Trends in Food Science & Technology 150 (2024) 104589

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Trends in Food Science & Technology

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Plant-derived food waste management, valorization, and recycling through

sourdough fermentation
Erica Pontonio a, * , Alessandro Stringari b , Raffaella Di Cagno b , Pasquale Filannino a ,
Carlo Giuseppe Rizzello c , Andrea Polo b , Olga Nikoloudaki b , Marco Gobbetti b
a
Department of Soil, Plant, and Food Sciences, University of Bari Aldo Moro, Bari, Italy
b
Faculty of Science and Technology, Free University of Bolzano, Bolzano, Italy
c
Department of Environmental Biology, “Sapienza” University of Rome, Rome, Italy

A R T I C L E I N F O A B S T R A C T

Handling editor: AR Jambrak Background: The food industry generates a vast amount of food waste. Nevertheless, several types of food waste, i.
e. those deriving from fruits, vegetables, grains, and other plant-based food production and processing chains,
Keywords: still contain valuable nutritional and bioactive compounds thus having the potential to be converted into value-
Food waste added products. Several approaches have been investigated as pre-treatment of food waste to improve the
Sourdough fermentation
nutritional, functional, and technological properties before to re-inclusion in food production. Sourdough
Lactic acid bacteria
fermentation, either spontaneous or through selected microbial strains, appears to be a suitable and sustainable
Nutritional properties
Functional features tool for upcycling plant-derived food waste.
Scope and approach: This review reveals the latest insights into the potential of sourdough fermentation to recycle
milling by-products, brewers’ spent grain, wasted bread, and miscellaneous plant wastes.
Key findings and conclusions: Sourdough biotechnology is suitable for improving the sustainability of several food
chains. Nevertheless, due to the significant effect of the presence, growth, and metabolic activity of specific
microorganisms on the quality of the final products, an accurate set-up and optimization of tailored fermentation
processes is highly suggested.

1. Introduction generate more than half of the total food waste (54%) with 70% of food
waste arising at household, food service and retail (Food waste and food
Food waste is any edible or inedible loss from its supply chain waste prevention – estimates, 2023).
(O’Connor et al., 2021). In the European Union, over 58 million tons of Stockpiling of food wastes poses social, environmental, and eco­
food waste (131 kg/inhabitant) are generated annually (Food waste and nomic issues. Global food system causes approximately one-fourth of all
food waste prevention – estimates, 2023). The most abundant part greenhouse gas emissions (FAO, 2019). Food wastes require large-scale
comes from fruits and vegetables, including cereals as the main in­ storage, transportation, and costly disposal applications (FAO, 2019).
gredients for worldwide staple foods, which generate by-products dur­ They have high oxidation potential and water content, and when
ing pre- and post-harvesting, preparation, and processing (FAO, 2019). generated from rendering plants and accidental failure of the cooling
The overall estimation is that one-third of worldwide food is wasted at system occur, they also suffer from contamination by spoiling and/or
various stages along the food supply chain (FAO, 2019). FAO’s Food pathogen agents (ICMSF, 2000). Given this impact, the United Nations
Loss Index estimates that globally, around 14 percent of all food pro­ set the Sustainable Development Goals to reduce food waste by 50% per
duced is lost from the post-harvest stage up to, but excluding, the retail capita by 2030. FAO believes that food waste recycling is the pillar of
stage (FAO, 2019). According to the UNEP Food Waste Index 2021, achieving sustainable development with the concept of a circular
around 931 million tons of food waste were generated in 2019–61% of economy (FAO, 2019).
which came from households, 26% from food service, and 13% from Solutions for food waste concern education programs focused on
retail – suggesting that 17% of global food production may be wasted at consumers’ healthy lifestyles, prevention, improvement of processing
these stages of the food supply chain. Similarly, in the EU, households efficiency, and valorization. Valorization is the most sustainable

* Corresponding author.
E-mail address: [email protected] (E. Pontonio).

https://doi.org/10.1016/j.tifs.2024.104589
Received 15 April 2024; Received in revised form 3 June 2024; Accepted 9 June 2024
Available online 10 June 2024
0924-2244/© 2024 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
E. Pontonio et al. Trends in Food Science & Technology 150 (2024) 104589

approach, which should convert wastes into products with added value, leavening agents. A recent systematic review (Arora et al., 2021) high­
thus limiting their disposal in landfills and eliminating greenhouse gas lighted the versatility of sourdoughs to ferment an extraordinary variety
emissions (Al-Obadi et al., 2022). Currently, food wastes are physically, of cereal, pseudo-cereal, and legume flours, as well as miscellaneous
chemically, or biologically transformed to get an array of biofertilizers agri-food by-products.
and soil amendments. Composting and anaerobic digestion are the main Here, we review the potential of sourdough fermentation to recycle
large-scale conversion techniques to exploit the potential of food wastes milling by-products, brewers’ spent grain, wasted bread, and miscella­
into biofertilizers and soil amendments. Other emerging conversions neous plant wastes. Sourdough fermentation should represent a natural,
such as dehydration, biochar production and chemical hydrolysis show low-cost, sustainable, and flexible approach to create added value
promising potentialities in agriculture and soil remediation (O’Connor within the food chain system (Fig. 1).
et al., 2021). Aiming at a “zero waste economy”, food waste can be used
as raw material for new products and applications. Nevertheless, it was 1.1. Recycling of milling by-products
observed that the food waste needs further processing steps before being
used. Overall, the use of microorganisms that guide the production of Milling is the principal conversion procedure in the cereal industry.
functional ingredients for novel food formulation has been suggested to Dry milling separates by-products such as outer fibrous materials and
be helpful for the development of low-cost bioprocesses of food wasted germs from the grain endosperm. Pearling gradually removes seed coat
and by-products leading to a transition toward a bio-economy model (testa and pericarp), aleurone and sub-aleurone layers, and germ to get
(Sabater et al., 2020). Lactic acid bacteria isolated from sourdough can polished grains. Wet milling delivers starch and gluten, leaving steep
be employed as a biotechnological starter to increase the safety of food solids, germ, and bran as by-products. It is estimated that circa 13% of all
industry by-products, to provide added value, to design the synthesis of food waste comes from these milling procedures, with 30% of the cereal
functional molecules in fermentable substrates, and to moderate the weight basis lost (FAO, 2019). While milling is inevitable for processing
technologies for safer alternative stock (e.g., food waste and baked goods, most, if not all, milling by-products have high biological
by-products) incorporation to the main food (e.g., bread) formulas. and nutritional value to be recycled (Cacace et al., 2022). Chemical
Sourdough is a mixture of flour and water, spontaneously fermented by and/or physical extraction, and purification procedures are options to
lactic acid bacteria and yeasts, and having acidification and leavening separate valuable compounds, but they are quite expensive and
capacities (Arora et al., 2021). Sourdough is one of the oldest examples contribute per se to environmental pollution. On the other hand, the
of natural starters, mostly used for making leavened baked goods as an direct reuse of untreated milling by-products as food ingredients causes
alternative to baker’s yeast and chemical leavening. Almost thirty years poor technological and unpleasant sensory attributes. Sourdough
of literature accumulated on sourdough show the undoubtedly techno­ fermentation, either spontaneous or driven by an ad hoc microbiota
logical, sensory, and nutritional advantages compared to the other (Arora et al., 2021), conjugates the potential to exploit the biological

