REVIEW

published: 06 July 2020

doi: 10.3389/fnut.2020.00075

Saccharide Characteristics and Their

Potential Health Effects in
Perspective
Fred Brouns*

Department of Human Biology, Faculty of Health, Medicine and Life Sciences, School of Nutrition and Translational Research
in Metabolism, Maastricht University, Maastricht, Netherlands

To understand the effects of saccharides on our metabolism and health, we need a

clear understanding of what they are, how they differ, and why some types are deemed
“less healthy” and others “better for health.” There are various ways to look at this topic.
Firstly, saccharides can be classified according to their degree of polymerization (DP). This
classification is useful when qualitative or quantitative analysis and calculation of intakes
are required or for food-labeling definitions. However, it does not account for the fact that
saccharides with a similar DP can differ in molecular composition, which will influence
digestion, absorption, and metabolism. Secondly, another approach widely used in
the biomedical and nutritional sciences is therefore a physiological classification, which
Edited by: addresses the rate and degree of digestibility and absorption, the glycemic response,
Jennie Cecile Brand-Miller,
University of Sydney, Australia and the metabolic fate. The individual health status also plays a role in this respect.
Reviewed by: An active, lean person will have a metabolic response that differs from an inactive
Javier Gonzalez, person with overweight and insulin resistance. However, this approach will not give a
University of Bath, United Kingdom
Sion Adam Parry,
complete answer either because the characteristics of the matrix/meal in which these
University of Oxford, United Kingdom carbohydrates (CHOs) are present will also influence the responses of our body. Thirdly,
*Correspondence: one can also rank CHOs by comparing their functional/technological properties, such
Fred Brouns as relative sweetness, viscosity, and solubility. Understanding CHO characteristics and
[email protected]
related physiological responses will help understand health and disease implications.
Specialty section: Therefore, a brief outline of different carbohydrate classifications is presented. This outline
This article was submitted to will be placed in the context of potential overall effects after consumption. The answer
Nutrition and Metabolism,
a section of the journal
to the question whether we should we eat less of certain sugars depends on the angle
Frontiers in Nutrition from which you look at this matter; for example, do you address this question from a
Received: 07 March 2020 single molecular characteristic point of view or from a meal quality perspective? Looking
Accepted: 01 May 2020
at one particular CHO characteristic will almost always lead to a different conclusion (e.g.,
Published: 06 July 2020
the labeling of fructose as toxic) than evaluating from a “total perspective” (fructose has
Citation:
Brouns F (2020) Saccharide adverse effects in certain conditions). Examples are given to help understand this matter
Characteristics and Their Potential for the benefit of justified dietary/food-based recommendations.
Health Effects in Perspective.
Front. Nutr. 7:75. Keywords: saccharide-characteristics, carbohydrate-classification, added sugars, free sugars,
doi: 10.3389/fnut.2020.00075 sugar-functionality, glycemic index, sugars and health

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Brouns Saccharide Characteristics and Health

INTRODUCTION TABLE 1 | Chemical classification of carbohydrates (9–11) *Maltodextrins are an

