Dietary fiber: Properties, Recovery & Applications

Chapter 4. Dietary fiber & metabolism

Ramona Suharoschi1, Oana Lelia Pop1, Romina Alina Vlaic2, Carmen Ioana Muresan1, Crina
Carmen Muresan2, Angela Cozma3*, Adela Viviana Sitar-Taut3, Romana Vulturar4, Simona
Codruta Heghes5, Adriana Fodor6, Cristina Adela Iuga5,7
1
Laboratory of Molecular Nutrition and Proteomics, Institute of Life Sciences, University of Agricultural Sciences
and Veterinary Medicine of Cluj-Napoca, 3-5 Manastur Street, Cluj-Napoca, 400372, Romania;
[email protected] (R.S.); [email protected] (O.L.P.); [email protected]
(C.I.M.)
2Department of Food Engineering, University of Agricultural Sciences and Veterinary Medicine of Cluj-Napoca, 64

Calea Floresti Street, Cluj-Napoca, 400445, Romania; [email protected] (R.A.V.);

[email protected] (C.C.M.)
3 Department Internal Medicine, Clinique IV, Faculty of Medicine, “Iuliu Hațieganu” University of Medicine and

Pharmacy, 6 Louis Pasteur Street, Cluj-Napoca 400349, Romania; [email protected] (A.C.);

[email protected] (A.V.S.T.)
4 Cellular and Molecular Biology, Faculty of Medicine, “Iuliu Hațieganu” University of Medicine and Pharmacy, 6

Louis Pasteur Street, Cluj-Napoca 400349, Romania; [email protected] (R.V.);

5 Department of Pharmaceutical Analysis, Faculty of Pharmacy, “Iuliu Hațieganu” University of Medicine and

Pharmacy, 6 Louis Pasteur Street, Cluj-Napoca 400349, Romania; [email protected] (S.C.H.); [email protected]
(C.A.I.)
6 Diabetes and Metabolic Diseases, Faculty of Medicine, “Iuliu Hațieganu” University of Medicine and Pharmacy, 6

Louis Pasteur Street, Cluj-Napoca 400349, Romania; [email protected] (A.F.)

7 Department of Proteomics and Metabolomics, MedFuture Research Center for Advanced Medicine, “Iuliu

Hațieganu” University of Medicine and Pharmacy, 4-6 Louis Pasteur Street, 23 Gh. Marinescu Street, Cluj-
Napoca, 400349, Romania; [email protected] (C.A.I.)

* Corresponding author: [email protected]

Abstract
A healthy diet rich in dietary fiber (DF) is considered benefic for human health and recommended
dietary intake of dietary fiber is established all over the world. The physiological effect of dietary
fiber is mostly related to its behavior during the digestion. In this chapter, the behavior of dietary
fiber in the human digestive tract is discussed and linked to its physiological effect with special
attention to four aspects of such behavior: (1) the modulation of bioavailability by the plant cell
walls, (2) the effect of dietary fiber on the rheological and colloidal state of digesta, (3) the binding
of dietary fiber with phenolic compounds, bile salts, mineral ions, and digestive enzymes, and (4)
dietary fiber fermentation in the large intestine and the corresponding effect on microbiota
composition.
Keywords: dietary fiber, digestion, metabolism, gut microbiota

1. Introduction

It is generally known that the population consuming diets rich in dietary fiber (DF) has a lower
incidence of chronic diseases. The digestion and absorption in the gastrointestinal (GI) tract of DF
is determined by the structural complexity of those. Importantly, all plant cell walls are composed
by DF that are digested selectively by microbiotic enzymes and not by host enzymes. Thus,
complex DF in plant cell walls function as both substrate for the microbiota and as a sustained
release delivery system of other plant-cell derived nutrients and biomolecules for both microbiotic
and host utilization. The cell wall structure is different from plant cells to another and determine
the specific site of plant cell wall degradation and nutrient release.

A healthy diet is generally considered to include DF as an important element and this has lead to
dietary recommendations for an optimal consumption level. The DF positive effect on health is
related to its behavior in the GI tract.

Foods rich in DF category are extremely large and diverse from whole foods (unrefined) to fully
refined foods, and currently no official difference based on the degree of refinement is made. Foods
with added refined nutrients (e.g., sugar, starch or oils) or any food which has been structurally
disrupted on the macroscopic level are considered as refined.