Fig. 1. Schematic approach of food waste fermentation.

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Table 1
Non-exhaustive list of the main advantages related to the application of spontaneous or selected sourdough fermentation to cereal milling by-products and their related
products.
Cereal source Fermentation type/species/strain Effects References

Bran
Wheat Levilactobacillus brevis E95612 and Kazachstania Increase of peptides and free amino acids concentration; increase of Coda et al. (2014)
exigua C81116 with enzymes protein digestibility, soluble fiber concentration. Decrease of pungent
flavor and bitter taste of fortified bread.
Lacticaseibacillus rhamnosus (R0011, ATCC 9595, Exopolysaccharides synthesis; increase of total phenolic content, Bertsch et al. (2020)
and RW-9595M) and Saccharomyces cerevisiae bioaccessibiity and antioxidant activity.
Lacticaseibacillus rhamnosus 1473 Decrease of phytic acid; increase of soluble arabinoxylans; improvement Spaggiari, Ricci, et al. (2020)
of aroma profile.
Enterococcus faecalis M2 Increase of soluble dietary fiber, alkylresorcinol, flavonoid and total Mao et al. (2020)
phenol contents, and antioxidant activity
Spontaneous fermentation Increase of soluble fiber water-extractable arabinoxylans and free ferulic; Manini et al. (2014)
total degradation of phytic acid.
Commercial baker’s yeast, Lactobacillus Increase of water extractable arabinoxylans and soluble dietary fiber. Zhao et al. (2017)
delbrueckii subsp. bulgaricus and Streptococcus Reduction of phytic acid content. Improvement of hydration properties
thermophilus and flavor.
Levilactobacillus brevis E− 95612 and Candida Increase of protein solubilization and in-vitro digestibility. Increase of Arte et al. (2015)
humilis E− 96250 with enzymes phytase activity and total phenols content.
Kazachstania exigua VTT C-81116 and Increase of free amino acids and phenolic compounds concentration. Savolainen et al. (2014)
Levilactobacillus brevis VTT E− 95612
Spontaneous fermentation Increase of folates and phenols bioavailability; pentosan solubilization. Katina et al. (2012)
Lactobacillus helveticus FAM22155 Reduction of aflatoxin B1 Zhang et al. (2021)
Companilactobacillus paralimentarius (PB 3, PB 4), Increase of radical scavenging activity of wheat-rye bread Mikušová et al. (2013)
Lactobacillus helveticus (PB 7), Levilactobacillus
brevis (PB 12A)
Liquorilactobacillus uvarum, Lacticaseibacillus casei Lowering content of mycotoxin (below the threshold for human Bartkiene, Zokaityte, et al.,
and Lacticaseibacillus paracasei consumption) and biogenic amine. 2021; Zokaityte et al., 2021
Pediococcus acidilactici LUHS29 (DSM 20284) Reduction in DON content in the sourdough by 44–69% and a removal of Zadeike et al. (2021)
15-acetyldeoxynivalenol (15-AcDON), alternariol (AOH),
deoxynivalenol-3-glucoside (D3G), toxins H-2 and HT-2.
Lactobacillus helveticus Decrease of aflatoxin B1 concentration. Zhang et al. (2021)
Wheat, Lactobacillus plantarum T6B10 and Weissella Increase of peptides and total free amino acids; Increase of total phenols Pontonio, Dingeo, et al. (2020)
barley and confusa BAN8 with commercial xylanase and antioxidant activity. Decrease of phytic acid. Improvement of the
emmer nutritional value of bread.
Oat Streptococcus thermophilus, Lacticaseibacillus Folic acid fortification. Korhola et al. (2014)
rhamnosus, Saccharomyces cerevisiae and Candida
milleri
Kazachstnia humilis Increase of fiber solubility. Degutyte-Fomins et al. (2002)
Rice Lacticaseibacillus rhamnosus and S. cerevisiae with Reduction of the cytotoxicity and inhibition of melanogenesis in B16F1 Chung et al. (2009)
enzymes melanoma.
Lactobacillus acidophilus GIM1.731 and Increase of total phenolic content and antioxidant activity. Liu, Cao, et al. (2017)
Lactiplantibacillus plantarum subsp. plantarum GIM
1.648 with enzymes
Weissella koreensis DB1 Increase of ornithine and citrulline content. Yeong et al. (2020)
Increases in total amino acid content and antioxidant activity of bread.
Lactiplantibacillus plantarum EM High cholesterol removal and strong antimicrobial activity. Moon and Chang (2021)
Levilactobacillus brevis Decrease of mycotoxin concentration. Anti-aflatoxigenic effect on Sadeghi et al. (2019)
aflatoxin B1.
Rye Baker’s yeast with enzymes Increase of phenolic compounds in bread. Koistinen et al. (2016)
Limosilactobacillus reuteri TMW 1.106 Increase of amount and speed production of exopolysaccharides. Kaditzky and Vogel (2008)
Weissella confusa Exopolysaccarides synthesis. Kajala et al. (2016)
Germ
Wheat Spontaneous fermentation Improve technological and sensory properties of doughs and breads. Marti et al. (2014)
Lactiplantibacillus plantarum subsp. plantarum Slow down lipid oxidation by decreasing the activity of lipase and Rizzello et al., 2010a,b
DSM 32248 and Furfurilactobacillus rossiae DSM lipoxygenase. Increase of in-vitro protein digestibility, the concentration
32249 of total phenols and amino acids, phytase and antioxidant activities
(sourdough and bread). Improve the texture and sensory properties of
bread.
Ex-vivo anti-proliferative effects colon and ovarian carcinoma cell lines. Rizzello et al. (2013)
Lactiplantibacillus plantarum and Lactobacillus Decrease of the activity of lipase and lipoxygenase and increase in Khosroshahi et al. (2022)
acidophilus antioxidant activity.
Saccharomyces cerevisiae 5022 and Increase of peptide and GABA concentrations and scavenging activity. Bayat et al. (2022)
Lactiplantibacillus plantarum 299v
Lactiplantibacillus plantarum dy-1 Antiproliferative effects and the induction of apoptosis of human HT-29 Zhang et al. (2015)
colon cancer cells.
Latilactobacillus sakei TMW1.22 and Degradation of wheat germ agglutinin. Tovar & Gänzle, 2021
Fructilactobacillus sanfranciscensis DSM20451T
Saccharomyces cerevisiae Cancer-fighting characteristics: inhibition metastatic tumor Boros et al., 2005; Saiko et al.,
dissemination and proliferation. 2009; Comín-Anduix et al.,
2002
Spontaneous fermentation Anti-aging activity. Zhao et al. (2021)
Rice Latilactobacillus sakei Enrich in GABA showing a positive effect on the sleep disturbance of Mabunga et al. (2015)
mice.
Bran and Germ
(continued on next page)

