Food Research International 104 (2018) 86–99

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Food Research International

journal homepage: www.elsevier.com/locate/foodres

Fruits and vegetables, as a source of nutritional compounds and T

phytochemicals: Changes in bioactive compounds during lactic fermentation
Axelle Septembre-Malaterreb,⁎, Fabienne Remizeb, Patrick Pouchereta
a
Laboratoire de Pharmacologie et Physiopathologie Expérimentale, UMR 95 Qualisud, Université de Montpellier, France
b
UMR QualiSud, Université de La Réunion, CIRAD, Université Montpellier, Montpellier SupAgro, Université d'Avignon et des Pays de Vaucluse, Sainte Clotilde, France

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

Keywords: Nutritional recommendations emphasize fruit and vegetable consumption. This is related to health-benefits
Fruits associated with bioactive nutritive molecules (nutrients, vitamins, minerals, fibers, …) as well as non-nutritive
Vegetables phytochemicals (phenolic compounds, flavonoids, bioactive peptides, …) content of these types of foods.
Phytonutrients Fermented fruit or vegetable products are part of various diets worldwide. Lactic acid bacteria (LAB) fermen-
Polyphenols
tation is common among the various fermentation processes used. It produces changes in both profile and types
Lactic fermentation
of bioactive compounds. Molecules such as bioactive peptides, short chain fatty acids or polysaccharides are
Health benefits
generated while sugar content or anti-nutritional compounds are decreased and phenolic compounds are con-
verted to molecules with added biological value. These transformations, associated with prebiotic and/or pro-
biotic potential supply as well as improvement of food components bioaccessibility and bioavailability, result in
modifications of health-related properties. Lastly, interactions between ingested fermented food, intestinal mi-
crobiota and their correlations to metabolomics profiles and health represent an important perspective deserving
to be further explored.

1. Introduction
Bioactive compounds in foods are numerous and chemically diverse
(Kris-Etherton et al., 2002; Rickman, Barrett, & Bruhn, 2007; Rickman,
Bruhn, & Barrett, 2007). In fruits and vegetables, molecules of nutri-
tional interest are fibers, vitamins, minerals, phenolic compounds in-
cluding flavonoids, phytoestrogens, sulfur compounds, monoterpenes
and bioactive peptides. Their level in fruits and vegetables is a function
of varieties, maturity and agronomical practices such as nitrogen ad-
dition and irrigation. Nutrients are defined as molecules required for
organism survival and growth. Dietary Reference Intakes (DRIs) are
defined for nutrients. Fruits and vegetables are good sources of fibers,
minerals especially potassium, and vitamins, mainly vitamins C and K
(Pennington & Fisher, 2010). More precisely, dark green leafy vege-
tables like spinach and cabbages provide at least 50% DRI for vitamins
C and K. Legumes and Allium family bulbs provide 50% DRI for vitamin
K while tomatoes and other red fruits provide 50% DRI for vitamin C.
More than 25% DRI for folate is found in dark green leafy vegetables
and legumes and for manganese and vitamin B6 in Allium family bulbs.
All fruits contribute significantly to vitamin C intake, but citrus fruits
may provide 40% of DRI. Composition tables particularly focus on
nutrients and every food may be described as “good source of” or
“contributing to” for each nutrient. These data may be found elsewhere
(Pennington & Fisher, 2010). Apart from nutrients, phytochemicals are
defined as the bioactive non-nutrient plant compounds in fruits, vege-
tables, grains and other plant food. They gather > 5000 chemicals.
In Western countries, fermented foods were the object of a renewed
interest (Ebner, Smug, Kneifel, Salminen, & Sanders, 2014). They are
associated with: (1) consumers concern for nutrition-health approach of
individual diet (vitamins, pre-biotic, pro-biotic, digestibility), (2) food
safety, as the process is natural and limit biohazardous contaminants,
(3) organoleptic modification of food with new aroma, texture or taste,
(4) shelf life extension in particular for perishable foods, (5) simplicity
of preparation since process is self-conducted with a limited number of
unitary operation, (6) valorization of unused raw vegetal material left
over by agricultural and food industries and (7) the process is cheap and
energetically compatible with sustainable development (Marco et al.,
2017; Montet, Loiseau, & Zakhia-Rozis, 2006; Motarjemi, 2002).
This is the consequence of the food transition that occurred in these
countries and currently arises in low and middle income countries.
Food transition toward a “Western” type of diet is characterized by an
increased intake of calories, sugar, saturated fat and animal proteins,
and a decreased consumption of fruits, vegetables and fibers. This
change of the dietary pattern is associated with a growing physical

⁎
Corresponding author.
E-mail address: axelle.malaterre-septembre@univ-reunion.fr (A. Septembre-Malaterre).

http://dx.doi.org/10.1016/j.foodres.2017.09.031
Received 29 April 2017; Received in revised form 27 August 2017; Accepted 9 September 2017
Available online 14 September 2017
0963-9969/ © 2017 Elsevier Ltd. All rights reserved.
A. Septembre-Malaterre et al. Food Research International 104 (2018) 86–99