Beneficial effects of DF on human health have been demonstrated, e.g. in immunity, diabetes,
cancer and cardiovascular diseases. Direct effect of DF on boosting the immune response by the
interaction of bacterial and fungal β-glucan with specific receptors of epithelial immune cells have
been demonstrated [22]. However, this is an exception not the rule. The review studies that
describe the physiological effect of DF related to behavior through digestion address one or few
aspects of this behavior and a comprehensive and detailed approach is needed. In this chapter, four
health-linked aspects of DF behavior in the GI tract are discussed: (1) the effect of plant cell walls
on bioavailability, (2) the effect of DF on the rheological and colloidal state of digesta, (3) the
binding of DF with phenolic compounds, bile salts, mineral ions and digestive enzymes and (4)
DF fermentation in the large intestine and the effect on gut microbiota composition.
 2. The modulation of bioavailability by the plant cell walls

A widely recognized fact is that the incidence of chronic diseases is inversely proportional to the
consumption of dietary fibers [1, 2]. Human health is modulated through synbiotics effects borne
from dietary fiber consumption. The pathway to understanding the physiological effect of the
dietary fibers is to spotlight their behavior during digestion. A whole mechanism is contributing
to human health when dietary fibers are ingested.
Usually, dietary fibers belong to the structural system of the plant, namely in the cell wall. The
key role of the wall cell is protection and the principal surface of interaction with the external
medium and environment. There are two layers that compose the plant cell wall: a primary cell
wall that forms during plant growth and a secondary cell wall that forms after plants have
completed their growth [3].
In most of the cases, the primary cell wall structure has a network of cellulose fibrils-hemicellulose
(xyloglucan, arabinoxylans, β-glucan, etc.) embedded in a network of pectins [4].
In this arrangement, there are different roles, namely: cellulose and hemicelluloses function as a
load-bearing structure, while pectin confers plasticity and controls the porosity of the cell wall.
Dick-Perez [5] in his work propose a new approach of the cell wall characterization, with a single
structure made from pectin, cellulose and hemicellulose. The cell wall of plans is different,
depending on the plant type. In grains, excepting rice, and seeds cell walls are comparatively
poorer in pectins and cellulose where as arabinoxylans and β-glucans dominate [6].
Secondary cell walls are made up of a network of cellulose and lignin whereas pectins are low or
absent [7]. From nutritional point of view, the primary cell wall, secondary cell wall has a no
significant amount of components of human diet. Pectins are also the main component of the
middle lamella, a thin layer that is responsible for the adhesion of adjacent cells in plant tissues.
From the chemical point of view, the cell wall structure is organized in different manners and is
related to factors as plant species and is affected by environmental and developmental factors [8,
9]. Cell wall structure and composition play an essential role in the bioavailability of micro- and
macronutrients for the human body. For these dietary fibers, present in the cell wall membrane, to
be suitable for absorption, intracellular molecules must pass through the physical barrier
represented by the cell wall. The nutrient entrapped in the cell are well protected and their release
is depended by the release ensured by the cell disrupture. However, the entrapped nutrients can be
released, in small proportions and depending on the size of the molecules, disrupt the pores of the
cell wall. Thereafter, digestive fluids that enter into the cell are able to start the digestion of
macromolecules inside the cell. The functions of the plant cell wall are directly influenced by
factors as the wall chemical composition, structure, digestion influence, industrial and home food
processing but also mastication [10, 11]. Dietary fibers that act as prebiotics, suffers minor
chemical changes to cell wall when exposed harsh gastric conditions [12].
Mandalari et al., [13] discovered a maximization of the cell wall thickness after 2, 6 and 12 hours
upon digestion and suggested a possible facilitation of diffusion through the swollen cell walls.
Broxterman et al [14] are talking about swelling of cell wall in raw carrots matrix and solubilisation
of pectins. In fruits and vegetables, in their raw state, reduction of particle size by extreme
mechanical grinding, milling or chewing increases the fraction of fractured cells which are exposed
on the particles exterior. This reveals frequently reported statement that the lower the particles size
the higher is their bioavailability.
In raw fruits and vegetables, with strong structure, where cell adhesion is quite strong, tissue
rupture happens mostly through cell walls. In this particular situation, under mechanical stress or
oral processing the number of fractured cells is considerable increasing. In fruits and vegetables
with weaker structure, after thermal treatment, cell adhesion is weakened and tissue fraction would
happen principally along cell walls, producing, upon mechanical stress, clusters of intact cells. On
the other hand, the solubilisation of pectins and implicitly of other constituents present in the cell
wall, may facilitate the permeability of the cell wall by increasing the size of the cell wall pores.
Aribas-Agusti et al [15] demonstrated that permeability of carrot cell wall towards molecules of
limited size is facilitated after heat treatment because of the solubilisation of native pectin.
In conclusion, a must be recall that nutrients that have not been absorbed in the small intestine
because cell entrapment can be released in the large intestine upon fermentation of cell wall
material by the local microbiota. Most of these compounds, e.g. polyphenols, may be absorbed by
the colon epithelium after conversion by the gut microbiota.
The connections among plant cell wall ingestion, effect of processing, microbial fermentation and
bioavailability in the large intestine is a subject that need further investigation. The understanding
of this mechanisms are of interest for scientists, health authorities, food engineers and chefs alike.
 3. The effect of dietary fiber on the rheological and colloidal state of digestion

