancient times, S. cerevisiae has had a long historical standing in human civilization and social development, mainly reflected in food production and fermentation such as bread, beer, and wine [19,20].
Based on numerous studies in this field, this research took bioethanol as an example, reviewed the significant advantages of S. cerevisiae as a quite efficient synthetic cell factory and the main techniques currently used to improve the production of bioethanol. In addition, the bottlenecks existing in the current bioethanol synthesis process using S. cerevisiae as chassis cells will be discussed in this paper, and a rational and feasible future outlook will be proposed. Hopefully, this study could provide the necessary basis for further research on bioethanol synthesis based on S. cerevisiae.

2. Saccharomyces cerevisiae: An Efficient Cell Factory

S. cerevisiae has conceivably received increasing attention in recent years in consideration of its inseparable relations with humankind. Advances in biotechnology, increased demand for synthetic biological products and dynamic environmental changes have enabled the continuous updating and optimization of S. cerevisiae strains [21]. With long-term selection and domestication, S. cerevisiae strains with specific functions have been gradually selected in the expected direction from the wild types [22]. Nowadays, S. cerevisiae is no longer confined to the fermentation of food and drink but has been designed as a cell factory for producing important pharmaceuticals, recombinant proteins and advanced biofuels based on the continuous innovation of different technical approaches [23,24,25,26]. S. cerevisiae has several prominent advantages compared with other prokaryotic and eukaryotic microorganisms, making it one of the most important cell factories today (Figure 1).

2.1. Finite Biosafety Concerns

S. cerevisiae is closely related to human diet and life, completely non-pathogenic, and has been confirmed as a food-grade microorganism in the long history [27]. In addition to the function of food fermentation, studies in recent years have expounded that S. cerevisiae also exhibits obvious probiotic properties. A study by Sun et al. showed that the application of engineered S. cerevisiae could attenuate DSS-induced colitis in mice via the suppression of macrophage pyroptosis and modulation of the intestinal microbiota [28]. S. cerevisiae also has significant probiotic effects on farmed animals. The feed digestibility and the number of pathogenic bacteria could be effectively improved by supplementing live S. cerevisiae cells with animal feed [29]. There is also direct evidence about S. cerevisiae-based probiotics as a novel anti-microbial agent for treating bacterial diseases [30,31]. In addition, S. cerevisiae has a significant biological control effect on citrus population disease through nutritional competition and the production of antifungal compounds [32]. Therefore, the high biosafety and probiotic activity are prominent advantage that distinguishes S. cerevisiae from other organisms, displaying great potential and necessary prerequisites as a cell factory.

2.2. Applicable Evolutionary Position

S. cerevisiae is a kind of single-celled eukaryotes, which not only has the characteristics of easy culture, rapid reproduction and toilless genetic manipulation similar to prokaryotes but also has the basic molecular and cell biological characteristics of typical eukaryotes. The applicable evolutionary position of S. cerevisiae has made it an indispensable experimental model for elaborating many regularities in modern genetics, cell biology and biochemistry of both eukaryotic and prokaryotic cells [33]. Also, S. cerevisiae is considered the most promising tool for exogenous gene expression [34,35], essential for remodeling biofuels’ in vivo synthetic pathways. Evidence also indicated that S. cerevisiae is helpful as a host for genetic engineering since it allows the folding and glycosylation of expressed heterologous eukaryotic proteins and can be subjected to many genetic manipulations [36].

2.3. Undemanding Cultivation Conditions

S. cerevisiae is relatively easy to be cultured on different types of media under laboratory conditions, allowing the evaluation of many phenotypes and the construction of different types of cell factories [37]. Generally, S. cerevisiae does not need a complex medium for growth and can be cultured in liquid and solid mediums. The nutrients required by almost all types of S. cerevisiae strains, including nutrient-deficient and gene-deletion mutants, are common, inexpensive, and readily available. The most commonly used media for S. cerevisiae are YPD (also called YPED) media, SC media, minimal media (supplemented with essential amino acids) and sporulation media [38]. Given the certainty of the nutrient requirements of S. cerevisiae, researchers can conduct in-depth research, such as regulating the nutrient metabolic utilization pathway and corresponding metabolites to improve product quality and yield [39,40,41].