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Table 1 (continued )
Cereal source Fermentation type/species/strain Effects References

Wheat Lactiplantibacillus plantarum subsp. plantarum Increase of fiber content, protein digestibility and nutritional indexes of Pontonio et al. (2017)
DSM 32248 and Furfurilactobacillus rossiae DSM fortified bread. Decrease of glycemic index.
32249
Maize Lactiplantibacillus plantarum subsp. plantarum Increase of the concentrations of free amino acids and peptides, the Pontonio et al. (2019)
T6B10 and Weissella confusa BAN8 antioxidant activity. Degradation of phytate.
Improve content of dietary fibers and proteins, the protein digestibility,
and the starch hydrolysis index in bread
Lactobacillus sakei MI401 and Pediococcus Improve technological properties of albumins and globulins and increase Zadeike et al. (2022)
acidilactici PA-2 the digestibility and free radical scavenging activity of prolamins.
Increase of free amino acids concentration and antioxidant activity.
Rye/wheat Kazachstania unispora and Kazachstania servazii Improve the complexity of the volatile molecules. Increase in short chain Siroli et al. (2022)
and Latilactobacillus curvatus fatty acids, antioxidant activity, total phenol content, bioactive peptides.
Decrease in phytic acid content and an increase in prebiotic activity.

value and eliminate the negative attributes of recycled bran and germ in content of bioavailable ferulic acid by 82% (Manini et al., 2014). Lactic
food preparations (Table 1). acid bacteria, including those populating the sourdough ecosystem,
express ferulic acid esterases (Gaur & Gänzle, 2023). Additionally,
1.1.1. Cereal bran Pediococcus acidilactici was found to significantly increase the contents of
With variations depending on the cereal species, bran is a concen­ caffeic acid and mainly gallic acid during fermentation (Zhang et al.,
trate of dietary fibers (mainly arabinoxylans), phenolic compounds, 2023).
proteins, and amino acids with high biological value, as well as minerals When combined with cell wall degrading enzymes, sourdough spe­
and vitamins (e.g., folates) (Chen et al., 2023). Unfortunately, these cies such as Lv. brevis and Kazachstania humilis abundantly released free
nutritive features has low bio-accessibility for humans, limiting its phenolic compounds (Arte et al., 2015). Lac. rhamnosus metabolized
recycling potential. The bound forms of dietary fibers and phenolic conjugated phenolic compounds and broke the linkage with cell wall
compounds weaken their bioavailability, digestibility and/or intestinal polysaccharides (Spaggiari, Calani, et al., 2020). The wheat bran
absorption (Holland et al., 2020). The formation of insoluble complexes solid-state fermentation that started with Enterococcus faecalis promoted
with phytate (Saurabh et al., 2021) and constraints imposed by the cell a remarkable free radical scavenging activity, which depended on the
wall matrix limit the digestion and bioavailability of proteins (Alzuwaid release of free phenolic compounds (Mao et al., 2020). The sourdough
et al., 2020) and minerals. Furthermore, the direct recycling of bran in fermentation with Lactobacillus acidophilus, Lactiplantibacillus plantarum
food preparations might be responsible for poor hygiene (mycotoxins and hydrolyzing enzymes modified the profile of phenolic compounds in
and other contaminants), and unpleasant rheology and sensory attri­ rice bran, thus increasing the levels of soluble ferulic acid (Liu, Zhang,
butes (Ma et al., 2021). Notwithstanding the potential of other micro­ et al., 2017). This processed rice bran was used as an ingredient for
organisms (Chen et al., 2021; Li et al., 2022; Wu et al., 2022), sourdough making functional foods, which, due to the enrichment of free phenolic
fermentation prior to recycling into food preparations, alone or in acids and flavonoids, showed remarkable antioxidant activity.
combination with enzymes, has the potential to overcome most of these Although the disruption of the cell wall matrix is the main factor for
nutritional, technological, and sensory constraints (Chen et al., 2021; Li increasing the bio-accessibility of bran proteins (Li et al., 2023), sour­
et al., 2022; Wang, Li, et al., 2022a,b). dough fermentation initially solubilized wheat bran proteins because of
The ad hoc fermentation with two typical sourdough species, Levi­ the activation of endogenous proteases at circa pH 4.0 (Arora et al.,
lactobacillus brevis, and Kazachstania exigua, in combination with xyla­ 2021). Following this primary proteolysis, peptidases by sourdough
nase, endoglucanase, and β-glucanase enzymes, was successful in lactic acid bacteria were responsible for the liberation of small-size
breaking the wheat bran cell wall, thus increasing the content of soluble peptides and free amino acids (Christensen et al., 2022), which were
arabinoxylans (Coda et al., 2014). Combining the spontaneous sour­ more readily absorbable at intestinal level comparing to native proteins
dough fermentation and the activity of endogenous or microbial en­ (Khubber et al., 2022). Processing rye bran with enzymes and sourdough
zymes, the same effect was found for rye arabinoxylans. After fermentation prior to recycling into bread making led to high in-vitro
fermentation with sourdough, the bran was recycled into the bread protein digestibility (Nordlund et al., 2013). Sourdough proteolysis also
formula, and volume and texture improved (Courtin & Delcour, 2002). favored the release of biogenic peptides (Pontonio, Verni, et al., 2020;
Usually, the extractability of arabinoxylans increased during sponta­ Verni, Verardo, & Rizzello, 2019) and degraded antinutritional factors
neous sourdough backslopping and reached the highest value when the such as trypsin inhibitors and phytic acid (Patterson et al., 2017; Pon­
microbiota of lactic acid bacteria and yeasts became mature and stable tonio, Dingeo, et al., 2020). Pre-fermented wheat bran was used to
(Manini et al., 2014). Soluble dietary fibers almost quadrupled, free enhance the nutritional profile of bread. Compared to nonfermented
amino acids increased as the hydrolysis of wheat bran proteins pro­ bran, the in-vitro protein digestibility increased by 40%, and enriched
ceeded, and the content of phytate decreased (Mao et al., 2020). Almost breads showed pleasant sensory and textural attributes (Pontonio,
the same efficiency was observed under solid-state fermentation of Verni, et al., 2020). The sourdough fermentation with an ad hoc
wheat bran steered by Lactobacillus delbrueckii subsp. bulgaricus and microbiota was also efficient for the treatment of rice bran. Weissella
Streptococcus thermophilus together with baker’s yeast (Zhao et al., koreensis enriched rice bran with ornithine, which is a healthcare sup­
2017). Mixing wheat bran and whey permeate, another agri-food plement to activate the immune system and liver function (Yeong et al.,
by-product, the fermentation with Lacticaseibacillus rhamnosus and 2020). Water-soluble extracts of sourdough fermented rice bran had an
Saccharomyces cerevisiae allowed the solubilization of fibers and anti-photoaging effect on human skin fibroblasts cultures (Seo et al.,
increased the level of free phenolic compounds (Bertsch et al., 2020). 2010). The rice bran co-fermented with Lac. rhamnosus and S. cerevisiae
The sourdough fermentation does not affect the total content of decreased the cytotoxicity and inhibited melanogenesis in B16F1 mel­
phenolic compounds but elevates the ratio between free and bounded anoma through the downregulation of microphthalmia-associated
forms. The sourdough fermentation of both peeled and native rye bran transcription factors (Chung et al., 2009). Rice bran fermented with
increased the content of free ferulic acid and other hydroxycinnamic Lp. plantarum attenuated the levels of cholesterol (45–68%) and exerted
acids liberated from the polymeric structure (Katina et al., 2007). It was inhibitory activities towards foodborne pathogenic bacteria and spoiling
estimated that the spontaneous sourdough fermentation increased the fungi (Moon & Chang, 2021).