inactivity. Together these factors promote higher prevalence and in- their antioxidant, hypoglycemic, hypolipidemic and anti-inflammatory
cidence of non-communicable degenerative diseases such as obesity, properties (Andriantsitohaina et al., 2012; Coban et al., 2012;
diabetes, cardiovascular pathologies, … (Popkin, 2015). These chronic Scalbert & Williamson, 2000; Tresserra-Rimbau et al., 2014).
illnesses seem to be partially correlated with induction of unbalanced
digestive system microbiota that favor diseases (Ramchandran, 2015). 2.2. Bioactive nutritive molecules
Fermented foods would represent a mean to decrease the risk or se-
verity of Western lifestyle associated diseases through improved nu- Fruits and vegetables are consumed as sources of water, carbohy-
tritional and functional values (Hasler, 2000, 2002; Stanton, Ross, drates and essential nutrients such as vitamins, minerals and fibers
Fitzgerald, & Sinderen, 2005). (Costa, Garcia-Diaz, Jimenez, & Silva, 2013).
This review will particularly focus on the fruits and vegetables Dietary fibers whether soluble of insoluble are a group of polymers
phytochemical, as bioactive non-nutrient compounds, their production with heterogeneous structures (polysaccharides, lignin). Fruits and ve-
through the fermentation process and their potential interest for human getables content in fibers usually ranges from 1 to 5.4 g per 100 g of
health. fresh weight. In consequence, fruits and vegetables are effective on
physiological parameters such as satiety, gastrointestinal tract phy-
2. Nutritional compounds and phytochemicals in fruits and siology, metabolic parameters (post-prandial lipemic response, long
vegetables term basal lipemia) and microorganism local population through pre-
biotic effects (Eswaran, Muir, & Chey, 2013). In addition, consumption
2.1. Fruit and vegetable composition and diet of fruits and vegetables helps covering fiber requirements with con-
tained energy intake when compared to refined foods (Chang,
Fruits and vegetables rich diet is well established for its efficiency to Alasalvar, & Shahidi, 2016).
promote human health, in particular to regulate the body weight Carbohydrate intake from vegetables varies from 1 to 9% (from
(Estruch et al., 2013; Mozaffarian, 2016). Consumption of fruits and aerial parts such as salads, to roots such as carrots, respectively)
vegetables is considered by many organizations (World Health Orga- whereas for fruits it varies from 5 to 22% of fresh weight (citrus or
nization—WHO, Food and Agriculture Organization— FAO, United strawberry to grapes or banana respectively). Major nutritional com-
States Department of Agriculture—USDA and European Food Safety positions of commonly consumed fruits and vegetables are listed in
Authority—EFSA) as a major public health issue and is the subject of Table 1 (US Department of Agriculture, Agricultural Research Service,
nutritional recommendations worldwide (Ragaert, Verbeke, 2016).
Devlieghere, & Debevere, 2004; Su & Arab, 2006). In fact, the Dietary
Guidelines for Americans published in 2016 advises that half our meal Table 1
should be composed of fruits and vegetables. Many health policies insist Major nutritional components of commonly used fruits and vegetables.
on increasing consumption of fruits and vegetables due to the resulting
Fruits and vegetables Fiber Carbohydrate Energy
health benefits (Djuric, 2011; Tiwari & Cummins, 2013).
Foods of vegetable origin are characterized by low caloric intake g/100 g fresh weight kcal
(due to their high water and low fat content). It is mostly composed of
high content of carbohydrates, fibers, minerals and micronutrients of Artichoke 5.40 10.51 47.00
Asparagus 2.10 3.88 20.00
interest, including carotenoids, vitamins, minerals and polyphenols
Broccoli 2.60 6.64 34.00
(Yahia, 2009). These bioactive compounds are considered as molecules Cabbage 2.50 5.80 25.00
with therapeutic potential that can exert actions on energy intake, Carrot 2.80 9.58 41.00
while decreasing excessive oxidative stress, pro-inflammatory state and Cauliflower 2.00 4.97 25.00
metabolic disorders (Siriwardhana et al., 2013). Indeed, a diet enriched Celery 1.60 2.97 16.00
Eggplant 3.00 5.88 25.00
in fruits and vegetables has a positive impact on several chronic con- Garlic 2.10 33.06 149
ditions, such as obesity, diabetes, cancer, cardiovascular and neurode- Leek 1,8 14,15 61
generative diseases (Leite et al., 2011). Lettuce 1,3 2,87 15
A growing interest is brought to the antioxidant micronutrients Onion 1,7 9,34 40
Potato 2,5 12,44 58
owing to the diverse biological properties that they would be able to
Pumpkin 0,5 6,5 26
exert. For decades, these compounds have been correlated to a risk Radish 1,6 3,4 16
reduction of chronic diseases such as cardiovascular disease, cancer, Spinach 2.20 3.63 23.00
diabetes, Alzheimer disease, cataracts and age-related functional de- Tomato 1,2 3,89 18
cline. Human cells are constantly exposed to oxidizing agents and a key- Zucchini 1,1 3,11 21
Apple 1,3 12,76 48
point is to balance this oxidative effect by antioxidant mechanisms. Apricot 2 11,12 48
Antioxidants in fruits and vegetables are mainly phytochemicals, and Banana 2,6 22,84 89
more specifically phenolic compounds and carotenoids. At 1 g/day, Cherry 2,1 16,01 63
polyphenols are the most abundant antioxidants in the human diet Clementine 1,7 12,02 47
Fig 2,9 19,18 74
compared to vitamin C (90 mg daily), vitamin E (12 mg daily), and
Grapes 0,9 18,1 69
vitamin A and its precursors carotenoids (5 mg per day) Litchis 1,3 16,53 66
(Scalbert & Williamson, 2000). These compounds constitute a hetero- Mango 1,6 14,98 60
geneous group of compounds. More than 8000 molecules were identi- Melon 0,9 8,16 34
fied and classified into five main classes according to their chemical Orange 2,4 11,75 47
Papaya 1,7 10,82 43
structure: flavonoids, phenolic acids, stilbenes, lignans, and curcumi- Peach 1,5 9,54 39
noids (Manach, Scalbert, Morand, Rémésy, & Jiménez, 2004). The Pear 3,1 15,23 57
considerable diversity of their structures put polyphenols apart from Pineapple 1,4 13,12 50
other antioxidants. Their bioavailability and biologic properties vary to Plum 1,4 11,42 46
Strawberry 2 7,68 32
a great extent and are affected by their chemical structure (Manach,
Watermelon 0,4 7,55 30
Williamson, Morand, Scalbert, & Rémésy, 2005). For several years
much interest has been attributed to plant products with high poly- Ref: USDA. (2017). National nutrient database for standard reference.
phenols content due to their extensive biological potential, including Values correspond to those of fruits and vegetables raw.

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A. Septembre-Malaterre et al. Food Research International 104 (2018) 86–99

Table 2
Macro and micronutrients composition of commonly used fruits and vegetables.

Fruits and vegetables Macronutrients (mg/100 g of fresh weight) Micronutrients (μg/100 g of fresh weight)

Vitamin C Phosphorous Potassium Magnesium Sodium Calcium Vitamin A Iron Selenium Copper Manganese Zinc

Artichoke 11,7 90 370 60 94 44 1 1280 0,2 230 260 490

Asparagus 5,6 52 202 14 2 24 38 2140 2,3 190 160 540
Broccoli 89,2 66 316 21 33 47 31 730 2,5 50 210 410
Cabbage 36,6 26 170 12 18 40 5 470 0,3 20 160 180
Carrot 5,9 35 320 12 69 33 835 300 0,1 40 140 240
Cauliflower 48,2 44 299 15 30 22 0 420 0,6 40 150 270
Celery 3,1 24 260 11 80 40 22 200 0,4 40 100 130
Eggplant 2,2 24 229 14 2 9 1 230 0,3 80 230 160
Garlic 31,2 153 401 25 17 181 0 1700 14,2 300 1670 1160
Leek 12 35 180 28 20 59 83 2100 1 – 480 120
Lettuce 3,7 28 187 3 28 36 370 860 0,6 30 250 180
Onion 7,4 29 146 10 4 23 0 210 0,5 40 130 170
Potato 11,4 38 413 21 16 9 0 730 0,5 120 140 290
Pumpkin 9 44 340 12 1 21 426 800 0,3 130 120 320
Radish 14,8 20 233 10 39 25 0 340 0,6 50 70 280
Spinach 28,1 49 558 79 – 99 469 2710 1 130 900 530
Tomato 16 29 212 8 42 5 75 470 0,4 60 90 140
Zucchini 34,1 93 459 33 3 21 25 790 0,3 100 200 830
Apple 4,6 11 107 5 1 6 3 120 0 30 40 40
Apricot 10 23 259 10 1 13 96 390 0,1 80 80 200
Banana 8,7 22 358 27 1 5 3 260 1 80 270 150
Cherry 10 15 173 11 0 13 3 360 0 60 – 70
Clementine 48,8 21 177 10 1 30 – 140 10 40 20 60
Fig 2 14 232 17 1 35 7 370 0,2 70 130 150
Grapes 3,2 20 191 7 2 10 3 360 0,1 130 70 70
Litchis 71,5 31 171 10 1 5 0 310 0,6 150 60 70
Mango 36,4 14 168 10 1 11 54 160 0,6 110 60 90
Melon 36,7 15 267 12 16 9 169 210 0,4 40 – 180
Orange 53,2 14 181 10 0 40 11 100 0,5 40 30 70
Papaya 60,9 10 182 21 8 20 47 250 0,6 40 40 80
Peach 6,6 20 190 9 0 6 16 250 0,1 70 60 170
Pear 4,3 12 116 7 1 9 1 180 0,1 80 50 100
Pineapple 47,8 8 109 12 1 13 3 290 0,1 110 930 120
Plum 9,5 16 157 7 0 6 17 170 0 60 50 100
Strawberry 58,8 24 153 13 1 16 1 410 0,4 50 390 140
Watermelon 8,1 11 112 10 1 7 28 240 0,4 40 40 100

Ref: USDA. (2017). National nutrient database for standard reference.

Values correspond to fruits and vegetables raw.