2.4. Strong Environmental Stress Tolerance

The tolerance of microorganisms to diverse stresses is very important for practical applications [42,43]. The living habitat of S. cerevisiae is complex and extensive, always facing a variety of adverse environmental perturbations. S. cerevisiae has developed a capacity to cope with harsh environments in long-term natural evolution and artificial application [44,45,46,47]. Based on the physiological and genetic characteristics of strong tolerance of S. cerevisiae, the rational modification could be carried out through genetic engineering to obtain more robust strains to cope with acute environmental changes, including oxidative stress, low temperature, heat shock, nutrient dropout, osmotic stress, antibiotic pressure, acetic acid stress, etc., in practical application [45,48,49,50,51].

2.5. Ease of Manipulation

As a most thoroughly studied eukaryotic model organism, research has been conducted on S. cerevisiae based on the well-annotated genome information. Mature genome manipulation and gene editing techniques for S. cerevisiae led to easy operation of building specific cell factories [52]. S. cerevisiae strain S288c was the first eukaryote to be genetically elucidated in 1996, which for the first time opened the opportunity for the global study of the expression and functioning of the eukaryotic genome [53] and created opportunities for further studies in comparative, functional, and evolutionary genomics, laying a solid foundation of systematically understanding and rationally engineering metabolism pathways [54]. On this basis, S. cerevisiae can be studied in depth and detail at multiple levels in the follow-up study [55]. The remarkable plasticity of the S. cerevisiae genome makes it possible for the elaboration of synthetic pathways and large-scale production of biofuels [18,33].

3. Synthesis of Bioethanol in Saccharomyces cerevisiae

Biofuel has the potential to achieve environmentally sustainable development, reduce reliance on imported resources, and meet the energy demand with economic growth [3,7]. Figure 2A shows the biofuel production levels in petajoules (PJ; 1 PJ = 1015 J) in leading countries in 2021, with relatively noteworthy differences between different countries. Bioethanol is a particularly compelling class of biofuels, considered a prominent alternative to fossil fuels in the 21st century due to its advantages of complete combustion, low exhaust emissions and contribution to the reduction of crude oil consumption and environmental pollution [56]. The current situation of bioethanol production in major countries in the world is shown in Figure 2B.
Bioethanol is mainly produced by microorganisms in a fermentation process that utilizes plant feedstock as raw materials (e.g., corn, sugarcane, sugar beets, sweet sorghum) [57]. According to the different production raw materials, bioethanol production can be divided into three categories: the use of grain, the use of lignocellulose and the use of algae for bioethanol production [58,59]. There are several microbial cell factories to choose from for fermenting the product ethanol based on different raw materials, including Escherichia coli [60,61], Zymomonas mobilis [62,63], Bacillus subtilis [64,65], S. cerevisiae [66,67], etc. Within all fermentative microorganisms, S. cerevisiae is regarded as an excellent industrial ethanologenic organism based on the outstanding advantages mentioned above. The process relies on the ability of S. cerevisiae to efficiently and completely ferment sugars from feedstock biomass into ethanol on a large scale [68].
Based on the latest research results, the main measures taken to produce bioethanol using S. cerevisiae as chassis cells can be summarized as follows (Figure 3).

3.1. The Addition of External Stimulants to the Medium

The nutritional conditions required by the growth and metabolism of S. cerevisiae are relatively simple. In fermentation to produce ethanol from biomass raw materials, adding exogenous nutrients in the medium is an effective stimulus to increase the ethanol yield. A study by Li et al. showed that adding collagen peptide to the media significantly increased the bioethanol yield under different glucose concentrations and fermentation times compared with the non-added group [69]. In second-generation bioethanol production, the finite tolerance of S. cerevisiae to the inhibitors in lignocellulosic hydrolysates remains a significant challenge. Adding a mixture of pyridoxine, thiamine, and biotin to propagation media could improve cell growth and ethanol yields during fermentation in both corn stover and wheat straw hydrolysate to a large extent [70]. This method is simple and easy and can achieve a significant increase in yield in a short time.

