General features to be considered in the selection of wine yeast.
\\n\\n
Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
\\n"}]',published:!0,mainMedia:{caption:"Highly Cited",originalUrl:"/media/original/117"}},components:[{type:"htmlEditorComponent",content:'IntechOpen is proud to announce that 191 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\nThroughout the years, the list has named a total of 261 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\nReleased this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
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In oenology, the availability of yeasts able to drive alcoholic fermentation (AF) process and bacteria that efficiently carry out malolactic fermentation is required. In fact, in the first phase of the wine production process the yeasts, mostly belonging to the genus
In the past, fermentation of fruit juice, like those of apple and pear to produce cider, grape to obtain wine, or grains to make beer and so on for any kind of alcoholic beverages, have carried out by indigenous and naturally occurring microorganisms present in the original “must” [3, 4, 5].
The first molecular evidence in a Chinese Neolithic village, dated back to 7000 BC, shows that the food processing activity has given rise, without awareness, to the evolution of the genus
Since the discovery of fermented beverages, their production process has undergone many evolutions, but initially the role of the microorganisms was unknown. Only in a second moment the choice of the best microorganisms to be used in a specific production, and their genetic improvement, become a conscious option. Hence, a certain degree of genetic yeast improvement was implemented in response to the requirements of wine production processes [3]. In fact, the scientific community proposed to the industry the use of starter cultures, that could be defined as a microbial (bacteria, yeast, mould) preparation containing a large number of live cells or resting forms of at least one species/strain that once added to a raw material leads to the production of a fermented food by accelerating and driving the fermentation process. The starter culture could contain unavoidable residues of additives and culture media [7, 8, 9, 10].
Regarding wine production, until 150 years ago, also the transformation of grape must into wine took place without knowing the biological agent driving the fermentation process. In the usual cellar practices, it was carried out the inoculation of the must with a small amount of matrix from a previous successful fermentation, that in wine production was called “pied de cuve” [9]. In 1864, the role of microorganisms in fermentation was discovered by Louis Pasteur thus paving the way to the modern microbiology. Further research developments, achieved through microbiology, ecology, biochemistry and recently, molecular biology, have elucidated the metabolisms and in particular the biochemical process of alcoholic fermentation (Figure 1), as well as the interactions among microbial communities involved in winemaking, the phylogenetic and taxonomy. Based on this knowledge, the key role of yeasts in determining the quality of wine is now universally accepted [1, 11, 12, 13].
Central metabolisms of alcoholic fermentation in yeasts.
These scientific achievements have made it possible to supply oenological products and starter cultures appropriate for the industry. In fact, beginning from the mid-1960, the production and use of
The importance of the adoption of yeast starter inoculation mainly consists in provide a faster beginning of AF. This is a stable and reproducible wine making procedure and, at the same time, ensures the absence of defects due to unwanted microorganism contamination [3, 9, 11]. The genetic selection of commercial ADY by the industry is based on the identification of specific technological and physiological features (Table 1) [3, 11, 15, 16].
Technological features | Desirable | Undesirable | Depending on process |
---|---|---|---|
Ethanol tolerance | x | ||
Complete fermentation of sugar | x | ||
Fermentation vigour | x | ||
Resistance to SO2 | x | ||
Type of growth in liquid media (Dispersed cells, Aggregates cells, Flocculence, Foam formation, Film formation, Sedimentation speed) | x | ||
Growth at high and low temperature | x | ||
Killer factor | x | ||
Fermentation by-products (e.g Glycerol, 2-Phenyl ethyl acetate, Ethyl butanoate, Isoamyl alcohol, β-Phenylethanol) | x | ||
Volatile acidity, Sulphuric compounds (H2S, SO2) | x | ||
Enzymatic activity (e.g. β-Glucosidase, Esterase, Proteolytic enzymes, Carbon-sulphur lyase) | x | ||
Ethyl carbamate precursor | x | ||
Effect on wine colour | x |
General features to be considered in the selection of wine yeast.
The discovery of DNA, together with the development of molecular techniques further contributed to the taxonomic classification and, in a more practical context, to the identification of useful and spoilage microbes [17].