Fruits and vegetables are a significant source of vitamins, minerals
and antioxidants. They may contain substantial amounts of potassium,
calcium (citrus, cabbages), magnesium, iron (leafy vegetables), copper
(leafy vegetables) and sulfur (cabbage, onions, garlic, leeks, turnips,
radishes). Fruits are usually low in sodium and contain trace minerals.
Fruits and vegetables are also rich in water-soluble vitamins such as
vitamin C (cabbages, leafy vegetables, tomatoes, citrus fruits, currants,
blackcurrants, strawberries …), vitamins of the B group and car-
otenoids, precursors of vitamin A. Micro and macronutrients composi-
tion of commonly consumed fruits and vegetables is listed in Table 2
(US Department of Agriculture, Agricultural Research Service, 2016).
2.3. Phytochemicals: non-nutritive
2.3.1. Diversity of compounds
Phytochemicals are secondary metabolites synthetized by plants,
also known as bioactive compounds, (Yao et al., 2004). They can be
described as chemicals compounds that may affect health, but are not
essential nutrients (Temple, 2000). Phytochemicals from vegetables
and fruits are used against free radicals and oxidative damage asso-
ciated with chronic diseases (Tiwari & Cummins, 2013). They include
several families of molecules, such as glucosinolates, flavonols, iso-
flavones, phenolic acids, flavones, phytoestrogens and carotenoids.
In general, these compounds demonstrate properties such as anti-
oxidant (D.-O. Kim, Padilla-Zakour, & Griffiths, 2006), anti-in-
flammatory (Vincent, Bourguignon, & Taylor, 2010), lipid profile al-
teration (Wang, Melnyk, Tsao, & Marcone, 2011) and antitumor effects
(Stan, Kar, Stoner, & Singh, 2008). Besides beneficial properties for
human health, phytochemicals are responsible for color, flavor and
odor (Miglio, Chiavaro, Visconti, Fogliano, & Pellegrini, 2008). Their
content is influenced by crop type, variety, environmental conditions,
location, germination, maturity, processing and storage (Björkman
et al., 2011; Carbone, Giannini, Picchi, Lo Scalzo, & Cecchini, 2011).
Phenolic compounds are one of the main class of plant secondary
metabolites and among the most abundant natural antioxidants in the
diet. Polyphenols would be involved in the prevention of various
pathologies associated with oxidative stress, such as cancer, neurode-
generative or cardiovascular diseases (Robards et al., 1997; Yao et al.,
2004). These health benefits are based on their structure characterized
by one or more phenolic groups bearing at least one hydroxyl group
(OH) (Hennebelle, Sahpaz, & Bailleul, 2004). Phenolic potential of
fruits depends on many external factors (climatic conditions, technical
itineraries, origins) and internal factors (physiological state of the fruit,
position of the fruit on the tree, genotype) (Dragovic-Uzelac, Levaj,
Mrkic, Bursac, & Boras, 2007; Renard, Dupont, & Guillermin, 2007)
(Renard et al., 2007). According to their structure, polyphenols can be
classified into 5 major families: phenolic acids, flavonoids, lignans,
stilbenes and curcuminoids (Panickar & Anderson, 2011). The main
food sources are fruits, vegetables, derived beverages (wine, tea, coffee,
and fruit juices), cereals, oilseeds and pulses. Fruits and vegetables
contribute to about one half of the total nutritional intake of poly-
phenols, the other half being provided by derived beverages (Brat et al.,
2006). In most cases, foods contain complex mixtures of polyphenols.