3.2. Optimization of S. cerevisiae Culture Components and Systems

Rational optimization and improvement of culture conditions and systems could effectively increase bioethanol production. A study by Pereira et al. showed an optimized medium based on corn steep liquor and other low-cost nutrient sources that significantly increased the final ethanol titer, yield, and yeast activity, providing valuable insights into cost-effective nutritional supplementation of bioethanol production [71]. In addition, the innovation of the fermentation culture system is also a helpful practice worth trying. By co-culture of S. cerevisiae and Pichia pastoris, the yield of bioethanol can be increased to a great extent [72]. This method of increasing yield requires trial and error to determine the most appropriate medium component and mixed culture system.

3.3. Breeding of High-Tolerance Strains

As a chassis cell, S. cerevisiae faces complex and varied pressures in bioethanol fermentation. Physicochemical conditions, substrate concentration, toxic effects of ethanol and other factors are essential elements affecting the final yield of bioethanol. Therefore, the screening and breeding S. cerevisiae strains with better tolerance is very important.
At present, the breeding of S. cerevisiae strains is mainly focused on improving the thermo-tolerance, glucose-tolerance, and ethanol tolerance, which is the most common type of stress faced by S. cerevisiae cells in the process of bioethanol fermentation [73,74]. The breeding of excellent strains is an essential prerequisite for achieving high bioethanol yield, and that’s why a considerable amount of research is currently focused on this through various physicochemical methods.

3.4. Precise Modification of the S. cerevisiae Genome

To increase ethanol yield, it is a research hotspot to edit and modify the genome of S. cerevisiae accurately from different metabolic pathways based on the transparent genetic background and mature gene operating system. There is a lot of relevant research, and remarkable progress has been made. Modification, knock-out and overexpression of key genes and promoters may be closely related to bioethanol yield. For example, disrupting the alcohol dehydrogenase (ADH2) gene via complete deletion of the gene and introducing a frameshift mutation in the ADH2 locus via CRISPR/Cas9 technology showed that the ethanol yield improved by up to 74.7% compared with the yield obtained using the native strain [75]; the introduction of glucose-proton symporter, fructose-proton symporter and extracellular invertase under the context of deletion of genes encoding hexose transporters, disaccharide transporters and disaccharide hydrolases resulting in a 16.6% increased anaerobic ethanol yield [76]; the knock-in and knock-out of key genes in the glucose metabolic pathway are also crucial for increasing bioethanol production [77,78]. This approach is more precise, more targeted, and more thorough, illustrating the applicability to promote the improvement of bioethanol production in S. cerevisiae via genome engineering.

4. The Bottleneck of Producing Bioethanol by Saccharomyces cerevisiae

Yeasts such as S. cerevisiae have been used in bioethanol production, especially in the brewery and wine industries, thousands of years ago. Numerous efforts have been aimed at comprehending and further improving yeast fermentation. Nevertheless, bioethanol production by fermentation in S. cerevisiae is not without obstacles. There are many external constraints, including environmental conditions, price factors, policy background, scale limitation, and complexity and uncertainty of the fermentation process, and these factors have evolved into the main bottleneck problems, inevitably preventing the increase of ethanol production. Based on the existing research results, some representative bottlenecks are summarized below (Figure 4).