Populations that consume a large amount of dietary fiber in their daily diet were reported to have
a lower incidence of cardiovascular disease compared to low-fiber consumers [16, 17]. This also
as beneficial effects in preventing and treating obesity[18, 19], insulin and diabetes mellitus [16]
and cancer [20]. Grain fiber-rich diets were shown to improve insulin resistance and the risk of
diabetes. The effects of grain fiber -rich diets on weight loss seem to be moderate [16].
Systolic and diastolic blood pressures have a decreasing trend under the influence of viscous
soluble fiber. Diets which include viscous fiber may play a role in reducing the cardiovascular
disease risk, thus contributing to an overall improvement in blood pressure [21].
Nutritional components are generally recommended for their intrinsic nutritional value and their
effect on the human body. The beneficial effect of dietary fibers on human health is due to their
behavior in the gastrointestinal tract. The beneficial effects of dietary fiber on human health due
to the bacterial and fungal interaction of β-glucan with epithelial immune cell-specific receptors
have been reported alongside with their contribution to the stimulation of immunological responses
[22].
The physiological behavior of dietary fibers in the tract is extremely varied, depending on their
interactions [23].
The type(s) of dietary fibers and their content in monosaccharide constituents may be different
according to the form of the monosaccharides (α or β), the type of chemical linkage, the length of
the linear polymer chain, the presence, distribution and composition of branches attached to the
linear chain. Oligosaccharides and polysaccharides can be differentiated based on the length of the
linear chain [24]. The polysaccharides that can be found in dietary fibers include resistant starch
and non-starch polysaccharides. Resistant starch enters the colon and escapes digestion. The
amount of resistant starch from the food differs from that reaching the digestive tract because
resistant starch may depend on the meal composition, storage and cooking conditions and on the
physiological factors [25].
Particle size, solubility, moisture and viscosity are the essential features of dietary fibers, which
help understanding their behavior during digestion [26]. The solubility of the fibers is dependent
on their chemical structure and interactions with water molecules. Solubility depends on the pH
and the ionic concentration of the environment. Thus, dietary fibers are classified into soluble
(pectins, inulin, mucilages, glucomannan, β-glycans, oligosaccharides, etc.) and insoluble fibers
(cellulose and hemicellulose, some types of resistant starch, lignin, etc.) [23, 27]. The most
important sources of soluble fibers are: fruit, berries, certain vegetables (e.g. pectins from guava,
carrots, beans, lentils; nuts); germ fraction from oat and barley products; guar; psyllium; and
insoluble fiber: whole-grain and bran products; also skins of fruit; cucumbers, tomatoes; hull of
grains; brown rice; legumes; nuts, almonds. Certain food products contain both soluble and
insoluble dietary fibers. Generally, soluble dietary fibers are viscous and while the viscous fibers
are insoluble - Figure 1 [27].
Viscosity is the main rheological property of fibers. This is defined as resistance to flow, and
dietary fibers have a variable ability to produce a viscous solution after dissolution in water. This
capacity depends to a great extent on the molecular weight and the concentration in dietary fibers
and is positively correlated with its solubility [23].
Despite the fact that dietary fibers are considered to be an inactive material in the digestive tract,
they undergo structural and chemical changes, which influence their behavior during digestion as
shown in Figure 2. Dietary fibers are part of the cell wall in plant tissues. In the case of preparations
or ingredients rich in dietary fibers, they appear as particles of varying size, purity and composition
and are either part of the cell walls or can be isolated from them. During mastication, mechanical
processes cause cell wall breakage and reduction to smaller pieces/fragments. The level of
fragmentation has a major impact on the bioavailability of nutrients. In their route through the
stomach and intestines, the dietary fibers absorb varying amounts of water and swell according to
their specific hydration capacity. This swelling is followed by partial solubilization [23].
Solubilization is limited when dietary fibers are part of cell wall structures due to strong chemical
interactions with other cell wall components. These interactions may become less strong as
digestion occurs and is in progress [28]. Favored by thermodynamic processes, the dietary fibers
which are soluble in liquid fraction are easier to absorb in the digestive tract. When in solution,
the dietary fiber molecules can remain dispersed or aggregated in varying proportions, and can
produce colloidal particles [29].
The rheological and colloidal state of digestion of dietary fibers is determined by their interaction
with digestive fluids and other dietary components.
The ability of certain dietary fibers to absorb water leads to the formation of certain viscous
solutions. This mechanism has the effect of lowering post-prandial blood glucose [30] and serum
cholesterol [31]. Recent research findings from in vitro analyses of digestion showed a slight
decrease in starch hydrolysis rate and diffusion coefficients for glucose (1.5-2.5 times) compared
to a 100-fold rise in digesta viscosity. They had an effect of delay of gastric emptying, which led
to a decrease in the post-prandial glycaemia [25].