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Cereal endogenous or microbial phytases are responsible for the butyric acid (GABA), a non-protein amino acid with functional features
hydrolysis of phytic acid in cereal bran. Wheat endogenous phytases are (Bayat et al., 2022; Rizzello, Nionelli, Coda, De Angelis, & Gobbetti,
almost inactive at neutral pH, but sourdough acidification promotes 2010; Rizzello, Nionelli, Coda, Di Cagno, & Gobbetti, 2010). One of the
their activation with the consequent decrease of the phytate content and most promising achievements for sourdough fermented wheat germ was
the increase of protein and mineral bioavailability (Ameur et al., 2022). represented by the cytotoxic activity towards cancer cell lines. Wheat
Phytases by several sourdough lactic acid bacteria and yeasts were germ fermented with Lp. plantarum dy-1 inhibited the proliferation of
characterized. Lac. rhamnosus had the capability to degrade phytate in HT-29 cells via apoptosis suggesting its use as a potential anticarcino­
wheat bran (Spaggiari, Ricci, et al., 2020), and Lp. plantarum and genic (Zhang et al., 2015). Avemar®, a commercial product made of
Weissella confusa decreased its level by 25%–60% when used as starters wheat germ fermented with S. cerevisiae (Boros et al., 2005; Saiko et al.,
for wheat bran fermentation (Moon & Chang, 2021; Pontonio, Dingeo, 2009), showed in-vitro anticancer and autoimmune properties on
et al., 2020). Also, the fermentation of rice bran with a functional strain various human cancer cell lines (Comín-Anduix et al., 2002).
of Lp. plantarum EM decreased the content of phytic acid by 50% (Moon Two-methoxy benzoquinone and 2,6-dimethoxybenzoquinone are
& Chang, 2021). The spontaneous sourdough fermentation of bran naturally present in wheat germ but in a glycosylated and non-active
increased the level of folates by over 100% and the highest detectable form. After activation through β-glucosidase activity of Lp. plantarum
levels coincided with the highest cell density of autochthonous lactic and Fu. rossiae, they exerted ex-vivo anti-proliferative effects on colon
acid bacteria and yeasts. Overall, the increase of folate was mainly due and ovarian carcinoma cell lines (Rizzello et al., 2013). Based on the
to synthesis by yeasts, while the liberation by lactic acid bacteria was metabolism of thiols, sourdough fermentation decreased the content of
strain-dependent. The ad hoc substitution of folate-consuming lactic acid agglutinins in wheat germ, which is one of the triggering factors for
bacteria with folate-synthesizing strains significantly increased the non-celiac wheat sensitivity (Tovar & Gänzle, 2021). A sourdough made
vitamin content in sourdough breads enriched with pre-fermented and combining wheat germ and bran was recycled and used in the bread
recycled bran (Mendez-Vilas, 2007). The capability of yeasts to syn­ formula (Pontonio et al., 2017). The bread was enriched in dietary fi­
thesize folate from oat bran was species dependent, with S. cerevisiae, bers, free amino acids, and phenolic compounds, and as shown by
Pseudozyma sp., Rhodotorula glutinis and Kluyveromyces marxianus lead­ in-vitro and in-vivo assays, it was classified as a low glycemic index food.
ing to the highest concentrations (Korhola et al., 2014). Intending to After fermentation with Latilactobacillus sakei, the rice germ fermented
increase the folate concentration in bran processing, sourdough was enriched in GABA, showing a positive effect on the sleep distur­
fermentation with yeasts was always the most efficient option. Oat bran bance of mice (Mabunga et al., 2015). Lp. plantarum and W. confusa were
fermented with St. thermophilus or Lac. rhamnosus, and S. cerevisiae and used to ferment raw maize or heat-treated germ and bran (Pontonio
K. humilis led to a folate concentration of 120 ng/g. The intake of 100 g et al., 2019). The concentrations of free amino acids and peptides, the
of fermented oat bran supplied 15% of the recommended daily dose of antioxidant activity, and the phytate degradation increased during
folate (Korhola et al., 2014). The enrichment of rye and wheat brans sourdough fermentation. Incorporation of these fermented maize
with exopolysaccharides (EPS) is another option with multiple out­ by-products in bread positively affected the content of dietary fibers and