4.1. The Utilization Dilemma of Fermentation Raw Materials

The first generation of bioethanol is produced from food crops (corn, wheat, sweet potato, etc.) with excellent fermentation characteristics [79]. However, food crops are extremely important resources with high production and storage costs, mainly reflected in using valuable and scarce available farmland and irrigation water. Therefore, in terms of cost, producing bioethanol from food crops is not an option based on the lowest cost and highest return.
Lignocellulosic biomass represents the largest resource pool worldwide and is the raw material for producing second-generation bioethanol with low cost and wide sources. One outstanding advantage of lignocellulosic biomass is that it can be obtained for ethanol production without competing for arable land and agricultural inputs with crops for human or livestock consumption [80]. However, despite intensive research exploring lignocellulosic ethanol, this option still accounts for <1% of global ethanol production [68]. The major drawback of these feedstocks is the recalcitrance to degradation of the lignocellulosic matrix, which is comprised of covalently and hydrogen-bonded cellulose and hemicellulose polymers that are further linked to lignin in its natural state [81,82,83]. In addition, S. cerevisiae, the most widely used ethanol-producing species in bioethanol production, has difficulty utilizing β-D-xylose and α-L-arabinose, the main pentoses in hemicellulose polymers [84,85]. This is an important reason that makes it impossible to expand the production of second-generation bioethanol further.
The third generation of bioethanol production is derived from microalgal biomass. Microalgae is currently a promising option for producing new forms of renewable energy with a wide range of sources and rich nutrients. Nevertheless, due to the particularity and complexity of microalgae, the process of using microalgae raw materials to produce bioethanol is relatively complicated, which needs to go through several steps such as cell wall breaking treatment, starch extraction, hydrolysis saccharification and the final fermentation process [86]. Besides, compared with the equipment and process for producing bioethanol from food crops, the production equipment of microalgal biomass for bioethanol synthesis is not mature enough, and many key technologies are still in the theoretical research stage. Also, how to effectively prevent bacterial contamination become another significant constraint in mass cultivation and impedes the industrial process [87].
At present, the largest source of feedstock for the production of bioethanol is still food crops [88]. How to crack the bottleneck of raw material utilization in bioethanol synthesis is a problem that needs to be solved in the future.

4.2. Limitations of Gene Editing Techniques in Saccharomyces cerevisiae

CRISPR-Cas9 gene editing technology to identify the target genome sequence through artificially designed sgRNA (guide RNA). It guides Cas9 protease to cut the DNA double-strand, resulting in double-strand breaks effectively. The CRISPR-Cas9 system of S. cerevisiae is mature enough compared to other chassis cells in synthetic biology, but there are still some problems impeding practical application. For example, chromatin affects the gene-editing efficiency of the Cas9 protein in S. cerevisiae because PAM accessibility in chromatin is a critical factor regulating Cas9 targeting and cleavage [89]. In addition, Cas9-based gene editing can sometimes lead to excessive cutting of DNA, which can lead to some mutations. gRNA is crucial for editing activity, but how to avoid unnecessary side effects of gene editing by rationalizing gRNA modification remains unclear. The low efficiency of the HDR-mediated genome will also significantly affect the efficiency of gene editing. This is undoubtedly a problem that needs to be paid attention to the improving bioethanol production through further genetic modification [90].
In conclusion, a gene-editing system of S. cerevisiae must be better constructed for improved bioethanol production.

4.3. Factors of Yield Constraint

The efficient production of bioethanol is desired for large-scale bioenergy applications, but it is severely challenged by ethanol stress, which is a mutual problem in the global context [91]. Bioethanol accumulates continuously and reaches a higher concentration in fermentation by S. cerevisiae. Although S. cerevisiae has been modified through mutagenic breeding, gene editing and other technologies to improve ethanol tolerance, it is still far from the expected value. It cannot reach the scale fermentation level of higher density.
On the other hand, looking at bioethanol production worldwide, it is distinctly found that the degree of marketization is appreciably high, with fierce competition and development gaps between different countries and regions [92]. In 2020 and 2021, the global bioethanol fermentation industry capacity showed a relatively downward trend compared to 2019 and 2020 (Table 1). Among the major countries producing bioethanol, the overall production of China, Brazil and Thailand has shown a downturn in the past two years. In addition to the impact of macroeconomic regulation policies and the effect of the COVID-19 pandemic, specific agricultural development among countries also significantly affects bioethanol production. Taking China as an example, it is difficult to form a large-scale and stable supply of bioethanol raw materials of first-generation bioethanol in China in terms of the challenge of food security, which is the biggest obstacle to bioethanol industrialization in China at present [93].
Figure 4. Main bottlenecks of bioethanol production by Saccharomyces cerevisiae.
Figure 4. Main bottlenecks of bioethanol production by Saccharomyces cerevisiae.
Fermentation 09 00709 g004

5. Future Perspectives

Global concerns about fossil fuel depletion and the environmental effects of greenhouse gas emissions have led to widespread fermentation-based production of bioethanol [94]. S. cerevisiae, the preferred microorganism for bioethanol production, given its convenient cultivation and genetic manipulations, can use biomass raw materials to synthesize bioethanol through specific metabolic pathways more effectively. Until now, bioethanol fermentation in S. cerevisiae represents the predominant product of industrial biotechnology [95,96].
While seeing the remarkable advantages of bioethanol production by S. cerevisiae, it is also urgent to recognize the significant challenges. To make substantial improvements better, the following aspects could be considered and strengthened in future research.