Viscosity and enzymatic/absorptive processes do not always have a linear relationship. Following
in vitro studies of gelatinised starch/guar gum mixtures on the small intestinal digestion, viscosity
during digestion was analysed and it was found that it did not decrease alongside with starch
hydrolysis [32].
The viscosity of dietary fibers during digestion can be increased together with the liquid fraction
and the changes affecting the physical properties in the structure of particles. Soluble dietary fibers
have the capacity to increase the viscosity of the liquid fraction during digestion (pectins, gums,
arabinoxylans and β-glucans) and this capacity is influenced by several factors. Rigid or rod-like
polymers have a higher viscosity than linear polymers. Thus, branched polymers tend to have a
smaller volume than linear polymers [33].
High Molecular Weight polysaccharides result in more viscous solutions than their
depolymerisation products. During food processing, dietary fibers may become depolymerised.
For instance, pectin is hydrolised during the process of thermal treatment. β-glucan may partially
become hydrolised during baking (e.g. viscosity decreases in the process of baking) [34]. Dietary
fiber molecules of the same class may largely vary in molecular weight depending on the extraction
procedure. Viscosity may also be influenced by differences in the chemical structure of DF from
the same class. For instance, viscosity of pectin solutions decreases [35]. Viscosity differs
substantially depending on the arabinoxylans extracted from rye, and it is related to molecular
weight of water molecules and to the degree of branching [36].
Viscosity of a dietary fiber solution varies depending on the dietary fiber type and the dilution of
dietary fiber with digestive fluids [37]. For instance, xanthan gum retained its viscosity very well
after mixing with digestive fluids as compared to other gums, such as locust bean, guar, fenugreek
and flaxseed [38]. Lazaridou et al. [39] reported that solubility in the intestines of β-glucan from
coarse barley flour is lower than that from fine barley flour. The ionic strength and pH of the
solution influence the viscosity of polysaccharide solutions. For instance, a multiple-course meal
may induce the occurrence of different polysaccharides in the digestive tract. This may have
synergic or antinergic effect on viscosity [37, 40].
Dietary fibers have the capacity to change the physical characteristics of the material they are part
of, depending on the disintegration kinetics of foods or on the distribution of particles during
digestion. Thus, insoluble dietary fiber may increase their viscosity during digestion and can slow
down nutrients absorption. This was reported for crystalline cellulose [41], wheat bran particles or
wood particles [32].
According to in vitro studies reported by Houghton et al. [42], the viscosity of alginate is the double
of that of an equivalent amount of alginate incorporated or co-digested with bread. Bread dietary
fiber form a barrier around starch granules, which denies access to α-amylase [43], but the same
fibers may also change the gluten network so that it can protect starch granules against α-amylase
[44]. In physiological conditions of the gastro-intestinal tract, gels have a solid consistency and
are formed of polymers cross-linked to create an interconnected network immersed in water or
another liquid medium [45]. From a rheological point of view, a gel is a viscoelastic system [46].
The hydrocolloid properties of dietary fiber have mainly been studied in the stomach but also in
the small intestine even though evidence was present with regard to the fact that β-glucan and
chitosan could form gels there [29]. The gel content formed in the stomach was reported to increase
satiety and decrease the gastric emptying rate [47].
Gels have to resist the mechanical stress that the gastric acidic fluids exert. Alginate forms such
gels in the stomach. However, its gelation in the stomach is extremely fast and the properties of
the gel are too sensitive to the conditions in the stomach [47, 48]. An alternative to alginate can be
replaced by gellan gum, a complex bacterial polysaccharide, and pectins [49]. These gels can also
be influenced by other nutritional components with which they come into contact, such as: a
mixture of whey protein and pectin which forms an intragastric gel at much lower concentrations
of poly saccharides [50]. Also xanthan gum and carragenans are able to form gels upon mixing
with gastric juices and whey protein isolates. Another effect of dietary fiber refers to their impact
on lipid emulsion stability. In the gastric and duodenal compartments, the polymer stabilization /
destabilization of emulsions depends on dietary fiber type, concentration, other biopolymers in
solution, pH and ionic strength and the nature of the emulsion-stabilizing surfactant [51].
Minekus et al. [52] reported the effect of partially hydrolysed guar gum on full-fat yoghurt in
which the lipolysis rate was reduced and cholesterol was absorbed. Beysseriat et al. [33] showed
that, in the small intestine (neutral pH), positively charged chitosan can destabilise corn oil in water
emulsion and induce coalescence through a bridging flocculation mechanism. In contrast,
negatively charged pectin cannot adsorb onto lipids but it can destabilise the emulsion by inducing
reduced flocculation. Espinoza-Ruiz et al. [53] also found increased coalescence of corn oil in
water emulsions in the presence of pectins extracted from banana, passion fruit and corresponding
decrease in the digestion rate of fats. Another type of DF, namely: methyl cellulose, can also induce
reduced flocculation in corn oil/water emulsions [53].