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A. Septembre-Malaterre et al.
2.3.2. Phenolic acids
Phenolic acids are classified in two classes of compounds: benzoic
acid derivatives (or hydroxybenzoic acids) and cinnamic acid deriva-
tives (or hydroxycinnamic acids).
Derivatives of benzoic acid are present in fruits and vegetables
mostly in the conjugated form (esters or glycosides) but can also be
found in the free form. Their content in edible plants is generally very
low, with the exception of some red fruits, black radishes and aromatic
plants (e.g., protocatechic acid in raspberries) (Cabrini et al., 2001).
Hydroxybenzoic acids can be esterified with glucose molecule to give
more complex structures called hydrolyzable tannins
(Clifford & Scalbert, 2000), such as gallotannins found in mangoes and
ellagitannins in berries (Clifford & Scalbert, 2000).
Hydroxycinnamic acids are extensively found in caffeic acid form
(D'Archivio et al., 2007) and are abundant in some vegetables such as
eggplant (D'Archivio et al., 2007; Nicholson, Tucker, & Brameld, 2008).
Para-coumaric acid is found in spinach (Clifford, 2000b). Hydro-
xycinnamic acids are rarely present in free form and are found mainly
in conjugated forms (glycosylated or esters) (Manach et al., 2004). The
most common conjugate is chlorogenic acid, which is found in many
fruits and in potatoes (Clifford, 2000b).
2.3.3. Flavonoids
Flavonoids are the most abundant polyphenols in the diet and >
4000 compounds have been identified (D'Archivio et al., 2007). Fla-
vonoids are found throughout the plant kingdom, especially in medic-
inal and aromatic plants, fruits and vegetables (Verhoeyen et al., 2002).
They may be subdivided into eight subclasses (Crozier,
Jaganath, & Clifford, 2009): flavones, Isoflavones, flavonols, flava-
nones, flavanols, proanthocyanidins, anthocyanins, chalcones/dihy-
drochalcones.
Flavones are mainly found in parsley and celery or moderately in
red pepper (Erdman et al., 2007). The glycosides of flavones, in parti-
cular luteolin and apigenin, are present in cereals such as millet and
wheat (Manach et al., 2004).
Isoflavones are contained almost exclusively in legumes and more
particularly in the Fabaceae. Soy and its by-products are the main
sources in human diet (Aedin Cassidy, Hanley, & Lamuela-Raventos,
2000). Three main molecules have been identified in soybean: genis-
tein, daidzein and glycitein (Coward, Smith, Kirk, & Barnes, 1998).
Isoflavone content is highly variable in foods derived from soybeans but
is found in large quantities in soy-based fermented products (Aedin
Cassidy et al., 2000).
Flavonols are the most common flavonoids in the plant kingdom,
with the exception of algae and mushrooms. Their main representatives
are quercetin, kaempferol, myricetin and isorhamnetin. They are gen-
erally present at relatively low concentrations (onion is the richest food
in quercetin). Fruits may contain between 5 and 10 different flavonol
glycosides. The four most frequent flavanones are naringenin, hesper-
etin, eriodictyol and isosakuranetin. They are characteristic of citrus
fruits (lemons, mandarins, grapefruits and oranges). Aglycones such as
naringenin confer a bitter taste in grapefruits, while rutinoids (narir-
utin, hesperidin, etc.) give a less pronounced flavor in oranges and
lemons (Leuzzi, Caristi, Panzera, & Licandro, 2000; Tomas-
Barberan & Clifford, 2000).
Flavanols and proanthocyanidins are the major flavonoids in the
human diet. They exist both in the form of monomers (catechins) and in
the form of polymers (proanthocyanidins). The main catechins present
in the fruits are catechin and epicatechin, while epigallocatechin, epi-
catechin gallate and epigallocatechin gallate are found in some legume
seeds, grapes and especially tea (Arts, van de Putte, & Hollman, 2000a,
2000b). Catechins are present in many fruits such as apricots and
cherries and also in red wine but green tea is the main source
(D'Archivio et al., 2007; Hara, 2011).
Proanthocyanidins are particularly abundant in fruits (apple, plum,
strawberry, …), wine, chocolate and some dried vegetables (common
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Food Research International 104 (2018) 86–99
bean, hazelnut) (Gu et al., 2004). Proanthocyanidins contribute to the
flavor of food. They form molecular complexes with salivary proteins,
are responsible for the bitterness of chocolate and astringency of certain
fruits (grapes, peaches, persimmons, apples, …) as well as some drinks
such as wine, cider, tea or beer (Rasmussen, Frederiksen, Struntze
Krogholm, & Poulsen, 2005; Santos-Buelga & Scalbert, 2000; Tanaka
et al., 1994).
Anthocyanins are found in red wine, varieties of cereals, vegetables
and tubers (eggplant, cabbage, beans, onions, radish), but they are
more abundant in fruits. Their content in food is generally proportional
to the intensity of the color of the product (high value in blackberries).
These values increase as the fruit ripens. Anthocyanidins are mainly
found in the skin of the fruit (Clifford, 2000a; Manach et al., 2004).
Chalcones are poorly present in the diet due to their transformation
into acidic flavanones, mainly during the extraction stages. The most
studied are phloretin and its glycoside, phloridine (phloretin 2′-O-glu-
coside) that are abundant in apples (Scalbert & Williamson, 2000;
Tomas-Barberan & Clifford, 2000).
2.3.4. Lignans
Lignans are present mainly in flax seed, which contains secoisolar-
iciresinol, as well as small amounts of matinesinol
(Adlercreutz & Mazur, 1997). Other cereals (triticale and wheat), fruits
(pears and plums), some vegetables (garlic, asparagus and carrots) and
legumes such as lentils also contain lignans but at 1000 times lower
than in linseed (Adlercreutz & Mazur, 1997). The intestinal microflora
metabolizes the lignans into enterodiol and enterolactone, thus they are
considered phytoestrogens (Heinonen et al., 2001), notably present in
apples (Scalbert & Williamson, 2000). Phytoestrogens present in fruits
and vegetables have attracted much interest due to their potential
protective effects against various disease conditions such as cancer,
cardiovascular disease (CVD), osteoporosis, and menopausal symptoms
(Bradford & Awad, 2007; John, Sorokin, & Thompson, 2007).
2.3.5. Stilbenes
Stilbenes are poorly present in human food and the main re-
presentative is resveratrol, which exists in cis and trans forms.
Trans‑resveratrol is mainly present in the skin of grape (Frémont,
2000). Concentration of trans‑resveratrol in wine is partly determined
by the steps of maceration with the skin and grape seeds (Fernández-
Mar, Mateos, García-Parrilla, Puertas, & Cantos-Villar, 2012; Romero-
Pérez, Ibern-Gómez, Lamuela-Raventós, & de La Torre-Boronat, 1999).
It is also present in red and white grape juice (Romero-Pérez et al.,
1999). In addition, trans‑resveratrol is also found in 72 other plant
species, such as blackberries and peanuts as well as medicinal plants
(Crozier et al., 2009; Hurst et al., 2008).
2.3.6. Curcuminoids
Curcuminoids are the majority compounds of the curcuma rhizome
with a content of 20 to 80 g/kg, responsible for the yellow color of this
spice and numerous biological activities such as antitumor and neuro-
protective effects (Aggarwal, Sundaram, Malani, & Ichikawa, 2007).
2.4. Other molecules with beneficial health effects
Glucosinolates is a large family of molecules composed of a β-D-
thioglucose group, a sulfonated oxime moiety and a variable side-chain
derived from amino acids (methionine, tryptophan or phenylalanine
and some branched chain amino acids). These compounds are present
in Brassica vegetables like cabbages, broccoli, cauliflower, mustard,
garden cress and Brussels sprouts. Glucosinolate biosynthesis is a very
complex process involving three distinct stages: side chain elongation,
nucleus structure formation and side chain changes, thus generating a
large variety of glucosinolates. Glucosinolates are not biologically ac-
tive, only their enzymatic derivatives are. They are inactive as long as
they are located into their sub-cellular compartments. However, the
A. Septembre-Malaterre et al.
endogenous enzyme myrosinase catalyses the hydrolysis of glycosidic
bond releasing glucose (sugar part) and bioactive breakdown com-
pounds (non-sugar part or aglycone). Myrosinase (EC 3.2.3.1) is acti-
vated upon plant tissue disruption, caused by harvest, pests, processing
or consumption (Johnson, 2002; Petropoulos, Gioia, & Ntatsi, 2017).
The biological properties of glucosinolate breakdown products, espe-
cially isothiocyanates, are related to their ability to block the cell cycle
and induce apoptosis in several types of cancer cells, exerting thus
anticarcinogenic effects. Indoles and isothiocyanates are responsible for
chemoprotective effects. Other health-promoting effects of iso-
thiocyanates are related to their antioxidant properties.
2.5. Anti-nutrients
Plants synthesize secondary metabolites to protect themselves from
various attacks (insects, pathogens, unfavorable growing conditions,
…). These compounds may have detrimental effects on consumers. For
example, potatoes contain alkaloid solanine, arsenic and nitrite, green
leafy vegetables presents toxic oxalates and peas contain phytic acid,
protease inhibitors and tannins (Habiba, 2002). Most anti-nutrients can
be reduced or destroyed by appropriate cooking method
(Fabbri & Crosby, 2016). Other compounds called antinutritional fac-
tors may exert specific biological activities depending on their struc-
ture. Legumes are the main sources of anti-nutritional compounds in
human diet. Common legumes include soybeans (Glycine max), black
gram (Phaseolus mungo), cow pea (Vigna unguiculata), dry beans (Pha-
seolus vulgaris), winged beans (Psophocarpus tetragonolobus), chick pea
(Cicer arietinum L.), horse gram (Dolichos biflorus), moth bean (Vigna
aconitifolia), pigeon pea (Cajanus cajan), favabeans (Vicia faba L. minor),
grain amaranth (Amaranthus spp.), lentil (Lens culinaris medic), jack-
bean (Canavalia gladiata) and grass peas (Lathyrus sativus). Legumes
consumed in fresh forms, like green beans, young beans in their pods
and green peas, are considered as vegetables. The anti-nutritional fac-
tors predominantly present in these types of foods are lectins, trypsin
and protease inhibitors, tannins, phytic acids, saponins, phytates, glu-
cosinolates, cyanogens, phytoalexins, etc. Some of these compounds
may reduce the bioavailability of certain compounds or inhibit diges-
tion enzymes.
Hereafter is a summary of the most studied anti-nutritional factors.
Protease inhibitors or enzyme inhibitors (trypsin) form stable
complexes with proteolytic enzymes which reduce/block their activity
in the intestine. Thus their main action is to reduce the digestibility of
proteins. These molecules can be classified into two broad classes:
Kunitz trypsin inhibitors and Bowman-Birk trypsin/chymotrypsin in-
hibitors.
Saponins, which are glycosides of triterpenoids or steroids, are
found mainly in legumes, roots but may also be present in sugar beets,
oats, and tea. They are characterized by a bitter taste, a surfactant ac-
tivity, their ability to haemolyze red blood cells, their effect on the
intestinal epithelium functions, favoring the passage of allergens or
disrupting cell renewal.
The phytic acid (or phytates) consists of an inositol radical esterified
by 6 phosphate radicals. They are present in monocotyledons (wheat,
rice), in dicotyledons (legumes, nuts and oilseeds). Phytates are closely
related to proteins. They demonstrate an influence on the functional
and nutritive properties of foods. Despite a recognized antioxidant ef-
fect, phytates are the main chelating agents of legume seeds. During
digestion, they form insoluble and non-digestible complexes with di-
valent cations such as (Mg2 +, Fe2 +, Zn2 +, Ca2 + …). They are con-
sidered mineral assimilation disturbing agents reducing mineral ele-
ments bioavailability. Phytates also influence the activity of enzymes
such as pepsin, trypsin and amylases. They can also form complexes
with proteins which reduces their solubility and digestibility. Phytates
can also interact with starch either directly through hydrogen bonds
formation with a phosphate group or indirectly via proteins which re-
sults in a decrease in starch solubility and digestibility. Despite their
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anti-nutritional effects, phytates are the only source of phosphorus in
legumes and their concentrations can reach 10% of dry matter (Bora,
2014).
Lectins are proteins that bind specifically and reversibly to some
carbohydrates. They are found in leguminous seeds (lentils, beans,