5.1. Redesign of Fermentation Culture System

A good fermentation culture system is critical to increase the production of bioethanol. As mentioned above, microalgae are an essential raw material for third-generation bioethanol production, but there is still a long way to go before their widespread application. Future research on microalgae culture equipment should be strengthened to promote the transformation and upgrading of microalgal raw materials and the construction of a resource-saving and environment-friendly society. For example, to obtain more microalgal biomass, new types of simple photobioreactors suitable for high-density cultivation of microalgae with low cost should be developed, as well as a series of equipment that can be scaled up for harvesting and cell wall breaking to reduce the drying of microalgae and the extraction of carbohydrates as much as possible, saving equipment investment and energy consumption [97].
Furthermore, the symbiotic system of microorganisms and microalgae has become a research hotspot in recent years due to the strong carbon sequestration capacity and rich nutrient factors of microalgae, which can provide most of the essential carbon sources as well as microelements required for microbial growth. There is some research on the symbiotic system of bacteria and algae [98,99], but there are few reports on its application to bioethanol fermentation. A novel symbiotic system between S. cerevisiae and microalgae could be constructed to optimise culture conditions and increase bioethanol yield effectively.

5.2. Targeted Regulation of Fermentation Pretreatment Process

It has been shown that the efficient pretreatment step of raw material fermentation is essential for converting renewable biomass into fuels and chemicals. Therefore, it is a practical scheme to modify fermentation raw materials by directional control of the pretreatment process. The direction design optimization based on machine learning can effectively regulate the pretreatment parameters [100]. The organic coupling of this technology with the pretreatment process of fermentation raw materials could be considered to improve the bioethanol yield continuously. For example, xylose is one of the most abundant sugars in cellulosic biomass but cannot be utilized by wild-type S. cerevisiae [100,101,102]. In the future, based on machine learning and reverse design optimization strategy, the fermentation pretreatment process of raw materials could be continuously optimized and improved to enhance the xylose utilization and conversion efficiency.

5.3. In-Depth Exploration of Important Regulatory Factors

Nowadays, the genetic manipulation of S. cerevisiae strains is very mature, and some metabolic pathways related to ethanol synthesis have been analyzed in depth and detail. Nevertheless, there is still a lack of mining and identification of some important regulatory elements, such as non-coding RNA (ncRNA). ncRNAs cannot be translated into functional proteins but confer important regulatory functions [103]. Many studies have shown that ncRNAs play crucial regulatory roles in all microbial growth and metabolism stages, and S. cerevisiae is no exception [104]. However, there are no relevant studies on whether ncRNA plays an important regulatory role in the bioethanol synthesis pathways of S. cerevisiae. Future studies could emphasize identifying key regulatory elements like ncRNAs closely related to bioethanol metabolism and synthesis in S. cerevisiae genome through high-throughput sequencing and -omics technology so that they could be flexibly used as crucial regulatory biological elements.

5.4. Construction of Cell-Free Synthetic Biological System Based on Saccharomyces cerevisiae Cell Extracts

Due to the complexity of the living cell system, the irrevocability of cell growth, the interference of intracellular noise and the obstruction of the cell membrane, the transformation of biological elements is greatly limited. Hence, the current bioethanol synthesis based on S. cerevisiae still faces problems such as unpredictability, incompatibility, and high complexity that cannot be underestimated.
In recent years, cell-free synthetic biological (CFSB) systems referring to the engineering science that implements the central principles of biology in vitro have come into sight, instructively complementing the study of synthetic biology. The core of this system is to break the shackles of cells, reintegrate cell resources in vitro, and focus on synthesising customized target products [105]. CFSB systems show significant advantages in toxicity tolerance, economic cost, synthetic efficiency, etc. CFSB systems based on cell extracts are widely used technical means at present, among which the CFSB systems based on E. coli cell extracts and wheat germ cell extracts are the most mature among prokaryotic and eukaryotic systems, respectively [106,107]. There is also some research on the cell-free synthesis system of S. cerevisiae, but they are just starting and not deep enough [108,109]. In the future, efficient CFSB systems based on S. cerevisiae cell extracts can be further explored and constructed to lay the specific foundation for providing maximum synthesis efficiency of bioethanol.