Figure 1. Classification of fibers, typical attributes and shared effect [27]

 Figure 2. Changes of structural properties of dietary fiber during digestion [23]

4. The binding of dietary fiber with phenolic compounds, bile salts, mineral ions and
digestive enzymes

4.1.Binding with phenolic compounds

The group of phenolic compounds are plant secondary metabolites included in different classes
defined by the number of phenol rings, by their linkage and type of attached functional groups. A
great number of compounds belongs to this group: phenolic acids, hydroxycinnamic acids,
xanthones, coumarins, acetophenones, flavonoids etc. From these ones, flavonoids are the most
important ones occuring in plants are phenolic acids, flavonoides, stibenes and lignans [54].
phenolic compounds, especially polyphenols, have potential health benefits associated with their
biological properties like: antioxidant [55], anti-estrogenic [56], immunomodulatory [57], anti-
carcinogenic [58], cardioprotective [59] activities. In additon, an important biological property is
the ability to interact and bind the macronutrients found in food.
Dietary fibers (DF) represent a vast category of polysaccharides with chemical and structural
diversity. DF can bind, adsorb or entrap other dietary compounds, thus having physiological
effects. phenolic compounds are omnipresent in vegetables, fruits, cereals and nuts being thus
found in plant-based food and beverages (beer, wine, tea, coffee). Their biological properties and
health effects will depend on their intake and bioavailability which in turn are conditioned by
different factors including their binding capacity to DF. phenolic compounds have hydrophobic
rings and hydrophylic groups able to bind to DF at different sites [60].
The nature of interactions between phenolic compounds and DF was intensively investigated and
reviewed [8, 60, 61]. The interactions between DF and phenolic compounds are driven by a
combination of hydrogen bonding, van der Waals forces, electrostatic attraction, hydrophobic
contact, strong covalent bonding (esterification) or physicochemical entrapment. For the phenolic
compounds the interactions are shaped by polymerization degree, flexibility of the molecule and
number of reactive groups (hydroxyl, glycosyl, galloyl etc). DF interactions are predominatly of
electrostatic and ionic nature [8]. Therefore, the DF-phenolic compounds interactions depend on
the type of DF and phenolic compounds. Moreover, when considering the physicochemical
entrapment of DF the particle size, porosity and surface characteristics will influence the amount
of entrapted phenolic compounds [62].
In the case of polyphenols found in apple the interactions are of ionic nature with pectins and
depend on the phenolic compounds molecular weight [63]. As for the interactions of bacterial
pectin-cellulose composites with anthocyanins and phenolic acids (PA) a two-steps process of
binding was found in an in vitro assay. In the first step, anthocyanins bound at neutral pH, the
binding being slightly higher for non-acetylated compared to acetylated anthocyanins, while in the
second step unspecific stacking of anthocyanins on pectin-cellulose composites occured slowly.
PA were binding via non-covalent interactions: to cellulose through hydrophobic contact, while to
pectins through H-binding and electrostatic interaction [64].
The binding between cellulose and phenolic compounds depends on the pH values and the
concentration of phenolic compounds [65].
Regarding the binding between phenolic compounds and β-glucans it has been shown that the
interactions mainly involve H-bonding [66, 67] and they depend on the phenolic compounds
structure, namely on the level of hydroxylation, glycosylation, respectively galloylation of the
phenolic compounds. Lo Piparo [68] showed that the binding of flavonoids to β-glucans is
positively correlated with flavonoids hydroxylation (being optimal at three hydroxyl groups and
having a decrease in binding with four or more hydroxyl groups), that glycosylation generally
limits binding (except for myricetin and daidzein) and that in the case of catechins, galloylation
increases the binding.
Taking into consideration that mainly the interactions between DF and PA are weak (non-
covalent), their formation and dissociation kinetics will respond to changes of conditions like pH
value and solvent quality [69] phenolic compounds begin to be released from the food matrix in
the upper part of the gastrointestinal tract either by direct solubilization at physiological conditions
(37 ℃, pH range between 1 and 7.5) or by enzymatic hydrolysis of the nutrients to which the
phenolic compounds are bound. However, an important amount of phenolic compounds pass
through the upper intestine undissociated from the DF. Binding to DF modulates phenolic
compounds bioavailability which has important physiological significance: the phenolic
compounds that escaped absorption in the small intestine will be released in the colon, whilst labile
phenolic compounds will be protected from the very acidic conditions in the upper part of the
gastrointestinal tract [60, 70] [71].
In consequence, DF has the role of being a PA carrier contributing to fiber-rich diet health benefits.
Thus, DF was used for the entrapment or encapsulation of phenolic compounds, for example
chitosan-entrapment of green tea polyphenols, nanoparticles composed of arabic gum and
maltodextrin with improved EGCG stability, guar gum matrix and pectin-
hydroxypropylmethylcellulose tablets for quercetin incorporation, etc [8].