comes. Sourdoughs made with ad hoc lactic acid bacteria, for instance proteins, the protein digestibility, and the starch hydrolysis index
Limosilactobacillus reuteri (Kaditzky & Vogel, 2008) or W. confusa, was (Pontonio et al., 2019).
the most efficient choice. Usually, the in-situ production of EPS in rye
bran reached the concentration of 2–3% on dry matter (Kajala et al., 1.1.3. Mycotoxins degradation
2016) and represented the way to improve the structure-forming capa­ Compared to whole grains, by-products from cereal milling suffer
bility of bran, making it a recyclable ingredient with prebiotic and from higher levels of mycotoxin contamination (Hoffmans et al., 2022).
in-vitro antitumor and immunomodulating activities (Korcz & Varga, Although using implemented good practices, the bio-accumulation of
2021). mycotoxins in milling by-products is almost inevitable, especially under
suboptimal storage conditions (Khodaei et al., 2021). Decontamination
1.1.2. Wheat germ before recycling is a priority. Available options include physical,
The annual world deposit of wheat germ is estimated to be circa chemical, and biological methods, with the latter attracting considerable
25,000,000 tons, which undoubtedly deserves valorization as it corre­ interest (Piotrowska, 2021). Although the effect of sourdough fermen­
sponds to the most nutritious part of the grain (Rizzello, Nionelli, Coda, tation is somewhat controversial, the degradation of aflatoxins, fumo­
De Angelis, & Gobbetti, 2010). The fast development of rancidity nisins, ochratoxins, deoxynivalenol, and zearalenone was demonstrated
because of the presence of unsaturated fats, lipoxygenases and lipases with various species of lactic acid bacteria (Bartkiene, Zokaityte, et al.,
severely limits the use of germ in baked goods processing (Li et al., 2021; Muhialdin et al., 2020; Sadeghi et al., 2019; Zokaityte et al.,
2016). Nowadays, sourdough fermentation is the most efficient solution 2021). The mechanism for mycotoxin decontamination occurs through
to stabilize and improve the nutritional attributes of germ (Khosroshahi the adhesion of significant amounts of mycotoxins to microbial cells
et al., 2022). and/or the degradation of mycotoxins into non-toxic metabolites
Lowering the value of pH by sourdough fermentation inactivates (Muhialdin et al., 2020; Piotrowska, 2021).
lipase and lipoxygenase enzymes (Marti et al., 2014; Rizzello, Nionelli, Extrusion combined with the fermentation by Lacticaseibacillus casei
Coda, Di Cagno, & Gobbetti, 2010). Compared to nonfermented wheat and Lacticaseibacillus paracasei, or Liquorilactobacillus uvarum reduced
germ, the sourdough fermentation with Lp. plantarum and Furfur­ the levels of several mycotoxins in wheat bran to levels below the
ilactobacillus rossiae almost inhibited the synthesis of volatile compounds threshold for human consumption (Bartkiene, Zokaityte, et al., 2021;
from lipid oxidation during 40 days of storage (Rizzello, Nionelli, Coda, Zokaityte et al., 2021). In addition, the sourdough fermentation with Lp.
Di Cagno, & Gobbetti, 2010). Other sourdough starters were proven to plantarum and P. acidilactici alone decreased the mycotoxin contamina­
behave similarly (Khosroshahi et al., 2022). In summary, the sourdough tion of whole wheat flour and rye by 10%–40% (Pontonio et al., 2021).
fermentation abolished the technological obstacles to incorporating Lv. brevis was capable of decreasing the contamination by aflatoxins B1,
wheat germ into the bread formula. Improved in-vitro protein di­ B2, G1, and G2 (Sadeghi et al., 2019), and P. acidilactici decontaminated
gestibility, decreased phytase activity, and extended shelf life were the deoxynivalenol (44–69%), and removed 15-acetyldeoxynivalenol,
main nutritional benefits derived from the germ supplementation into alternariol, deoxynivalenol-3-glucoside, and toxins H-2 and HT-2
wheat bread formula (Rizzello, Nionelli, Coda, Di Cagno, & Gobbetti, (Zadeike et al., 2021). Usually, the highest level of decontamination
2010). The sourdough fermentation also decreased the content of was observed during long-time (48 h) sourdough fermentation with
anti-nutritional factors such as phytic acid and raffinose and raised the mixed sourdough starters (Zadeike et al., 2021). Under solid-state
content of free amino acids to 50%, mainly lysine (Lys) and γ-amino fermentation of wheat bran, Lactobacillus helveticus synthesized