peas). Lectin ingestion reduces the absorption efficiency of digested
products. Being partially resistant to proteolytic enzymes in vivo, Lectin
action on the intestine is manifested by severe lesions of the brush
border membrane, a reduction of the size of the villi, an abnormal
development of microvilli and an increase in cell turnover, followed by
a significant reduction of the digestion and absorption capacities of the
intestine. Weight loss, nutrient malabsorption, and long-term growth
retardation are the results of ingesting lectin-rich foods.
Oligosaccharides are present at high levels in legume seeds (up to
20%). The main compounds found in legumes are verbascose, stachyose
and raffinose. Oligosaccharides are not digested by the human or-
ganism since it does not produce the suitable enzyme for their diges-
tion: the alpha-galactosidase. Thus, once consumed, these compounds
are fermented by the digestive flora of the colon, causing gas produc-
tion and flatulence. Their ingestion would have prebiotic effects by
promoting the proliferation of Bifidobacterium and Lactobacillus. These
bacteria help balancing the intestinal flora and preventing some dis-
eases. Positive effects are modulated by intake.
Anti-nutritional factors have negative effects on the nutritional
quality of the foods. As a result, it is necessary to eliminate or inactivate
them before the plant matrix is ingested by man or animals. This can be
carried out either by selection of the plant genotype or through post-
harvest treatment (germination, boiling, leaching, extraction fermen-
tation, etc.). The structure of anti-nutritional factors and their chemical
properties, particularly thermal lability, determine the physical process
that will be most effective in reducing or eliminating them, thereby
minimizing adverse biological effects.
3. Traditional lactic fermented foods from fruits and vegetables
Nutrient and phytochemical content of fruits and vegetables de-
pends on cultivar, agricultural practices and ripening stage as reported
for cabbage (Singh et al., 2006) and olive (Othman, Roblain, Chammen,
Thonart, & Hamdi, 2009). In cabbage, it was shown that some phyto-
chemical concentration can vary up to 10-fold as a function of cultivar.
After harvest, and even more, after cutting, fruits and vegetables are
rapidly spoiled due to the development of yeasts and molds, but also
because of endogenous reactions (like anaerobic catabolism or pecti-
nolytic activity) and mechanical damages (Barth, Hankinson, Zhuang,
Breid, & Breidt, 2009; Toivonen & Brummell, 2008). Food transforma-
tion technologies aim at either increasing shelf-life of products, chan-
ging the taste or both. A consequence is the modification of composi-
tion. For instance, decrease of water content, through drying or thermal
treatments, promotes hydrolysis or polymerization. Among food pro-
cessing technologies, fermentation aims at increasing shelf-life and
developing pleasant sensory characteristics. Fermentation belongs to
some of the most ancient food processes in human history. Since Pa-
leolithic and Neolithic eras, products such as bread, wine, beer and
cheese were developed and consumed by ancient populations world-
wide. Indeed, very soon, humans learned to exploit this endogenous
process that need very limited equipment or energy, and later on to
induce it. This phenomenon is natural and occurs spontaneously if raw
material, whether vegetal or animal, is stored of left without appro-
priate conditions of conservation. Microorganisms naturally present on
the substrate, develop their fermentative activity. This leads to the
transformation of the initial material and to modification of biochem-
ical composition. Fermentation causes considerable changes that affect
the organoleptic properties (taste, texture and in a lesser extend color),
the nutritional value and the microbial safety of food. Therefore, human
interest in fermentation lays on the four potential advantages for food:
(1) improved shelf life and safety, (2) improved nutrition health
A. Septembre-Malaterre et al.
properties, (3) organoleptic modification and (4) production of active
principles of interest (Haaland, 2007; Hussain et al., 2016;
Katongole & Nicholas, 2008; Young-Hee, Tung-Ching, & Young-Chul,
2005). Fermented foods exist for millenniums and are consumed all
over the word.
A fermentation system is fundamentally composed of three ele-
ments: (1) the microorganisms, (2) the substrate and (3) the environ-
mental conditions. Microorganisms used for fermentation are very di-
verse. They include bacteria, yeasts and molds. Fermentation is based
on their action on the substrate that provides them nutrients for growth
and energy purpose. Various types of fermentation can occur (lactic,
alcoholic, propionic, malolactic, butyric, …) that share the same basic
feature of being an anaerobic catabolism of organic compounds whose
yield is lower than respiratory chain in term of energy production.
Fermentation is traditionally initiated by simple processing steps,
such as salt addition or anaerobic condition, which lead to an im-
balance of natural microbiota. Consequently, to salt addition and en-
closure of food in hermetic jars, anaerobic salt-resistant microflora is
favored and the most rapid to grow colonize the food. Amazingly,
fermentation has been implemented worldwide empirically with si-
milar methods and has resulted in a very large diversity of food pro-
ducts with different tastes (Franz et al., 2014; Lan et al., 2013; Nuraida,
2015; Peres, Peres, Hernández-Mendoza, & Malcata, 2012; Sagdic,
Ozturk, Yapar, & Yetim, 2014; Singh, Lee, & Lee, 2017). Among the
most popular fermented vegetables, cabbage is in first place: sauerkraut
in Europe and Western Asia, Kimchi in Eastern Asia, Dhamuoi in
Vietnam and Cortido in Latin America (Kusznierewicz et al., 2008; Park
et al., 2012). Fermented leaves are another popular foods, like black
tea, cassava leaves or mustard leaves (Fessard, Bourdon,
Payet, & Remize, 2016; Lan et al., 2013; Paludan-Müller, Huss, & Gram,
1999). Besides, radish, cucumber, olives, carrots, onions, Jack-fruit and
beetroots are fermented and called pickles (Gardner, Savard,
Obermeier, Caldwell, & Champagne, 2001; Juvonen et al., 2015;
Saeedi, Shahidi, Mortazavi, Milani, & Yazdi, 2015). Among fruits, fer-
mented lemon, but also durian fruit, and green mango are produced (Di
Cagno, Coda, De Angelis, & Gobbetti, 2013; Randazzo et al., 2016;
Sagdic et al., 2014). Lastly, fermented beverages like non-dairy kefir
and Kombucha (fermented brewed tea) result from complex microbial
fermentation ecosystem including LAB and yeasts (Chakravorty et al.,
2016).
Although in some cases the microbial species driving fermentation
are not precisely known, in many cases LAB are involved. Usually tra-
ditional fermented vegetables and fruits harbor a multi-species eco-
system. The most frequent species of LAB involved on fruits or vege-
tables fermentation are Lactobacillus plantarum, Lb. brevis, Lb.
rhamnosus, Lb. acidophilus, Leuconostoc mesenteroides, Lc. citreum, Lc.
fallax, Lc. kimchi, Pediococcus pentosaceus, P. acidilactici, Weissella con-
fusa, W. cibaria. During the last three centuries, with the improvement
of scientific knowledge and tools, fermentation evolved from a tradi-
tional small scale community centered process to an industrialized
controlled large scale market process with associated constraints among
which safety and constant repeatable quality. Nowadays, especially in
Western countries, fermentation process is well controlled and starters
are increasingly used (Corsetti, Perpetuini, Schirone, Tofalo, & Suzzi,
2012; Hammes, 1990; Leroy & De Vuyst, 2004). However, 90% of fer-
mented foods over the word are still produced with natural auto-
chthonous flora (Tamang, Shin, Jung, & Chae, 2016). The development
of starters emerged to ensure a constant foods sensory quality, but also
to improve safety. Other functional properties of starters, used for se-
lection purposes, are degradation of toxic and anti-nutritional com-
pounds, synthesis of bioactive molecules including antioxidants, en-
hancement of the bio-availability of nutritive compounds and
contribution to probiotic effect of foods. Physiological impacts that
were demonstrated to promote human health can be non-specific nu-
tritional impact (provide essential nutrients needs, correct a deficiency)
and often can go beyond with a specific physiological impact on a body
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function whether at cellular, organ or system levels. Apart from the
probiotic effect, the different ways by which LAB affect the nutritional
quality of foods are: (i) increase of nutrient density, mostly due to a
decrease of sugar content, (ii) hydrolysis of polymers from the raw
material and bioactive compounds content, (iii) biosynthesis of bioac-
tive molecules, (iv) degradation of toxic or anti-nutritional factors and
(v) synthesis of promoters for absorption and uptake.
4. Changes in bioactive compounds over lactic fermentation
4.1. Increase of nutrient density
The increase of nutrient density by lactic fermentation is mostly due
to a decrease of sugar content. Catabolism of disaccharides like sucrose
or lactose has been extensively studied for many LAB species
(Gänzle & Follador, 2012; Melanie Kostinek et al., 2005; Lynch et al.,
2015). Catabolism of disaccharides and of oligosaccharides implies the
transport of these molecules inside the cell where hydrolytic enzymes
are located. Starch is a polysaccharide for which extracellular enzymes,
amylases, are produced. Sucrose is one of the main oligosaccharides in
fruits. Interestingly, for short duration fermentation processes i.e. 17 to
24 h, under acidic conditions, sugar concentration essentially decreases
over prolonged storage under refrigerated conditions (Di Cagno et al.,
2009, 2011). In these conditions, metabolic activity of LAB is required
to ensure intracellular pH homeostasis and thus a high viability. This
metabolic process requires energy for proton-motive force main-
tenance. One could suppose that sugar consumption over storage is
related to this requirement of energy. On the opposite, sucrose, glucose
and fructose were totally depleted over cabbage or leek fermentation
when fermentation lasted several weeks (Jung et al., 2013; Wouters,
Bernaet, et al., 2013; Wouters, Grosu-Tudor, Zamfir, & De Vuyst, 2013).
Conversely, hydrolysis of polysaccharides occurs thanks to numerous
glucosidases and glycosyl hydrolases which release monomers of su-
gars.
Fermentation also contributes to increase protein, peptides and
amino acid content in foods from legumes. This leads to the opportunity
of using fermented vegetal products as an alternative and more sus-
tainable source of proteins to animal proteins in North and South
countries (Oguntoyinbo et al., 2016). Therefore, such possibility might
contribute to food security in countries with limited access to meat
protein sources but also in Western diet to replace animal proteins,
thereby limiting associated health, ethical and environmental issues.
4.2. Hydrolysis of phenolic compounds
Most of studies have focused on changes of antioxidant activity of
foods over fermentation (Table 3). From collected data, a high diversity
of patterns is observed. For instance, for cabbage, opposite trends in
antioxidant values over fermentation were obtained in different studies,
but the starter strains, the fermentation conditions and the antioxidant
assay differed between the studies. All these factors could easily explain
the observed apparent discrepancies. As a whole, a lot of studies, per-
formed with different substrates, with spontaneous fermentation or
different starter strains and with different assays to evaluate anti-
oxidant activity, did not succeed demonstrate predictable changes in
antioxidant activity over fermentation. However, a common conclusion
for several works is that the selection of starter can contribute to
maintain or to increase antioxidant activity when compared to spon-
taneous fermentation. Changes in antioxidant activity point out that
modifications of composition occur over fermentation. One of the main
mechanism that could explain antioxidant activity variation is the re-
lease of bioactive compounds from conjugated phytochemicals. Meta-
bolism of phenolic compounds by LAB has been reviewed by Rodriguez
et al. (2009). Interestingly, among LAB, Lb. plantarum is of great interest
since it possesses enzymes leading to the production of high-added
value compounds, such as powerful antioxidants (hydroxytyrosol and
 Table 3
Antioxidant activity modulation over lactic fermentation of fruits and vegetables.