industrially hydrolyzed starch product.
Sugars and other carbohydrates (CHOs) have many
Classification Sub-group Examples
characteristics, ranging from molecular composition to
functional, physiological, and biochemical behavior. As any Sugars • Monosaccharides • Glucose, fructose galactose,
individual characteristic of a given CHO can influence its (DP 1-2) • Disaccharides mannose, arabinose, xylose,
physiological properties, it should be viewed in the context • Sugars alcohols/polyols erythrose, and others.
of all other characteristics. For example, different sugars can • Sucrose, isomaltulose, lactose,
maltose, trehalose, and others.
be similar with respect to their monomer composition but • Sorbitol, mannitol, lactitol,
may differ in the bonds between these constituents. Ingesting xylitol, erythritol
sucrose, which delivers the monomers glucose and fructose Oligosaccharides • Maltodextrins* • *Contain: glucose, maltose
for absorption, can lead to different gastrointestinal and (DP 3-9) (Malto-oligosaccharides) gluco-oligosaccharides
post-absorptive effects compared with ingesting glucose or • Non-digestible • Raffinose, stachyose,
fructose as a single source. In a solid, liquid, or viscous matrix, oligosaccharides fructo-oligosaccharides (FOS),
• Starch arabino-oligosaccharides
the same sugars will show different physiological responses.
(AXOS), and others.
For this reason, it needs to be acknowledged that looking • Amylose, amylopectin, and
at one particular CHO characteristic will almost always lead modified starches.
to a different conclusion about potential health effects than Polysaccharides • Non-starch • Pectin, cellulose, hemicellulose,
looking from a “total” perspective as regards the effect of (DP >9) polysaccharides (NSP) hydrocolloids (Arabic gum,
the carbohydrate in a certain meal/pattern and lifestyle. A • Resistant starch (RS) guar gum, others).
• RS type 1,2,3, and 4
consequence may be that misinterpretations and misconceptions
are being created by interpreting the effects of saccharides During hydrolysis, a mixture of gluco-oligosaccharides, maltose, and glucose is formed.
on health in a strongly reductionistic way (1–4). In this light, The quantity of glucose and maltose present in “maltodextrins” depends on the extent of
hydrolysis (rate x time).
the reader will be provided with condensed information on
specific compositional characteristics of CHOs, especially sugars,
which have physiological and metabolic effects. The individual
characteristics will be discussed in the context of what they
Same Degree of Polymerization but
mean for the potential overall effects on health and disease, and
why food authorities are shifting to more qualitative food-based Different Effects
guidelines (5–8). When present in disaccharides, the bonds of the composing
monomers (α or β glycosidic bond) can differ, which will
affect the rate of digestion and absorption. In Table 2,
the chemical classifications and molecular characteristics of
CHEMICAL CLASSIFICATION OF selected CHOs (types, bonds) are given, along with some
SACCHARIDES AND ITS MEANING selected characteristics of digestion, absorption, distribution, and
metabolic fate.
Saccharides can be ranked according to the characteristics To explain how CHOs with a similar monomer composition
of their molecular composition. This ranking includes can differ in their degree of digestion and absorption, we will give
individual monomers (monosaccharides) and the number two examples: [1] sucrose and isomaltulose, and [2] amylose and
of bonds. For example, sucrose is comprised of two amylopectin starch.
monomers, glucose and fructose, which are linked by an
α1,2 glycosidic bond, having a degree of polymerisation 1) The disaccharides sucrose and isomaltulose are both
(DP) of 2. Chemical classifications that are commonly composed of the two monomers glucose and fructose.
used by nutrition and food safety authorities (9) are However, the linkage between the two monomers differs.
as follows: Sucrose has an α-1,2 bond, whereas isomaltulose has an
α-1,6 bond (see Figure 1). Due to its more stable α-1,6
1) Sugars (monosaccharides and disaccharides, DP 1–2) glycosidic bond, hydrolysis by small intestinal disaccharidases
2) Oligosaccharides (DP 3–9) is slow. In human small intestinal mucosa homogenates
3) Polysaccharides (DP ≥ 10). as an enzyme source, the hydrolysis rate was 26–45%
Within these categories, dietary CHOs can be further compared with sucrose (21). The result is a lower glycemic
subclassified as presented in Table 1 below. and insulinemic response (22), and consequently a reduced
Dietary CHOs can be further subclassified as presented in rate of metabolism (23).
Table 1 below. 2) One may wonder why the example of amylose and
amylopectin starch is being discussed alongside sugars. The
reason for including the example of starch is that sugars
Abbreviations: AGEs, Advanced glycosylation end-products; CHO, carbohydrate;
CHOs: carbohydrates; DP, degree of polymerization; GI, glycemic index;
deliver their constituent monomers to the intestinal cells for
MRPs, Maillard reaction products; SCFAs, short chain fatty acids; SSBs, sugar absorption as a digestive fate. In the light of the generally
sweetened beverages. accepted definition that sugars are all CHOs with a DP

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TABLE 2 | Chemical and physiological characteristics of sugars and other glycemic carbohydrates.

CHO Type Digestive In gut Enterocyte In blood Possible metabolic fate options GI
enzyme lumen uptake

Glucose Monosaccharide - Glucose - Glucose Used as fuel, stored as glycogen and/or 100
converted to other metabolites
Fructose Monosaccharide - Fructose - Lactate, glucose, Partially converted to lactic acid and 19
fructose glucose, used as fuel or stored as
glycogen, and fatty acids used as fuel or
triacylglycerol stored as lipid
Sucrose Disaccharide: glucose -fructose, Sucrase Glucose, Glucose, glucose, lactate, see glucose and fructose above 65
α1-2 bond fructose fructose fructose
Isomaltulose Disaccharide: glucose -fructose, Isomaltase Glucose, Glucose, See fate of glucose and fructose above 32
α1-6 bond fructose fructose
Galactose Monosaccharide - Galactose - Galactose Liver conversion to glucose, see fate of 25
glucose above
Lactose Disaccharide: glucose -galactose, Lactase Glucose, Glucose, Glucose, See fate of glucose and galactose above 45
α1-4 bond galactose galactose galactose
Honey Glucose 30.3%, fructose 38.4%, Sucrase Glucose, Glucose, Glucose, lactate, See glucose and fructose above 50
sucrose 1.3% fructose fructose fructose
Maple syrup Sucrose 98%, glucose 1%, Sucrase Glucose, Glucose, Glucose, lactate, See glucose and fructose above 54
fructose 1% fructose fructose fructose
HFCS 55 Fructose 55%, glucose, 43% α-Dextrinase Glucose, Glucose, Glucose, lactate, See glucose and fructose above 58
gluco-oligo saccharides 3% fructose fructose fructose
Starch Glucose polymers: amylopectin Amylase from Maltose, Maltose, Glucose See fate of glucose above 40–110*
α1-4 and α1-6 bonds. Amylose saliva, glucose glucose
α1-4 bonds pancreas
Maltodextrins Glucose polymer, α1-4 glycosidic α -Dextrinase Glucose, Maltose, Glucose See glucose above 110
bonds maltose glucose
Maltose Disaccharide: glucose-glucose, Maltase Glucose Glucose Glucose See glucose above 105
α1-4 glycosidic bond
Trehalose Disaccharide: glucose-glucose, Trehalase Glucose Glucose Glucose See glucose above 70
α1-1 glycosidic bond
Sorbitol* Sugar alcohol - Sorbitol - Sorbitol Liver conversion to fructose and glucose, 4
see above