This also allowed the development of genetic improvement programs aiming at increasing genetic variability using diverse techniques (e.g. intra- or inter-specific hybridization) and by genetic engineering techniques, mainly focused on improving the yeast qualitative characteristics [18, 19, 20]. In the last decades, genetically modified yeast was also obtained by insertion of useful genetic determinants of different species in
More recently, a new technology to engineer the genome of microorganisms, based on CRISPR/Cas9 system, has been developed. Vigentini et al. [23] applied this editing system in engineering of wine yeast to obtain genotypes with low production of urea through the deletion of DNA coding for arginine permease. This character is important because urea represent a precursor of ethyl-carbamate (EC) which is considered probably carcinogenic to humans [23, 24, 25, 26].
Despite these scientific developments, the current appreciation of local, natural and organic food and wines by consumers has led again to the exploitation of spontaneous fermentation [27]. In fact, organic producers and some consumers consider the use of industrial yeast starter as a non-organic or non-natural practice. Moreover, due to the use of the same commercial strain for various wine style in different winemaking geographical areas, a standardisation of wine sensory characteristics is possible and negatively considered. These criticisms are justified, but, on the other hand, a spontaneous fermentation has to deal with the risks of loss quality related to potential stuck, uncontrolled microorganism development, spoilage and off-flavour production. These problems are only partially addressed by technological strategies aimed at controlling the process [8, 9]. Another aspect to be considered is the wine safety: the uncontrolled development of unwanted microorganisms could lead to the production of toxic compounds, such as biogenic amine, ethyl carbamate or mycotoxins which could negatively impact on human health [8, 9, 28].
As reported by the International Organisation of Vine and Wine (OIV), from winemaking point of view, there is a constant requirement to improve the wine style to answer to the consumer’s demand for natural products and to compete in the globalised market [29, 30, 31]. As in the past, even today the scientific answers to these new market demands can be found by moving to specific yeasts selection. Massive propagations of yeast isolated from their own vineyard in order to inoculate the must, is an alternative strategy for winegrowers that combines unique sensory attributes with safe fermentations. Furthermore, the exploitation of indigenous yeasts is emerging as a marketing plan in several wine regions because the wines are perceived with more complex taste and flavour [9, 32].
The research of wild strains of
A strategy to find
Based on these ideas, the approach of propagation of the autochthonous yeasts for wine production encounters the consumer needs as well as the main winemakers’ target: terroir-yeast in the production of more complex tasting wines with a certain stylistic distinction, while preserving quality [36, 37, 38].
The aim of this chapter is to describe the methods applied for the selection of wine yeasts particularly on the indigenous
Considering the oenological objectives described, the selection of indigenous yeasts must be planned and involves experiments aimed to isolate and propagate yeasts, and to test various oenological feature on laboratory and pilot scale (Figure 2).
Scheme of a selection process of indigenous
The vineyard soil would represent a reservoir of genetically different
Several studies on spontaneous fermentations demonstrated the occurrence of an ecological succession with continuous shifts of the microbiota composition until the end of the process [42]. Due to the extreme condition of the must, especially high sugar concentration (250 g/l), low pH (3.5), nutrient availability and high osmotic pressure, the fermentative yeasts result to be more favoured compared to the species coming from the vineyard.
Performing the grape harvest at ripening time allows to obtain a good degree of yeast biodiversity representing an excellent starting point for the strain selection [32, 43]. The practice of experimental scheme of grape sampling may vary according to the vineyard feature and economic considerations. In optimal situation, the criteria that could be respected have been described by Setati et al. [41]. In detail, it’s recommended to:
Pay attention at any factor which can affect the microbiota community of the vineyard: climate conditions, microclimate (cooler and wetter area may contain a greater population of yeasts), geographical location, microbial vectors, vineyard management (conventional, integrated, organic or biodynamic farming), disease and pests, chemical and pesticides treatment, soil management, and so on [41, 45];
Collect bunches in proximity of harvest, in order to take the highest
A good method to sample is based on the Theory of Sampling (TOS); where a two-dimensional yield is linearised into an elongated one-dimensional lot from which to extract samples at equidistant intervals [41].