4.2.Binding with bile salts (bile acids)

Bile acids play an important role in lipid metabolism [72]. Bile acids are synthesized in the liver
from cholesterol and are secreted via the bile in the small intestine. Most of the bile acids are
reabsorbed via the enterohepatic circulation by active and passive mechanisms in the ileum [73],
while the unabsorbed ones are excreted in the feces [74]. The deficit of bile acids is compensated
by de novo synthesis in the liver, therefore presumably leading to reduced cholesterol levels [75].
The soluble and insoluble DF have different bile acids binding properties [76]. Gunness [77]
reported that DF interact with the bile acids impeding their reabsorption. Kahlon [78] evaluated
the in vitro bile acids binding of various raw and after different cooking of vegetables. High bile
acids binding values were obtained by steaming the vegetables, while the lowest ones were in the
case of boiling them. In the case of steaming, the bile acids binding values were as it follows: beets
18%, okra 16%, eggplant 14%, asparagus, collard greens, mustard greens and kale 10-13%,
cauliflower, green beans, carrots, spinach, brussels sprouts and broccoli 6-8%, green bell pepper
and cabbage 4% and turnips 1.2%. Under small intestine conditions the isolated pectins interacted
with bile acids [79]. bile acids binding capacity of several DF types, inulin, β-glucan, guar gum
and Konjac glucomannan (KGM), was significantly improved by ultrafine grinding [80]. Bile
acids binding to DF was demonstrated by in vivo and in vitro [81] studies, respectively clinical
trials. The interactions between DF and bile acids are influenced by particle size, its surface,
viscosity, pH and temperature of the media.
The DF interaction with bile acids seems to be a complex process involving hydrophobic and
hydrophilic effects. Moreover, polyphenols were shown to reduce bile acids solubility during
digestion. Ogawa [82] was able to show by using NMR that bile acids and tea polyphenols were
in close proximity interacting by H-bonding, which could potentially influence the solubility and
enterohepatic recirculation of bile acids.
It is important that not all bound bile acids are excreted from the body. A part of the DF is
fermented by the microflora and during this process the bound bile acids are released and
enzymatically converted. Despite changed physicochemical properties, a part of the bile acids may
be reabsorbed even from the large intestin [83].

4.3.Binding with mineral ions (MI)

 • The binding capacity of DF with mineral ions was intensively studied using in vitro and
in vivo assays and was recently reviewed [83]. It was shown that diverse factors may
influence the mineral bioavailability:
• the amount of different chelators (as phytate, tannins, citrate and amino acids);
• the mineral concentration, size, and valency;
• the potential association of phytic acid with proteins; the heat treatment that could affect
the binding;
• the pH value;
• the presence of other metal ions that may compete and the fermentability of the fiber
components in the colon that could lead to a potential uptake of the minerals in the gut
etc.

Especially the last factor is of interest as there are discrepancies between in vitro and in vivo studies
of DF-mineral ions binding. Davies studied both in vivo and in vitro the amount of zinc bound to
purified pectin and found that up to 86% of the zinc was bound in vitro, nonetheless no effect on
zinc availability was observed in vivo. The different in vivo and in vitro results may be related to
absorption of released minerals from the DF binding following fermentation of DF in the large
intestine [84].
The interactions between DF and MI are driven mainly by electrostatic attraction or complex
formation involving the carboxyl, hydroxyl and amino groups of DF as main functional groups.
Nair [85] reported that highly metoxylated pectines bind less MI than the low metoxylated pectines
due to the number of available carboxilic acid moieties. The electrostatic interactions between DF
and MI involve mainly divalent cations [86]. Pectins, xanthan gum and carragenans bind a higher
amount of divalent cations compared to agar and guar gum [87]. Reinhold [88] reported that FeII
and FeIII retention takes place due to the formation of poorly soluble fiber-mineral element
polymers.
Nevertheless, ionic strength has to be taken into account as the ion exchange capacity will be low
in the acidic conditions of the stomach while it will be high in the neutral conditions of the small
intestine. For example, a study of Schlemmer revealed that pectins and gums do not bind divalent
cations at ionic strength of around 0.1.
The interaction between DF and MI leads to various consequences, the most important one being
related to the bioavailability of MI. An increased calcium absorption was reported when adolescent
girls were supplemented with inulin [89]. These findings were supported by a study of Abrams[90]
on pubertal girls and boys supplement with inulin.