Raw material Microorganisms Condition Antioxidant assay Observation Reference

Cabbage (Kimchi) Spontaneous (W. koreensis, L. brevis, Leu. gelidum) Short term: < 7 d; over TPC Values in over ripened kimchi are higher
ripened: > 2 y at 4 °C DPPH than in short term fermented
White cabbage Leu. mesenteroides CECT 219 or L. plantarum CECT 748 or both 7 d at room temperature ORAC Increase from 100% to 190%
A. Septembre-Malaterre et al.

Cabbage L. plantarum ATCC8014; L. rhamnosus ATCC9595 or L. brevis 24 h 37 °C, pH-controlled at 7 TPC, TFC 15–24% decrease, strain-dependent
ATCC8287 DPPH 5–13% decrease, strain-dependent
FRAP Slight decrease, strain-dependent
Leek L. plantarum IMDO 788, L. sakei IMDO 1358, or Leu. With salt 2.5%, 18 °C 3 weeks ORAC Increase by 17 to 51% Wouters, Bernaet, et al.,
mesenteroides IMDO 1347 DPPH Constant to high decrease depending on 2013
starter
TPC Slightly higher
Korean leek (Allium tuberosum Rottler) W. confusa LK4 or L. plantarum LK8 48 h 30 °C TPC Increase of TPC over 48 h with LK8
DPPH No significant change
FRAP Decrease in spontaneous fermentation
whereas 20% increase in observed for the
others
Soybean L. plantarum KFRI00144, L. delbrueckii subsp. lactis 48 h 37 °C DPPH Increase by 1.9–2.3 fold Young-Hee et al., 2005
KFRI01181, Bifidobacteria thermophilum KFRI00748, or B. ABTS Increase by 2.5–3.0 fold
breve K-101
Tomato juice E. faecium/faecalis POM3, L. brevis POM2 and L. plantarum 17 h 25 °C, then 40 d 4 °C ABTS Constant whereas control TAA decreased by Di Cagno et al., 2009
POM1 and POM35 2.5 to 3-fold
ABTS
L. plantarum LP254
Decrease
Red (RS) and green (GS) smoothies P. pentosaceus SWE5, L. plantarum POM1, Pr3 and W. cibaria 24 h 25 °C, followed by 30 d DPPH Decrease by 10% until end of storage Di Cagno et al., 2011
B6 (RS) OR L. plantarum K3 and F6, and L. pentosus P1 (GS) 4 °C TPC Constant

92
Carrots, fennels, melons, onions, Commercial water 48 h 25 °C TPC Slight to 49% (fennel) decrease, juice
tomatoes or strawberries Kefir (mix of L. fermentum, L. kefiri, Lc. lactis, Leu. mesenteroides DPPH dependent
and Saccharomyces cerevisiae) Moderate changes (increase or decrease),
juice dependent
Apple, quince, grape, kiwifruit, prickly Commercial water 48 h 25 °C TPC Slight to 53% (grape) decrease, juice Randazzo et al., 2016
pear or pomegranate juice Kefir DPPH dependent
Slight to 19% (grape) decrease, juice
dependent
Sweet cherry puree added with stem P. pentosaceus SWE5 and L. plantarum FP3 36 h 25 °C TPC No variation Di Cagno et al., 2011
infusion DPPH Decrease by 13% over started fermentation,
by 30% for unstarted
Pomegranate L. plantarum C2, POM1 or LP09 120 h at 30 °C, then 30 d at TPC, DPPH Decrease, but in a lesser extend for started
4 °C juice
Pear juice (four cultivars) L. acidophilus 72 h 37 °C TPC Significant decrease within 24 h
DPPH Increase or decrease depending on cultivar
and time
Pineapple juice L. plantarum 1OR12 and L. rossiae 2MR10 24 h 25 °C DPPH Spontaneous fermented juice is less
antioxidant than started
Pineapple Leu. pseudomesenteroides S14, W. cibaria S27 48 h 25 °C ORAC, ABTS, DPPH, No significant change Increase by 25% for Fessard et al., 2016
hemolysis S14
TPC Slight increase for S27
LDL oxidation
Tea L. plantarum ASCC 292 and L. brevis NPS-QW 145 48 h 37 °C TPC, HPLC Increase in phenolic acids, derivatives, and Zhao & Shah, 2016
flavan-3-ols; decrease in flavonols
DPPH Increase by 9–11%
Cellular ox Increase by 8–15%