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E. Pontonio et al. Trends in Food Science & Technology 150 (2024) 104589

Fig. 2. Main advantages of fermentation on brewers’ spent grains and wasted bread.

enzymes capable of degrading aflatoxin B1 into four low-toxin de­ an ingredient for wheat bread formula. Compared to control bread
rivatives without a lactone ring structure (Zhang et al., 2021). without addition, the bread enriched with BSG had a higher content of
proteins and fibers, and positively influenced the gut microbiota func­
1.2. Brewers’ spent grain tionality. Along with an increase of several essential amino acids, a
consistent increase of GABA was also observed during the simulated
Malting for making beer leaves brewers’ spent grains (BSG), whose digestion (Koirala et al., 2022). BSG was also used for making a beverage
fermented by Lp. plantarum. The beverage had antioxidant potential
elevated organic load is critical for environmental pollution (Teixeira
et al., 2020). The recycling of BSG has gained attention due to biogenic because of the high content of phenols and flavonoids, which were
released through acidification and microbial metabolism (Gupta et al.,
components such as alkaloids, plant growth factors, food-grade pig­
ments, and phenolic compounds (Verni, Pontonio, et al., 2020). BSG is 2013). The combination of xylanase and sourdough fermentation with
Lp. plantarum released free phenolic compounds and liberated encrypted
also rich in cellulose and arabinoxylans and contains proteins up to 20%,
particularly rich in lysine (Verni, Pontonio, et al., 2020). The high biogenic peptides with antioxidant activity and protective effect toward
oxidative stress on human keratinocytes (Verni, Pontonio, et al., 2020).
moisture content (circa 80%), along with the richness in polysaccharides
and proteins, make BSG highly prone to microbial contamination with a Given these attributes, the bioprocessed BSG was used for making pasta.
Compared to pasta containing nonfermented BSG, the enriched pasta
short lifespan (7–10 days) (Kavalopoulos et al., 2021). Various conser­
vative treatments of BSG have been assessed (Lynch et al., 2016), and had higher protective effects against oxidative stress caused by human
among these, sourdough fermentation was one of the most promising colon carcinoma cells under simulated gastro-intestinal digestion
(Fig. 2). (Schettino et al., 2021). Positive effects of fermented BSG were also
The sourdough fermentation with lactic acid bacteria enriched BSG observed when used as an ingredient in bread formula. The rheology and
with dextran and oligosaccharides (Koirala et al., 2021). The supple­ sensory attributes of the enriched bread improved, and the nutritional
mentation with sucrose (4% w/w) and fermentation with Leuconostoc profile was optimal because of the supplementation with proteins, fi­
pseudomesenteroides and W. confusa yielded 11 g/kg of dextran, which bers, and essential amino acids (Waters et al., 2012). The enrichment of
improved the BSG viscosity and taste. This fermented BSG was used as bread with spontaneously sourdough-fermented BSG also decreased the

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E. Pontonio et al. Trends in Food Science & Technology 150 (2024) 104589