As general principles, in the environment and in the vineyard agroecosystem too, yeast populations suffer from spatial and temporal fluctuation, so grape samples should be taken in several locations to gather a sufficient amount of
Then, grape bunches should be placed in sterile bags avoiding the contamination with microorganisms unrelated to the sample, and transferred to the laboratory and processed as soon as possible according to the experimental protocol [41].
After the harvest of bunches, the spontaneous fermentation must be started, crushing the grapes. In order to avoid the contamination of the cultures, sterile conditions must be ensured by using sterilised or disposable equipment. In this step, di-ammonium phosphate (DAP) can be used as yeast nutrient and SO2 in the form of potassium metabisulphite can be added to promote the dominance of
Due to the ethanol tolerance of
The dilution of fermenting must or wine at the end of AF is critical to evaluate a reasonable number of colonies in the solid artificial media. However, a compromise with the risk to lose biodiversity with the dilution procedure must be found, so that the sample should represent the yeast population in each vinification. Usually, the sample is diluted until 10−5 or 10−6 and aliquots of these suspensions are plated. Wallestein Laboratory (WL) agar solid media allowing to differentiate among yeast species on the basis of different colours of the colonies is usually used for yeast growth (Figure 3). The incubation temperature must be 24–26° C.
Some
The genotypes loss during the isolation phase, is a problem to deal with during the selection procedure. As the different
One of the main goals in microbiology is to obtain a valid identification of microorganisms. Traditionally, before the application of molecular biology techniques, yeasts have been identified by morphological and physiological criteria. These methods are basically labor-intensive, time-consuming, and usually provide doubtful identifications. This is due to similar colony morphology, to the influence of culture conditions on yeast physiology and to the presence of different teleomorphic and anamorphic forms in the same species [50, 51].
The progress in molecular biology allowed to develop fast and efficient methods to identify both species and strains. Methods based on DNA technique, some of these based on DNA Polymerase Chain Reaction (PCR) proved to be the most effective identification tool. Allozyme patterns, DNA–DNA hybridization, electrophoretic karyotyping, microsatellite analysis, nested-PCR, random amplified polymorphic DNA (RAPD) and mitochondrial DNA restriction analysis are the molecular biology techniques which first contributed to yeast identification [50, 51, 52, 53, 54, 55, 56, 57, 58]. As an example, electrophoretic karyotyping is based on the weight analysis of the yeast entire genome according to the species [52]. Other examples of molecular analysis are: insertion site polymorphism of delta elements, simple nucleotide polymorphism (SNP), amplified fragment length polymorphism (AFLP), intron splice sequence amplification, PCR of intron of mitochondrial genes, ribosomal DNA sequencing [12, 54, 57, 59, 60].
Moreover, the genome of
In this paragraph we will describe more in detail the most relevant techniques for the identification and characterisation of
Quesada and Cenis in 1995 [53] and Baleiras Couto et al. in 1996 [54] used this method in the taxonomic identification of wine yeast strains both at genera and species level [53, 54]. In 2010, Capece et al. have used a RAPD-PCR with M13 primer to execute a fingerprint on 341 isolates obtaining 130 indigenous strains [43]. This technique can be applied both for interspecific and intraspecific characterisation [55]. The advantage of using RAPD is that it is rapid and easy to assay and there is no need of knowing the DNA sequence, but the main drawback is the low reproducibility.
In 1994, some authors focused the attention on mitochondrial DNA (mtDNA) for fast characterisation of
For the identification at species level, the main used technique is based on the amplification of the rDNA Internal Transcribe Spacer (ITS) region and subsequent digestion with restriction enzymes. This is a specific type of RFLP also called Amplified Ribosomal DNA Restriction Analysis (ARDRA). The amplified target region includes the conserved gene coding for the 5.8 rRNA subunit and the two flanking non-coding and variable internal transcribed spacers named ITS1 and ITS2 [64, 65].