5.5. Exploration of Quorum Sensing in Bioethanol Fermentation

Nongenetic approaches to alter metabolism may have the advantages of general applicability and simple control. Quorum sensing is an essential way for bacteria to carry out intra-species or inter-species communication, which uses the secretion and transmission of signal molecules to make bacteria respond to changes in cell density and flora composition in the environment [110,111,112]. Because of its relatively simple structure and transparent mechanism, the system is often used as a gene module to characterize the complex intracellular response mechanism and the interaction of different bacteria, which has far-reaching significance for synthetic biology research [113]. Quorum sensing also plays an essential role in the process of bioethanol synthesis by S. cerevisiae. Studies showed that the ethanol yield could be improved by adding quorum-sensing molecules to inhibit the cell growth of S. cerevisiae [114]. However, it is unknown whether quorum sensing plays an important role and how it works in the mixed fermentation system of microorganisms and microalgae. In the future, it is worth further exploring and elaborating on whether quorum sensing could realize the interactive regulation of the two communities (e.g., S. cerevisiae and microalgae) in the fermentation system while regulating the spatiotemporal behavior of the S. cerevisiae community in the process of bioethanol fermentation, to provide new insights for improving the bioethanol production.

Author Contributions

H.Z. contributed to the writing and editing of this manuscript. P.Z. and T.W. contributed to the investigation of this study. H.R. contributed to the development and correction of this manuscript. She is also the corresponding author of this paper. All authors have read and agreed to the published version of the manuscript.


This research was funded by Tianjin Education Commission Scientific Research Project (grant number 2022KJ004), Tianjin Municipal Science and Technology Bureau (grant number 22ZXJBSN00010) and National Key Research and Development Program (grant number SQ2022YFE013001).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Conflicts of Interest

The authors declare no conflict of interest.


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Figure 1. Main advantages of Saccharomyces cerevisiae as an efficient cell factory.
Figure 1. Main advantages of Saccharomyces cerevisiae as an efficient cell factory.
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Figure 2. Leading countries based on biofuel (A) and bioethanol (B) production worldwide in 2021, respectively. (Data source: accessed on 25 June 2023).
Figure 2. Leading countries based on biofuel (A) and bioethanol (B) production worldwide in 2021, respectively. (Data source: accessed on 25 June 2023).
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Figure 3. Major strategies for promoting bioethanol synthesis in Saccharomyces cerevisiae.
Figure 3. Major strategies for promoting bioethanol synthesis in Saccharomyces cerevisiae.
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Table 1. Annual world fuel ethanol production (Mil. Gal.) (Data source: accessed on 25 June 2023).
Table 1. Annual world fuel ethanol production (Mil. Gal.) (Data source: accessed on 25 June 2023).
Region201620172018201920202021% of World Production
United States15,41315,93616,09115,77813,94115,01655%
European Union1190125013001350128013505%
Rest of World5876447096556508203%
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Zhang, H.; Zhang, P.; Wu, T.; Ruan, H. Bioethanol Production Based on Saccharomyces cerevisiae: Opportunities and Challenges. Fermentation 2023, 9, 709.

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Zhang H, Zhang P, Wu T, Ruan H. Bioethanol Production Based on Saccharomyces cerevisiae: Opportunities and Challenges. Fermentation. 2023; 9(8):709.

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Zhang, Hongyang, Pengcheng Zhang, Tao Wu, and Haihua Ruan. 2023. "Bioethanol Production Based on Saccharomyces cerevisiae: Opportunities and Challenges" Fermentation 9, no. 8: 709.

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