phytate content (Ktenioudaki et al., 2015). technological features, and creating added functional value along the
BSG was also used as a solid-state substrate to cultivate and preserve food chain.
(up to 10 weeks at circa 6 Log cfu/g) sourdough cultures. The fermented When wasted bread is not edible and recycling into the food chain
BSG was used (10% p/p) in an experimental baking trial leading to implies microbiological risks (e.g., presence of mycotoxins), alternative
sourdough breads characterized by optimal organic acids ratio and biotechnological solutions are exploited. It is known that lactic acid
acceptability according to a consumer-based study (Vriesekoop et al., bacteria act as plant growth-promoting microorganisms (PGPM), indi­
2021). rectly favoring nutrient acquisition in the alkaline soils characterizing
the Mediterranean area. Besides, lactic acid bacteria act as biocontrol
1.3. Wasted bread agents, improving the ability of the host plant to withstand biotic and
abiotic stress or producing compounds that directly stimulate plant
On a global scale, bread is one of the major wasted foods. Industrial growth (Lamont et al., 2017). A bioprocessed wasted bread, which was
waste is generated because of substandard products, crust removal for obtained by enzyme treatments coupled with sourdough fermentation
sandwich production, and unsold bread at retail (Verni, Rizzello, & was used as an amendment in pot trials. The aim was to evaluate the
Coda, 2019). In most cases, the huge amount of wasted bread is still modification of the soil physicochemical properties and the plant
under safe conditions for human consumption. Indeed, innovative, and growth-promoting activity using escarole (Cichorium endivia var. Cuar­
sustainable recycling solutions must focus on keeping wasted bread - tana) as the indicator crop (Cacace et al., 2022). Compared to non­
within the food chain. Recycling wasted bread through sourdough amended soils, the supplementation with sourdough fermented wasted
fermentation has shown an undoubted potential. bread raised the content of soil organic carbon up to 37% and total ni­
Recycling of wasted bread to prepare a culture medium (wasted trogen up to 40%. In addition, the lower pH and the higher organic acid
bread medium, WBM) for the cultivation of lactic acid bacteria was content favored a higher availability of Mn, Fe, and Cu. Escaroles
considered (Verni, Minisci, et al., 2020). The protocol included bread showed improved growth with a higher number of leaves.
(20%) homogenization with water and hydrolysis with amylases. Its Recently, a biotechnological protocol for recycling bread waste at
suitability as a valid alternative to common synthetic media was the industrial level has been set up and patented (Ampollini et al., 2023).
demonstrated for almost all sourdough lactic acid bacteria reaching the The approach includes the preliminary drying and fine milling of bread
most common cell densities after cultivation. The sourdough was also resulting from sandwich production (bread crust). A slurry was obtained
proposed to ferment wasted bread and recycle it as an ingredient for with waste bread powder and water, and food-grade proteases were
bread making. Sourdough was made from wasted bread crumbs. Several added to the semiliquid formulation to promote fermentation. Lactic
lactic acid bacteria demonstrated the capability of using the bread acid bacteria (Lp. plantarum) and yeast (S. cerevisiae) strains were
crumb as a fermentation substrate. Lp. plantarum and P. acidilactici were selected for sourdough fermentation in automatic fermenters. It was
capable of a long-time (48–96 h) fermentation without the genesis of demonstrated that waste bread sourdough can be added up to 50% of the
off-flavors (Gélinas et al., 1999). A sourdough made with wasted bread final weight of bread dough without negatively affecting the textural
was used as an aroma and flavor enhancer for making bread. The for­ properties of the bread. Sensory properties were moreover those typical
mula included 50% whole wheat bread crumb and fermentation with Lp. of a sourdough bread.
plantarum. The synthesis of lactic and acetic acids resembled that of the
traditional sourdough fermentation, and a positive effect on glycemic 1.4. Miscellaneous plant wastes
index and insulinemic responses was hypothesized (Poutanen et al.,
2009). Bread is mainly subjected to fungal spoilage during its shelf life. Miscellaneous plant by-products and wastes comprise a very het­
Salts of propionic and sorbic acids and ethanol are the most common erogeneous set of matrices (e.g., fruit and vegetables hulling and peeling
chemical preservatives to extend the shelf life, although their negative wastes, pomaces, food surplus) sharing an unquestionable nutritional
consumer perception and the healthy recommendations to markedly value. Nevertheless, they are characterized by sources heterogeneity,
decrease their use. A protocol to synthesize bioactive peptides from variation of their chemical composition, and microbial instability. Being
wasted bread was standardized (Nionelli et al., 2020). After hydrolysis rich in carbohydrates, organic acids, mineral salts, and vitamins, these
by microbial proteases and fermentation with Lv. brevis, the wasted compounds can be utilized as substrates for contaminant (either path­
bread showed a broad inhibitory spectrum against some of the most ogenic or not) microorganisms. Furthermore, the high fiber and
diffuse fungal-contaminating species. Nine antifungal peptides, phenolic contents can negatively affect the sensory profile and techno­
encrypted in wheat protein sequences, were released during sourdough logical properties of fortified foods (Kosseva, 2020). The exploitation of
fermentation and identified as the main responsible for fungal inhibi­ their potential needs to overcome these weaknesses, either through
tion. The shelf-life of the bread was extended up to 10 days (Nionelli technological treatments or exploiting fermentation processes, with the
et al., 2020). The direct recycling of wasted bread negatively affects the latter proven to be an effective, sustainable, and mild approach.
specific volume and softness of the newly manufactured bread. Gelati­ Apples are among the most widely processed fruits, which inevitably
nized starch and denatured proteins are not reassembled into a new generate large amounts of residues. Such availability together with high
gluten network (Immonen et al., 2020). The hydrolysis of gelatinized contents of fibers and phytochemicals make these by-products and
starch improved the viscoelastic properties of wasted bread (Immonen wastes ideal ingredients to enrich the wheat bread formula. Fermenta­
et al., 2021), but the most suitable option is the use of sourdough lactic tion of apple by-products with a binary culture of Weissella cibaria and
acid bacteria capable of synthesizing EPS (dextran and β-glucan) in-situ S. cerevisiae was proposed for recycling in breadmaking at a rate of
(Immonen et al., 2020). The dextran synthesis occurring during sour­ 5–10% (w/w of flour) (Cantatore et al., 2019). The fruit and vegetable
dough fermentation of wasted bread compensated for the adverse effect by-products addition in baked goods causes low acceptability because of
of the recycled bread increasing the specific volume and decreasing the the poor sensory and rheological attributes. The dilution of gluten with
crumb hardness. EPS synthesized by Weissella species also exerted fruit or vegetable by-products implies a decreased capability of the
antioxidant and prebiotic activities (Kavitate et al., 2020). A blend of dough to retain gas, which leads to low volume and high hardness.
wasted bread and wheat bran was used as the substrate to synthesize Fortification with fruit or vegetable by-products does not exceed 10%
GABA (Verni et al., 2022). Lp plantarum yielded the highest content of (w/w of flour) (Gómez & Martinez, 2018). The formula enriched with
GABA (circa 800 mg/kg). This fermented wasted bread/bran slurry was 5% apple by-products fermented with sourdough improved the wheat
used as an ingredient for the manufacture of GABA-enriched bread. In dough stability and water absorption and did not interfere with the
conclusion, tailored sourdough fermentations with selected lactic acid bread specific volume and color (Cantatore et al., 2019). The sourdough
bacteria are key options for recycling wasted bread, gaining suitable fermentation maximized the hydration properties of apple by-products

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E. Pontonio et al. Trends in Food Science & Technology 150 (2024) 104589