This method was described by Guillamón et al. in 1998 [64], Granchi et al. [50] and Esteve-Zarzoso et al. in 1999 [51] and is used in oenological yeast species identification still today [50, 51, 64, 65]. According to Guillamón et al. [64], the method is based on a first step of amplification targeting the nuclear rRNA gene region by using primers ITS1 and ITS4. This region includes the coding zone for the RNA ribosomal 5.8S and two non-coding regions at its ends (ITS1 and ITS2) (Figure 4). PCR products show a high length variation according to the different species leading to a preliminary discrimination among yeasts after agarose gel electrophoresis. The second step consists in PCR product digestion using three enzymes, endonucleases,
Nuclear rRNA gene and region of DNA amplification through PCR using primer ITS1 (5′-TCCGTAGGTGAACCTGCGG-3′) and ITS4 (5′-TCCTCCGCTTATTGATA TGC-3′).
In general, this technique is highly reproducible and allows the discrimination of large number of samples.
Focusing on
Microsatellite markers, based on Simple Sequence Repeats (SSRs) scattered throughout the genome [69, 70, 71, 72, 73], represent the “gold standard” for this discrimination. Microsatellites are short DNA motifs, 2–6 bases (e.g GATA, GACA, etc.), tandemly repeated five to fifty times (Table 2). Their sequence lengths are intra- and interspecific polymorphic across species [56, 69, 70, 71, 72, 73]. Moreover, SSRs are characterised by higher mutation rate than the rest of the genome, representing a formidable tool for the genetic differentiation of
Locus | SSR Motif | Open Reading Frame Coordinates | Primer sequences (FW: forward; |
---|---|---|---|
ScAAT2 | TAA | YBL084c | FW:CAGTCTTATTGCCTTGAACGA |
ScAAt3 | TAA | YDR160w | FW:TGGGAGGAGGGAAATGGACAG |
C5 | GT | VI-210250/210414 | FW:TGACACAATAGCAATGGCCTTCA |
C3 | CAA | YGL139w | FW:CTTTTTATTTACGAGCGGGCCAT |
C8 | TAA | YGL014w | FW:CAGGTCGTTCTAACGTTGGTAAAATG |
C11 | GT | X-518870/519072 | FW:TTCCATCATAACCGTCTGGGATT |
YKR072c | GAC | YKR072c | FW:AGATACAGAAGATAAGAACGAAAA |
SCYOR267c | TGT | YOR267c | FW:TACTAACGTCAACACTGCTGCCAA |
YKL172w | GAA | YKL172w | FW:CAGGACGCTACCGAAGCTCAAAAG |
ScAAT1 | TTA | XIII-86902/87140 | FW:AAGCGTAAGCAATGGTGTAGATACTT |
C4 | TAA+ TAG | XV-110701/110935 | FW:AGGAGAAAAATGCTGTTTATTCTGACC |
C9 | TAA | YOR156c | FW:AAGGGTTCGTAAACATATAACTGGCA |
ScAAT5 | TAA | XVI-897051/8970210 | FW:AGCATAATTGGAGGCAGTAAAGCA |
C6 | CA | XVI-485898/485996 | FW:GTGGCATCATATCTGTCAATTTTATCAC |
YPL009c | CTT | YPL009c | FW:AACCCATTGACCTCGTTACTATCGT |
SC8132X (YPL009C) | GAA | XVI-536776/536705 | FW: GGTGACTCTAACGGCAGAGTGG |
SCPTSY7 | TTA | XIII-86953/87057 | FW: AAAAGCGTAAGCAATGGTGTAGAT |
Some simple sequence repeat motif and primers’ origin and sequence for
In 2016, Börlin M. et al. [74] characterised the population structure of more than 653 isolates of
Rex et al. [76] in 2020 have validated a SSRs molecular markers method for
The strain identification based on SSRs polymorphisms analysis with multiplex PCR application has been used for rapid and low budget procedure too [46]. As an example, Vaudano and Garcia-Moruno [46] performed the typing of 30 commercial wine strains. The discrimination was achieved by performing a multiplex PCR using primers designed on three highly polymorphic loci: SC8132X, YOR267C and SCPTSY7 and subsequent gel electrophoresis and band pattern analysis and comparison.
Then, this analysis was employed in a dominance study between two co-inoculated strain at different temperature of fermentation, 15°C and 20°C. This trial was finalised to control the ability of these
Methods such as the latter can be used for applicative purpose both in oenology and in wild yeasts selection. In particular, molecular marker supports the screening of the large number of yeasts isolated from natural fermentation [75, 76].