4.4.Binding with digestive enzymes

Ji [91] evaluated the α-amylase and glucoamylase activity inhibition by cellulose nanocrystals
(CNCs) using visible absorption spectroscopy, Fourier transformed infrared spectroscopy,
fluorescence quenching method and circular dichroism which showed an interaction between the
tested enzymes and CNCs involving changes in the secondary structure of the enzymes. α-amylase
is also inhibited by guar gum [32]. A DF found in brown algae, called fucoidan, has an inhibitory
effect on α-amylase and β-glucosidase [92]. Hansen [93] reported that lignin could totally inhibit
the activity of isolated trypsin and chymotrypsin, having no effect on enterokinase. The inhibition
effect depends on incubation time, fiber concentration and enzyme level.
In vitro studies and tests on rats showed that pectin has an inhibitory effect on pancreatic lipase
and trypsin [94]. The lipase catalytic activity inhibition could be explained by protonation of the
enzyme active site by the pectin carboxylic groups [93]. The same mechanism of inhibition was
proposed in the case of alginate which can also inhibit pancreatic lipase. Alginate has an inhibitory
effect on pepsin due to electrostatic interaction between the negative net electric charge of alginate
and the positive net electric charge of pepsin in the acidic conditions of the stomach.
It should be taken into account that some of the digestive enzyme inhibitory effect observed for
DF is actually related to the phenolic compounds carried and released under certain conditions in
the grastrointestinal tract. phenolic compounds were reported to posses inhibitory activity towards
enzymes such as amylase, glucosidase, pepsin, trypsin and lipases and the subject has been studied
extensively [95]. The inhibition of amylase and glucosidase is leading to a reduction in post-
prandial hyperglycemia, whilst the synergy between DF and phenolic compounds could have an
important role to play in the amylase activity inhibition and therefore to contribute to the
management of type II Diabetes in humans. A recent study of Perez [96] evaluated the biological
properties of an ethnic food common in Latin America, Prosopis nigra, and reported that the DF-
bound phenolics have a potential of lipase activity control.
 5. Dietary fiber fermentation in the large intestine and the corresponding effect on
microbiota composition