Fig. 3. Schematic reconstruction of putative pathways characterizing the fermentation (30 ◦ C for 24 h) of a brewer spent grain -based medium by Leuconostoc
pseudomesenterides and Lactiplantibacillus plantarum strains. Red colored pathways were unique to Leuc. pseudomesenteroides, while blue colored pathways were unique
to Lp. plantarum. Black colored pathways were present in both species. Genes that were over-expressed compared to growth in standard reference medium (MRS
broth) are shown in green (Acin-Albiac et al., 2022). (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of
this article.)

because of the synthesis of EPS, and the partial breakdown of insoluble microbial enzymes (e.g., feruloyl esterase, xylanase) with appreciable
fibers, which resulted in an increased porosity and surface area. properties (dos Santos Costa et al., 2021). Date seed flour was proposed
Enrichment with sourdough-fermented apple by-products also provided as an innovative ingredient for sourdough breadmaking (Ameur et al.,
bio-accessible polyphenols (Tlais et al., 2021). In addition, apple 2022). Autochthonous lactic acid bacteria and yeasts were isolated from
pomace was used to fortify sourdough started with a selected culture of date seeds and selected based on various technological criteria. A mixed
Fructilactobacillus florum DSM 22689 and baker’s yeast (single and starter, comprising Leuconostoc mesenteroides, Lp. plantarum and
co-culture). The sourdough nutritional value was positively affected by S. cerevisiae, was used to prepare type I sourdough after consecutive
the incorporation mainly thanks to the presence of valuable compounds refreshments. An aliquot of durum wheat flour was replaced by date
like organic acids and fibers. Additionally, the abundance of glucose and seed flour. The sourdough fermented bread showed an increased radical
fructose in apple pomace led to higher microbial viability and growth scavenging activity because of the consistent release of free phenolic
(Martău et al., 2021). Research efforts also focused on the potential of compounds, while the perceived bitterness and astringency diminished
plant by-products as sources of antimicrobial compounds to extend the due to microbial degradation of tannins. Being an economic and sus­
bread shelf life. By-products having relevant protein contents represent tainable alternative to animal-based proteins, rapeseed proteins isolated
ideal substrates for the biosynthesis of antimicrobial peptides. The from defatted rapeseed press cake have the potential for food applica­
sourdough fermentation of palm kernel cake with Lc. casei favored the tions. Rapeseed protein concentrations higher than 5% in wheat
accumulation of antifungal peptides, which, once added to the bread breadmaking led to detrimental effects on textural and sensory attri­
formula, were effective in counteracting the spoilage up for to 10 days butes. Rapeseed proteins became suitable ingredients for making sour­
(Asri et al., 2020). Another constraint to recycle non-conventional in­ dough bread when fermented with W. confusa, which synthesized
gredients is their eventual association with increased levels of acryl­ dextran in-situ. Compared to control bread, the sourdough fermentation
amide during baking (Bartkiene, Bartkevics, et al., 2021). Oat has a improved the in-vitro protein digestibility, enriched the free amino acid
higher potential for acrylamide formation than wheat because of the profile mainly with lysine, methionine, and isoleucine, and guaranteed
higher levels of soluble carbohydrates and free asparagine. Sourdough suitable specific volume and low crumb hardness (Wang, Li, et al.,
fermentation of oat by-products prevented acrylamide accumulation in 2022a,b).
baked goods (Bartkiene, Bartkevics, et al., 2021). Solid-state fermenta­
tion and the subsequent recycling as ingredients for breadmaking 1.4.1. Lactic acid bacteria glycosyl hydrolases
demonstrated how agri-food by-products also served to deliver One of the main secrets beyond the sourdough fermentation as

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E. Pontonio et al. Trends in Food Science & Technology 150 (2024) 104589

valuable biotechnology to recycle plant by-products and wastes con­ Writing- Reviewing and Editing, Supervision. Alessandro Stringari:
cerns the hydrolysis of complex polymers (e.g. arabinoxylans, cellulose, Data curation, Investigation. Raffaella Di Cagno: Writing- Original
and hemicellulose fraction) and the bio-accessibility of phenolic com­ draft preparation, Investigation. Pasquale Filannino: Writing- Original
pounds. Sourdough lactic acid bacteria have the aptitude to metabolize draft preparation, Investigation. Carlo Giuseppe Rizzello: Data cura­
diversified carbon sources. This aptitude relies on key metabolic path­ tion, Investigation, Writing- Original draft preparation. Andrea Polo:
ways mediated by an extensive set of glycosyl hydrolases. These en­ Original draft preparation. Olga Nikoloudaki: Data curation, Investi­
zymes undertake the ambivalent role of releasing a wide range of gation. Marco Gobbetti: Conceptualization, Methodology, Writing-
phenolic compounds from glycosylated precursors and degrading the Original draft preparation, Supervision.
resulting sugar moiety. Glycosyl hydrolases also have the potential to
remove undesired bitterness or release aroma compounds. Encoding
genes for glycosyl hydrolases are widespread among lactic acid bacteria. Declaration of competing interest
Phospho-β-glucosidase genes show a high degree of redundancy. Phos­
pho-β-glucosidases catalyze the degradation of phosphorylated gluco­ The authors report there are no competing interests to declare.
sides and fibers-related disaccharides (e.g., cellobiose and gentiobiose),
which are activated through the phosphoenolpyruvate (PEP)-dependent Data availability
carbohydrate phosphotransferase systems (PEP-PTS) (Acin-Albiac et al.,
2021). Recently, the complete framework describing the metabolism No data was used for the research described in the article.
drift of Lp. plantarum and Leuc. pseudomesenteroides caused by the
lignocellulosic BSG was provided by implementing molecular and phe­ Acknowledgements
nomics approaches (Fig. 3) (Acin-Albiac et al., 2021, 2022). As
confirmed by gene overexpression, Lp. plantarum preferred arabinose The authors thank Lena Brigitte Marie Granehäll for language
among pentosans, while Leuc. pseudomesenteroides mainly uses xylose. proofreading.
The phenotype switching towards galactose metabolism suffered the
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