When different genotypes have been identified, the analysis of the phenotype represented by physiological tests and micro-vinification assay is the following stage of the procedure. The physiological tests are for example: production of hydrogen sulphide, killer toxin synthesis, SO2 sensitivity, nitrogen requirement [32, 77].
An interesting test consists in the
In micro-vinification, the resulting wine is then evaluated through chemical analysis of basic features and volatile compounds [45]. Then, the behaviour of the native strains selected was monitored on a pilot scale in comparison with a known yeast used as control.
An example of this pilot test has been performed in 2019 in Lebanon and aimed to identify the most efficient indigenous starter from three autochthonous
In any described cases the evaluation of technological characters (Table 1) at the end of AF for each indigenous strain considered was always performed, generally using official OIV methods, standards Methods (ISO) or a multiparameter analyser. The more relevant features to be considered are: fermentation trend, ethanol production (%V/V), total acidity (g/l tartaric acid equivalent), volatile acidity (g/l acetic acid equivalent), pH, free and total SO2 (mg/l), residual sugar (g/l glucose + fructose). For the microbiological stability of wine is essential a residual sugar less than 2 g/l.
Concerning the volatile acidity, it is positive a low-producer yeast, 0.2–0.4 g/l in acetic acid. High producer strains of sulphur compounds are discarded in the selection. SO2 tolerance is a positive selection criterion [79]. The killer factor is traditionally studied, but its relevance is controversial as it seems that under fermentation conditions it has no influence on sensitive yeast [80].
The evaluation of the phenotype concerns also the wine aromatic profile derived from the secondary metabolism of yeasts. The production of volatile compounds is also affected by the composition of must, in particular depending on the biochemical precursors derived from vine variety. For example, the release of the volatile thiol 4-mercapto-4-methylpentan-2-one (4MMP) from its grape-derived cysteine-bound precursor is carried out by enzymes that possess carbon-sulphur lyase activity and it dependents on yeast [15].
Some volatile compounds belong to the category of higher esters and higher alcohols are shown in Table 3 [34, 43, 48, 81, 82, 83, 84, 85, 86, 87, 88]. In wines, esters can be formed by two different processes: fermentative ones, that involve enzymatic esterification performed by yeast, and storage for long periods that leads to chemical esterification. These two processes can concur in the synthesis of the same ester. The concentration in wine ranges from 10 to 20 mg/l. Higher alcohols are produced by yeasts, both from sugars directly and from grape amino acids through the Ehrlich reaction. They are mostly of fermentative origin and can be found in wines in quantities ranging from 150 to 550 mg/l. The main fermentative higher alcohols, part of the so-called “Fusel oils”, are isobutyl alcohol (2-methyl-propan-1-ol) and amyl alcohols (mixture of 2-methyl-butan-1-ol and 3-methyl-butan-1-ol). At concentration lower than 300 mg/l they participate in the aromatic complexity of the wine; at higher concentrations their penetrating odour masks the wine’s aromatic finesse. Acetic esters of these alcohols, especially isoamyl acetate, have a banana fragrance that may play a positive role in the aroma of some young red wines (primeur or nouveau) [79].