TPC: total phenolic content, TFC: total flavonoids content, DPPH: 2,2-diphenyl-1-picrylhydrazyl assay, ORAC: oxygen radical absorbance capacity, FRAP: ferric reducing antioxidant power, ABTS: 2,2′-azino-bis(3-ethylbenzothiazoline-6-sulphonic
acid) assay, LDL: low density lipoprotein.
Food Research International 104 (2018) 86–99
A. Septembre-Malaterre et al.
Decarboxylation
Gallic acid
pyrogallol) or food additives approved as flavouring agents (4-vinyl
phenol and 4-vinyl guaiacol) (Rodriguez et al., 2009). Pyrogallol is
formed by Lb. plantarum in two steps involving first tannic acid hy-
drolysis by a tannase activity leading to gallic acid which is subse-
quently transformed by a gallate decarboxylase activity in pyrogallol
(Fig. 1). Tannase, named tannin acyl hydrolase (E.C. 3.1.1.20), is an
esterase acting on hydrolysable tannins and releasing glucose and gallic
acid. In 2004, Vaquero et al. (Vaquero, Marcobal, Muñoz,
Marcobal, & Muñoz, 2004) investigated lactic wine-isolates and tannase
activity was only detected for Lb. plantarum. However, a subsequent
study (Kostinek et al., 2007), on a larger panel of bacterial isolates,
successfully detected tannase activity in Leuconostoc spp. and in Weis-
sella spp.
Feruloyl esterase enzymes (E.C. 3.1.1.73) are also known as ferulic
acid esterases (FAE), cinnamoyl esterases and cinnamoyl ester hydro-
lases, release ferulic acid or p-coumaric acid from conjugated phenolic
acids. Cinnamic acids (i.e., ferulic and caffeic acids) that are esterified
to the vegetable cell walls should be enzymatically released to be ab-
sorbed by intestinal cells (Scalbert, Manach, Morand,
Rémésy, & Jiménez, 2005). Releasing of gallic and ellagic acids, of fla-
vonoids, and of flavan-3-ols, was related to esterases activity (Curiel
et al., 2015; Filannino et al., 2016; Zhao & Shah, 2016). Ferulic acid
esterase activity was particularly searched in probiotic bacteria
(Esteban-Torres, Reveron, Mancheno, de las Rivas, & Munoz, 2013;
Hole et al., 2012; Jo, Kim, & Baik, 2014;
Palaniswamy & Govindaswamy, 2016). This activity increases the con-
tent of free assimilable phenolic acids, but also bound phenolic acids
released from fibers. However, the increase of bioavailability may be
counterbalanced by phenolic acid decarboxylase activities, also present
in LAB (Filannino, Bai, Di Cagno, Gobbetti, & Gänzle, 2015). Beta-glu-
cosidase is one of the major enzymes responsible for the hydrolysis of
flavonoid conjugates during bacterial fermentation. This enzyme was
detected in many LAB species (Lee, Han, & Kim, 2012;
Michlmayr & Kneifel, 2014; Pyo, Lee, & Lee, 2005).
As a whole, molecular nature of phenolic compound can be mod-
ified through fermentation leading to new derived compounds with
biological activities potential including modification of microbiota po-
pulations and gut immunoglobulin levels (Massot-Cladera et al., 2014;
Parkar, Trower, & Stevenson, 2013; Rodríguez, Landete, de las
Rivas, & Muñoz, 2008). In addition, polyphenols bioavailability can be
positively influenced by glucosidase, over fermentation, thereby in-
creasing in situ radical scavenging potential as well as putative stimu-
lation of natural antioxidant body defenses (Escudero-López et al.,
2014; Michlmayr & Kneifel, 2014).
4.3. Derivatives of glucosinolates
Cabbage content in glucosinolates is high (Kusznierewicz et al.,
2008). These substances and their decomposition products are believed
to act as anticarcinogens and were shown to inhibit tumor cell growth.
The main glucosinolates hydrolysis product in fermented cabbage is
ascorbigen. It was shown recently that cabbage fermentation by LAB
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Fig. 1. Tannins gallic acid hydrolysis into pyrogallol
during fermentation.
Pyrogallol
resulted in an increase of antioxidant activity (Kusznierewicz,
Śmiechowska, Bartoszek, & Namieśnik, 2008). The gradual release of
substances with antioxidant activity from fresh cabbages was recorded.
Similar observations were made with fermented Chinese cabbage (E. K.
Kim et al., 2011; Sun, Chou, & Yu, 2009): an increase in radical
scavenging activity was noticed after cabbage fermentation compared
to control fresh cabbage. Total phenolic compounds level increased
more than flavonoids content after fermentation. A first clinical study
with overweight and obese patients that consumed fermented cabbage
showed positive effects on various factors associated with metabolic
syndrome. In fermented cabbage, particularly high differences were
found for isothiocyanate levels from decomposition of sinigrin, glu-
coiberin and glucoraphanin (Tolonen, Rajaniemi,
Pihlavaa, & Johansson, 2004). The observed changes can be related to
Lactobacillus ability to degrade glucosinolate in allyl-isothiocyanate
(Llanos Palop, Smiths, & Brink, 1995). Knowledge of the bioavailability,
transport and metabolism of glucosinolates after consuming Brassica
vegetables is an essential prerequisite for understanding the mechan-
isms of their protective effects in humans. The occurrence of such
compounds in food determines its functional and health promoting
nature.
4.4. Bioactive peptides
Bioactive peptides have been mainly studied from milk or whey
hydrolysis during lactic fermentation (López-Fandiño, Otte, & van
Camp, 2006), but also from fermented soybeans (Gibbs, Zougman,
Masse, & Mulligan, 2004), and more recently from grapes (Aredes
Fernández, Stivala, Rodríguez Vaquero, & Farías, 2011) and cereal
flours (Coda, Rizzello, Pinto, & Gobbetti, 2012). Milk and derived whey,
soybeans and cereals are rich in proteins, but in each of these foods the
proteins are different. In all cases, LAB succeed in releasing bioactive
peptides. Bioactivity of many peptides, either naturally present in raw
food materials or released by food fermentation was reviewed in the
literature (García, Puchalska, Esteve, & Marina, 2013). Bioactive pep-
tides are generated during fermentation process through enzymatic
proteolysis, performed by single or multiple, specific or unspecific
proteases, of larger protein molecules usually not bearing any bioac-
tivity (Humiski & Aluko, 2007). Health impacts include modulation of
immune system with associated positive influence on inflammatory
processes. Reports have also shown specific activities on cardiovascular
system such as inhibition of conversion enzyme and anti-atherosclerosis
effects thereby preventing hypertension co-morbidity (Sirtori, Galli,
Anderson, & Arnoldi, 2009). Bioactive peptides can regulate specific
physiological functions, such as the reduction of blood pressure or
possess free radical-scavenging activities. More specifically, the angio-
tensin I-converting enzyme (ACE) regulates blood pressure. Its inhibi-
tion by bioactive peptides can exert an anti-hypertensive effect. The
antioxidant effect of peptides was related to their amino acid compo-
sition, conformation and hydrophobicity. These peptides are inhibitors
of lipid peroxidation, exert a direct radical scavenging activity or che-
late metal ions that catalyze the generation of oxygen reactive species
A. Septembre-Malaterre et al.
Phytate
Fig. 2. Phytate enzymatic degradation during fermentation process.
Adapted from Lei & Porres, 2003.
(Rizzello et al., 2016).
4.5. Secondary metabolites, short chain fatty acids and vitamins
Secondary metabolites generated by fermentation process are no-
tably interesting regarding health. Isoflavones, from soja for example,
were demonstrated with anti-cancer properties, limiting the onset of
breast carcinogenesis (Celligoi, Santos, da Silva, & Baldo, 2015). Poly-
saccharides, polyunsaturated fatty acids and polyphenols were corre-
lated with direct significant health effects on diabetes and cardiovas-
cular diseases including coagulation and inflammation. Conversely,
compounds such as sorbitol or mannitol do not demonstrate direct but
indirect effects through their lower caloric impact when compared to
glucose whose glycemic index is elevated. Therefore, these compounds
reduce excessive glucose pressure on obese and diabetic subjects,
sparing over solicited functions such as endocrine pancreas and insulin
signaling in insulin-sensitive tissues such as muscles, liver and adipose
tissues (Michalak & Chojnacka, 2015).
Short chains fatty acids are produced from hydrolysis of lipids. They
might be produced during the fermentation process and are used as
energy source by colon cells and leads, for example, to anticarcinogenic
properties of the final product correlated with a higher content in bu-
tyric acid (Parra-Matadamas, Mayorga-Reyes, & Pérez-Chabela, 2015;
Pereira-Caro et al., 2015).
Fermentation of fruits and vegetables is frequently associated with
an increased content in vitamins. They notably include vitamins of the
B group as well as others such as vitamin K. Vitamins are compounds
without energetic value that are mandatory for human health as these
molecules are co-factors of numerous enzymes involved in all types of
metabolism contributing to organs function and structure. LAB are
particularly interesting in this matter as it is possible to promote im-
provement of a predesigned B vitamin ratio content, of the fermented
product, as a function of the starter composition used (Capozzi, Russo,
Dueñas, López, & Spano, 2012).
4.6. Synthesis of exopolysaccharides
Exopolysaccharides (EPS) are long chains of homo- or hetero-
polysaccharides, consisting of (branched) repeated units of sugars or
sugar derivatives produced outside microbial cells.
Homopolysaccharides are formed of repeated units of either glucose or
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Inositol
fructose, classified as α-D-glucans (dextran, mutan, reuteran, and al-
ternan) and β-D-glucans, whereas those containing fructose are fructans
(levan and inulin-types). Heteropolysaccharides are composed com-
monly by glucose, galactose, and rhamnose and in some cases by N-
acetyl-D-glucosamine and N-acetyl-D-galactosamine. It was demon-
strated that LAB belonging to the genera Lactobacillus, Lactococcus,
Leuconostoc, Pediococcus and Weissella, in particular W. cibaria, W.
confusa and W. hellenica, are producing a large diversity of EPS, de-
pending on the carbon source (Fessard et al., 2016; Juvonen et al.,
2015; M. J. Kim, Seo, Hwang, Lee, & Park, 2008; Pan & Mei, 2010).
The most studied health effect is indirect and called prebiotic: EPS
stimulate the growth of probiotic bacteria. They may also be involved
in biofilm formation, thus helping ingested bacteria to survive in the
intestinal tract. EPS are also proposed to have direct effects, such as
immunomodulation effects, antioxidant activity and cancer prophylaxis