DF was defined as edible parts of the plants or analogous carbohydrates which have the capacity
to resist human digestion in the small intestine and can be completely or partially fermented by the
microbiome found in the large intestine. The fermentation depends on numerous structural and
physicochemical parameters. The fermentation process was associated with lower colonic pH
values, increased bacterial mass in the large intestine, a reduced number of harmful bacterial
species, vitamins and antioxidant compounds production, immune system stimulation etc.
Generally, insoluble DF (cellulose, lignin) is poorly fermentable by the large intestine microbiome
due to lack of water retention capacity. Soluble DF (pectin, guar gum, xanthan gum, β-glucan,
inulin, oligosaccharides) is fermentable and represents the main source of energy for the
microbiome. Both fermentable and non-fermentable DF have health beneficial effects. The non-
fermentable DF is lowering the transit time, is increasing the luminal content and has a dilution
effect on the toxic and carcinogenic compounds. An increased soluble DF consumption is
correlated with an increased microbiome development in the large intestine [1]. The fermentation
of DF by the microbiome leads to production of short chain fatty acids (acetate, propionate,
butyrate), amines, phenols, ammonia, water and gases, bacterial mass growth and energy release
[60].
The human intestinal microbiota is comprising trilions of bacterias which play an important role
in the development and function of diverse physiological processes. The microbiome is composed
of more than 400 different bacterial species which possess more than 150 times more genes, thus
more catalytic enzymes, compared to the human genome [97, 98]. The key factor influencing the
human intestinal microbiota is the nutrition, especially during infancy but also throughout life. A
dietary pattern rich in saturated fat and simple carbohydrates, low in DF, was associated with
incresed risk of developing cardiovascular diseases, colorectal cancer, obesity and diabetes [99,
100].
Some DF (inulin and other oligofructoses) have prebiotic properties which lead to increased
colonization of beneficial bacterial species like Bifidobacteria and Lactobacillus in the large
intestine which suppress the pathogenic bacterias [101, 102]. These bacterial species can produce
short chain fatty acids and stimulate the immune system. The DF with prebiotic effect are of great
importance for the infants as studies suggest that when supplemented with prebiotic fiber mixture
the postnatal immune system development was promoted, the bowel function was improved and
atopic dermatitis or respiratory infections were reduced [103]. The DF with prebiotic effect was
reported to have anxiolytic, [104] antidepressant and cognitive effects also [105-109].
Inulin inhibits the growth of harmful bacteria like Salmonella, E.coli and Listeria [110]. Pectin
was reported to reduce acute intestinal infection and to slow down diarrhea in infants and children
[111]. Guar gum was shown to be fermented in the human colon by the microbiota inducing
beneficial effects like improved bowel function and constipation relief [112]. Pectin is releasing
mainly acetate in the colon which is presumed to enter the peripheral circulation leading to altered
fibrin quality, a risk factor involved in cardiovascular diseases [113].
Short chain fatty acid productions is leading to low colonic pH values which improves the mineral
ions solubility and therefore facilitates their absorption [114, 115]. Furthermore, the low pH
inhibits the growth of potentially harmful bacteria (E.coli, Bacteroidetes) and favours the growth
of butyrate-producing Firmicutes [116].
The digestion and absorption of phenolic compounds associated with DF is not complete in the
small intestine, thus a part enters the large intestine. Perez-Jimenez [117] showed that the
procyanidin content in diet is understimated as the non-extractable phenolic compounds (phenolic
compounds bound to DF) measurement is crucial in assesing the dietary intake of phenolic
compounds. phenolic compounds are released from the DF by the bacterial enzymes and can be
metabolized, thus a major part of the DF associated phenolic compounds (a percentage of 42% of
the dietary phenolic compounds) becomes bioaccesible in the large intestine [60]. The unabsorbed
phenolic compounds and their fermentation metabolites (urolithins, phenylacetic, phenylpropionic
and phenylbutiric acids) contribute to an antioxidant environment in the large intestine [60, 118,
119]. This free radicals scavenging could be involved in the protection against cancer in the human
colon [120]. For example, Janicke [121] studied the effect of hydroxycinnamic acids on colonic
cancer epithelial cells (Caco-2 cells) using global gene expression analysis of mRNA with focus
on the genes which are responsible for cell proliferation and cycle progression and reported that
different functional groups and pathways were indeed affected, the S phase was delayed and genes
regulating centrosome assembly and S phase DNA damage checkpoints were affected. Also,
Lizarraga [122] studied the effect of grape DF on mice and reported the downregulation of
pathways associated with cancer and obesity like nuclear receptor signaling, lipid biosynthesis and
energy metabolism and the upregulation of antioxidant and detoxification enzymes, apoptotic,
immune system and tumor supression genes. Therefore, a new concept was introduced, the
„antioxidant DF”, which is defined as DF containing an important amount of associated natural
antioxidant compounds [123]. Additionally, the DF associated phenolic compounds have positive
effects on lipid metabolism.

6. Conclusions

Nowdays is a general consensus that a diet rich in DF have a health effect by preventing against
diseases. Compared to other nutrients the beneficial health effect of DFs depends on the
modulation of digestion rather than the nutritional characteristics or the triggering of specific
biological activities. In this review, the behavior of DF during food digestion has been discussed
and correlated to the physiological effects on health. Despite acctual knowledge of DF’s behavior
in the GI tract due to the last year realistic and reliable in vitro digestion models, there are still
several research directions for the future studies.
The increasing prevalence of non-communicable diseases with an inflammatory cause has led to
the understanding of the gut microbiota as an intrinsec regulator of host immune responses. Diet
as a microbiota-modulating factor by the content and composition of DF hence a health-
modulating factor. A revision of the current nutritional guidelines considering both the molecular
content and molecular structure of nutrients for sensitive persons (e.g., insulin resistant and insulin
sensitive individuals, etc.), could be valuable for the restoration of beneficial bacteria and
microbiota diversity, for the general population.

Author Contributions: Conceptualization, R.S.; Writing-Original Draft Preparation, O.L.P.,

R.A.V., C.I.M., A.V.S.T., and A.C.; Writing-Review & Editing, R.S., R.V., C.C.M., S.C.H.,
O.L.P., C.I.M.; Vizualization: R.A.V., R.V., C.C.M; Supervision: R.S., C.A.I., A.F.
Funding: This work was supported by the UEFISCDI, PN research grants PN-III-P2-2.1-CI-
2018-1367, PN-III-P2-2.1-CI-2018-1569, PN-III-P2-2.1-CI-2018-1134.

Acknowledgements: We are grateful to Rosita Gabbianelli, PhD, for her editorial assisstance.
Conflicts of Interest: The authors declare no conflict of interest.

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