Volatile Compound | Aroma descriptor | Olfactory threshold | Concentration in Wine | References |
---|---|---|---|---|
Ethyl acetate | Fruitiness, varnish | 7.5 mg/l* | 22.5–63.5 mg/l | Swiegers et al. 2005 [81] |
Isoamyl acetate | Banana, pear | 0.03 mg/l* | 0.1–3.4 mg/l | Swiegers et al. 2005 [81] |
Ethyl butanoate | Fruity | 0.02 mg/l* | 0.01–1.8 mg/l | Swiegers et al. 2005 [81] |
Ethyl 3-hydroxybutyrate | Fruity, grapefruit, winy | — | — | |
2-Phenyl ethyl acetate | Floral, rose, hyacinth, honey | 0.25 mg/l* | 0–18.5 mg/l | Swiegers et al. 2005 [81] |
Methyl hexanoate | Pineapple | — | — | |
Ethyl hexanoate (ethyl caproate) | Green apple, pineapple | 0.05 mg/l* | 0.03–3.4 mg/l | Swiegers et al. 2005 [81] |
Ethyl 2-methylbutanoate | Strawberry | — | — | |
Ethyl heptanoate | Grape | — | — | |
Ethyl octanoate (ethyl caprylate) | Fruity, floral, wax | 0.02 mg/l* | 0.05–3.8 mg/l | Swiegers et al. 2005 [81] |
Ethyl decanoate (ethyl caprate) | Fruity, apple, soap | 0.2 mg/l** | 0–2.1 mg/l | Swiegers et al. 2005 [81] |
Ethyl dodecanoate (ethyl laurate) | Waxy | — | — | |
Ethyl lactate | Buttery, butterscotch | — | — | |
Propanol | Alcoholic, pungent, harsh, fermented, weak fusel, musty, yeasty | 500 mg/l*** | 9.0–68 mg/l | Swiegers et al. 2005 [81] |
3-Methyl-1-pentanol | Fusel, cognac, wine, cocoa, green, fruity | — | — | |
Butanol | Fusel, spiritous | 150 mg/l* | 0.5–8.5 mg/l | Swiegers et al. 2005 [81] |
Isobutanol | Fusel, Ethereal, winey | 40 mg/l* | 9.0–174 mg/l | Swiegers et al. 2005 [81] |
Isoamyl alcohol | Solvent, Varnish, nail polish, ripe fruit, harsh | 30 mg/l* | 6.0–490 mg/l | Swiegers et al. 2005 [81] |
Amyl alcohol | Almond | — | — | |
1-Hexanol | Mowed grass, herbaceous, green | 4 mg/l*** | 0.3–12.0 mg/l | Swiegers et al. 2005 [81] |
2,3-Butanediol | Fusel, cognac, wine, cocoa, green, fruity | — | — | |
2-Phenylethanol | Dried rose, floral | 10 mg/l* | 4.0–197 mg/l | Swiegers et al. 2005 [81] |
Benzyl alcohol | Jasmine | — | — | |
3-(Methylthio)-propanol (Methionol) | Cauliflower | 1 mg/l**** | 0.17-2.4 mg/l | Ferreira et al. 2000 [82] |
3-Mercapto-1-hexanol | Passion fruit, grapefruit, | 6*10-5 mg/l***** | 0-1.28 * 10-2 mg/l | Tominaga et al. 1998 [83] |
Some volatile compounds from
Aqueous solution 10% ethanol.
Synthetic wine.
Wine.
Red wine.
Aqueous solution 12% ethanol.
Usually, the analysis of volatile is performed by gas chromatography equipped with Mass Spectrometer as detector (GC–MS) [43, 48, 81, 82, 83, 84, 85, 86, 87, 88].
The last examination at the end of a pilot scale production is the sensory evaluation performed by a panel test. That consist in the personal evaluation of wine descriptors fulfilled by a group of judges trained in the recognition of organoleptic features (appearance, odour, taste, texture) (ISO 1993). The panel, in short, quantifies the level of descriptors using an intensity scale as required by the ISO 2003 standard b. The sensory session must be performed in standard condition of the room, glasses, temperature, time, so that the environment does not affect the judges [34, 43, 48, 81, 82, 83, 84, 85, 86, 87, 88]. An example of sensory analysis results is shown in Figure 5. This sensory examination could be useful to predict the consumer appreciation. At the end of this process, all the data obtained by every test must be statistically analysed. The strain or strains which show the best performance and which better meet the enologist’s preferences, can be used in an industrial scale assay.
Comparison of sensory profiles of two (A and B) red wines fermented with two different indigenous strains of
In winemaking, the role of yeast is fundamental for a good fermentation process. There is a high biodiversity among the
In this chapter a workflow aimed to select indigenous
In consideration of the increasing appreciation by consumers of wines connoted by organoleptic complexity also linking with the territory of origin, the selection of indigenous
This chapter is funded by Università degli Studi del Piemonte Orientale Amedeo Avogadro.
The authors declare no conflict of interest.
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