For a review of fructose, see Tappy and Lê (12). For a review of lactose and galactose, see University of Waterloo (13). One example of a low-caloric/low-glycemic sugar replacer is
given. In the gut, sorbitol, a sugar-alcohol, is slowly absorbed (25–80% of the consumed dose) by facilitated diffusion. Absorbed sorbitol passes the liver, where it is converted to fructose
and glucose (14). The unabsorbed fraction is transported to the large bowel, where it is fermented. When sorbitol is consumed in high doses, potential side effects can occur as a result
of osmotic water shifts from blood into the gut, resulting in rumbling, loose stools, or diarrhea (extensive details about polyols can be found in Livesey (14), Ghosh and Sudha (15),
Rice et al. (16). For a review of low- and non-caloric/non-glycemic sweeteners compared with caloric sweeteners, see Rogers et al. (17). *The glycemic index of starchy foods varies
according to the molecular content of amylose, amylopectin, fiber, presence of protein, and characteristics of the food matrix, resulting in a range of reported values. For extensive
glycemic index data [see (18)], International Tables of Glycemic Index and Glycemic Load Values, the online University of Sydney searchable data GI; http://www.glycemicindex.com/
foodSearch.php. For further extensive details, see Queen Mary University London (19), Nomenclature of Carbohydrates (Available online at: https://www.qmul.ac.uk/sbcs/iupac/2carb/
00n01.html#0121) and nomenclature of sugar alcohols (20).

of 1–2, it becomes clear that both “sugar” and “starch” α-amylase is responsible for the breakdown of the starch into
deliver “sugars” to the intestinal cells for absorption. In dextrins (maltotriose, DP3) and maltose (DP2), which are
terms of metabolic responses, especially when comparing in turn digested by epithelial maltase, resulting in glucose
“sugars” with “starches,” it is good to have a clear comparative monomers. It is often suggested that the amylose content is
view. Plant starch generally contains 20–30% by weight of the most important factor in determining the rate of digestion
amylose and 70–80% by weight of amylopectin. Amylose and absorption as well as the related glycemic response, but
(Figure 2A) contains linear chains of approximately 300– recent research shows that the picture is more complex (25).
3,000 glucose monomers in length, connected by α-1,4 bonds. It appears that the interaction between the molecular and
In amylopectin, there is also a linear basic structure in which granular structure (helix formation, number of pores, size of
glucose monomers are linked by α-1,4 bonds (Figures 2B,C), the molecule, amylopectin sidechain length distribution and
but there are side branches along this linear base initiated with crystalline structure, the latter two being the most important)
α-1,6 bonds. This situation results in a molecule with many causes the variation in the rate of digestion across botanical
branching endpoints and a more open structure in which sources (25). The latter leads to relatively rapid digestion
digestion enzymes can act, compared with the more closed and a significant blood glucose response. The potential of
linear helix formation of amylose. The digestive enzyme starch to affect the blood glucose response, expressed as a

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FIGURE 1 | Molecular structure of isomaltulose. By using the microbial enzyme “glucosyl transferase” for rearranging the bond structure from α1-2 in sucrose, as
base substrate, to α1-6, isomaltulose is formed.