(Caggianiello, Kleerebezem, & Spano, 2016; Franco-Robles & López,
2015; Kwak, Cho, Noh, & Om, 2014; Pan & Mei, 2010). These molecules
scavenge hydroxyl and superoxide anion radicals, and non-radical
oxidant molecules (hydrogen peroxide, singlet oxygen). In vivo, as
tested in mice diet, EPS can stimulate the production of reactive oxygen
species detoxification enzymes, such as NADPH oxidase, catalase and
superoxide dismutase.
4.7. Degradation of anti-nutritional factors
The use of fermentation as an integral part of food detoxification
processes is widely practiced. Lactic fermentation using Lb. plantarum is
involved in lowering anti-nutritional factors and detoxifying food. The
content of proteases and trypsin inhibitor was also reduced by fer-
mentation. LAB, through their enzymatic equipment, help neutralize
anti-nutritive factors such as phytates, saponins, tannins, cyanogens or
trypsin inhibitors (Fig. 2) (Lai, Hsieh, Huang, & Chou, 2013).
For example, galactosidase is able to reduce anti-nutritive factors
such as phytic acid, tannins, trypsin inhibitors (Adeyemo & Onilude,
2014). This effect can be associated with modification of minerals
bioavailability (Arslan & Erbas, 2015; Gaspar & Crespo, 2015) and pro-
teins whose digestibility is also favored (Kaur, Jha, Sabikhi, & Singh,
2014). In addition, galactosidase is recognized to metabolize cyano-
genic glucosides present in some vegetal matrixes such as cassava roots,
bitter almonds or whole sorghum (Oguntoyinbo et al., 2016).
A. Septembre-Malaterre et al.
5. Bioavailability and interactions
5.1. Bioavailability of bioactive compounds
One key issue for bioactive compounds provided by diet is their
bioavailability. Bioavailability can be defined as the rate and extent to
which the compound is absorbed and becomes available for host cel-
lular metabolism. It involves gastrointestinal digestion which releases
the compounds from foods, absorption by epithelia cells, tissue dis-
tribution and bioactivity. Release from food is highly dependent of in-
testinal microbiota, digestive enzymes, and for fermented foods of in-
gested bacteria. Molecular structure of the bioactive molecule
considerably affects its absorption: high molecular weight molecules
are less absorbed and therefore cleavage of the glucoside moiety is
important for absorption of conjugated phenolic compounds.
Interactions with other food components, such as proteins, play also a
role (Bohn et al., 2015; Aedín Cassidy & Minihane, 2017; Rein et al.,
2013). From those elements, few changes in antioxidant activity as-
sayed by in vitro tests were recorded. Deeper investigation of the
composition variation and associated in vivo testing to evaluate the
effective antioxidant effect are required. As reviewed above, thanks to
the activity of various enzymes, fermentation of fruits and vegetables
tends to improve bio-accessibility and/or bio-availability of various
type of compounds such as proteins, amino-acids, vitamins and anti-
oxidants compounds (such as polyphenols).
5.2. Prebiotics
Prebiotics are defined as “a substrate that is selectively utilized by
host microorganisms conferring a health benefit”. This updated concept
includes not only carbohydrate but also non-carbohydrate (free fatty
acids, …) types of molecules. Also, a reasonable proof of causal link
between the prebiotic and health benefits (gastrointestinal tract, car-
diovascular system, …) (Gibson et al., 2017). It is noteworthy to specify
that fruits and vegetable are a less valuable source of prebiotics than
cereals. Among prebiotics, EPS have the ability to promote specific
microorganisms' over other ones. The most known prebiotics such as
fructooligosaccharide or inulin promote growth of specific probiotic
bacteria. It should also be mentioned that prebiotics tends to positively
influence the composition of the enteric microflora thereby sustaining
optimal health through probiotics (Arslan & Erbas, 2015). Prebiotic
were recorded with significant biological activity through modification
of the host gut microbiota and reduction of insulin-resistance and as-
sociated elevated blood pressure as well as overweight in obese patients
(G R Gibson & Roberfroid, 1995; Glenn R. Gibson, Probert, Loo,
Rastall, & Roberfroid, 2004). Isomaltooligosaccharide, produced during
fermentation of Kimchi, demonstrated prebiotic potential on resident
bacteria of colon (Cho et al., 2014).
5.3. Probiotic effect
Probiotics are defined as “live microorganisms that, when ad-
ministered in adequate amounts, confer a health benefit on the host”
(FAO/WHO, 2001; FAO/WHO, 2002). This definition was com-
plemented by recommendations from a consensus panel of expert in
2014. It included “microbial species that have been shown (…) to
confer benefits to health” but excluded live cultures from fermented
foods unless a health benefit was proven (Hill et al., 2014). Therefore
fermented foods may be considered as potential probiotic but they have
to be first demonstrated to have a positive influence of the consumer
health (Hill et al., 2014; Roberfroid, 2000). Fermented food can be
considered as a potential delivery system for probiotics in the sense that
this type of food contains living bacteria of potential interest for human
health. The positive impact on the consumers is not limited to the di-
gestive system as it may also positively influence other system function
such as immune or respiratory system for example (Ray & Montet,
95
Food Research International 104 (2018) 86–99
2015). Among probiotics, LAB were described to promote im-
munomodulation, cancer prevention, lower cholesterol absorption and
therefore cardiovascular disease prophylaxis but also reduced food al-
lergies (Aragón, Perdigón, & de Moreno de LeBlanc, 2014). In addition,
these bacteria can modify and positively influence human gut micro-
biota whose composition disturbances are known to be correlated with
obesity, chronic low grade inflammation, oxidative stress and asso-
ciated risk such as diabetes and cardiovascular diseases. These positive
impacts on health should nonetheless be moderated since persistence of
probiotics and their effects do not seem to last durably. The mechanism
of action of probiotics effects on health, whether local (intestine) or
global (whole body and functions) seems to be complex and still poorly
understood in spite of the various hypothesis that probably all con-
tribute, at various degrees, to the observed health benefits. Bacter-
iocins, for example, are protein derived molecules, produced by bac-
teria during the fermentation process. These compounds have an
antimicrobial activity aiming at neutralizing microorganism competi-
tion to promote dominance and supremacy of the fermentation species
over other bacteria. This ability helps preventing food contamination by
undesirable or potentially pathogen bacteria. It should also be men-
tioned that the impact of bacteriocins is further consolidated by
changes, such as pH levels and ethanol production, in reducing the
contamination risks by biohazardous microflora. A missing data re-
mains to be determined about probiotics: their safety/efficacy dose
range ratio (Parvez, Malik, Ah Kang, & Kim, 2006). Therefore, fer-
mented foods and associated probiotics interest for health needs further
investigations to better understand the modality of their influence
taking into account host microbiota specificity and the concepts of di-
rect local effects and indirect global effects on the human body
(Tamang, Watanabe, & Holzapfel, 2016).
6. Conclusion
Food consumption of vegetable materials, whether fruits, vege-
tables, leaves, stems, can prevent chronic diseases such as metabolic
syndrome, cancer, etc. These benefits are due to the presence of fibers
and micronutrients, such as vitamins, minerals, but also phytochem-
icals. Low fruit and vegetable consumption is one of the top ten risk
factors for mortality. Up to 1.7 million lives could be saved each year
with adequate consumption of fruits and vegetables at the world level.
Therefore, in addition to fresh products, fermented foods now represent
an additional source of consumption of fruit and vegetable with more
diversified and sometimes improved health properties. The lactic fer-
mentation of these vegetable materials makes it possible to propose an
alternative mode of intake, with new sensory properties, for foods de-
veloped according to a sustainable approach. Lactic fermentation
modifies the composition of the fermented materials and by the meta-
bolic microbial action can improve the beneficial health benefits of
food. The improvement of the functional properties of foods by lactic
fermentation results from several mechanisms: elimination of anti-nu-
tritional factors, production of metabolites with a positive effect
(bioactive peptides, exopolysaccharides), improvement of the bioa-
vailability through polymers hydrolysis (esters of phenolic compounds),
increased vitamin, mineral and phenolic compounds, leading to an in-
crease in the antioxidant capacity of the product and possibly an effect
on the intestinal flora and the intestinal absorption of bioactive com-
pounds. The improvement of the antioxidant properties of food by
fermentation and their effects on the state of cellular oxidation or
cancers development are reported in the literature. However, it seems
that these properties strongly depend on the plant material - microbial
strain couple and the molecular mechanisms underlying these ob-
servations are poorly documented. Indeed, very few bibliographic data
regarding the bioavailability and the potential health effects of the
newly formed compounds by lactic fermentation exist. Development of
tools for global analysis, such as metabolomics, proteomics and tran-
scriptomics, would considerably help in that respect. Determination of
A. Septembre-Malaterre et al.
food composition on fresh fruits and vegetables, on fermented foods
and furthermore in the gastrointestinal tract would be useful to eval-
uate the extent of molecular changes at each step. Moreover, enzyme
activity is the cornerstone promoting these changes. Thus, an extensive
knowledge of these activities and interactions between ingested food,
fermented food microorganisms and intestine microbiota is required.
This has become accessible with the development of global molecular
tools. Such approaches just start to be applied, as performed to in-
vestigate the mechanisms of pectin degradation (Despres et al., 2016)
by a prominent human gut bacterium.
Conflict of interest
The authors declare that there is no conflict of interest.
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