FIGURE 2 | Molecular structure starch: amylose starch (A), amylopectin starch (B,C) [figures source (24): starch, retrieved Jan 2020].

glycemic index (GI) value, can therefore vary considerably wide range of GI values for different varieties of rice, cereals,
depending on the content of amylopectin and amylose (26– potatoes and derived products, ranging from relatively low to
28). Interestingly, despite only small differences in amylose high GI values (18). For this reason, one cannot establish a
content, in vitro cumulative starch hydrolysis shows that generic GI value for starchy foods. These aspects are important
wheat starch is more rapidly digested than potato starch to understand for situations in which a rapid or sustained
(being the most resistant starch), with corn, high-amylose delivery of glucose to the circulation and tissues is required (e.g.,
corn, and pea starch having intermediate values (25). sports nutrition or compensation of insulin dosage-induced
hypoglycemia in diabetes patients), or generally to be avoided
Accordingly, rapidly digestible (available) CHOs, slowly (type 2 diabetes).
digestible (available) CHOs, and non-digestible (non-available) In infant nutrition, sports nutrition and sometimes in clinical
CHOs (dietary fibers) can be ranked (18, 29–31). Along similar nutrition, maltodextrins resulting from industrial enzymic starch
lines, digestible starch (glycemic) and resistant starch (not degradation are used, having a mixed content of glucose
digested, non-glycemic) are both polysaccharides composed oligosaccharides, maltose, and glucose. It is often suggested that
of glucose monomers and are both present in starchy foods, these maltodextrins are complex CHOs which result in a low
but they differ strongly in bioavailability. As a result, there is a and sustained glycemic response. However, there are no data

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to support this suggestion. In fact, the enzymic digestion of the GI value, glucose usually serves as the reference food with a
maltodextrins appears to take place at a high rate, which is also glycemic index of 100. A food portion containing an amount of
reflected by comparable post-ingestion insulin responses as well 50 g of available CHOs is ingested and the area under the blood
as oxidation rates during exercise compared with glucose [(32); glucose response curve is divided by the area resulting from the
Table 3]. ingestion of 50 g of glucose. Full details on this matter can be
found in Brouns et al. (34).
Table 3 gives some examples of the glycemic index values
GLYCEMIC INDEX CLASSIFICATION AND of foods. It is important to understand that the glycemic
ITS MEANING index value in isolation cannot fully explain the physiological
impact of CHO-based foods and beverages on health and
The potential of CHOs to raise the level of blood glucose is often
disease. For example, the ingestion of 5 grams of glucose
expressed as a glycemic index (GI) value. A high value refers to a will not induce measurable hyperglycemia, despite its high
strong elevation of blood glucose and is often seen as less healthy, GI value of 100. However, the ingestion of 50 grams will
whereas a low value is often seen as beneficial. When determining increase blood glucose very significantly. Thus, any GI value
should be interpreted in the light of the quantity ingested.
For this reason, the concept of the “glycemic load” of CHO-
TABLE 3 | The glycemic index value of the plain carbohydrate tested vs. glucose containing meals has been defined as a relevant approach. In
as reference-control.
addition, it needs to be noted that the GI value of any food
Glucose GI−100 prepared using these CHOs as a meal component is highly
French baguette GI−95 influenced by other factors that affect the rate of ingestion as
French fries GI−75 well as the subsequent transit, digestion and absorption, see
Fructose, mean of three studies GI−15 Figure 3. Examples are the content of enzyme inhibitors (e.g.,
Macaroni, white boiled, mean of three studies GI−50 α-amylase inhibitors) present in the CHO source, the overall
Potato boiled, mean of seven studies GI−53 macronutrient composition (quantity and type of CHO, fat,
Ripe banana, mean of nine studies GI−48 protein), the content and characteristics of dietary fibers (e.g.,
Sourdough rye bread GI−53 soluble, viscous, insoluble, bulking), the level of processing (e.g.,
Spaghetti, white boiled. Mean of eight studies GI−41 level of refinement, such as the separation of bran and germ
Sweet potato GI−61 during milling, resulting in “refined” white flour), as well as the
Sucrose GI−67 matrix effects (e.g., liquid vs. solid, starch in a compact elastic
White rice, mean of eight studies GI−59 spaghetti structure vs. starch in a well-cooked soft potato). In
White wheat bread, mean of seven studies GI−70 the case of drinks, energy content and osmolality are factors
Whole grain rye bread, mean of four studies GI−58 which can significantly affect the gastric emptying rate as well as
Data Source: Atkinson et al. (18) and University of Sidney (33) online searchable data GI, the related supply to the gut for absorption, depending on the
International Tables of Glycemic Index and Glycemic Load Values. concentration (35).

FIGURE 3 | Factors that play a role in gastrointestinal transit, digestion, and absorption of saccharides.

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There is still one other point that needs to be addressed, For this reason, it is important to have a basic understanding of
especially related to sugars. The GI value of fructose (27) is very the flow: type of carbohydrate → molecular characteristics →
low and that of sucrose (36) is moderate. Thus, in terms of the physiological aspects (digestion, absorption, and metabolic fate)
viewpoint that a low to moderate GI is beneficial for health, one → effects on health.
might conclude that fructose and sucrose are preferable for health To give an example, oral glucose appears in blood as
to starches that have a much higher GI value. Based on current glucose and drives glycemia in a 1:1 ratio depending on the
knowledge, this point is hard to substantiate. The suggestion dose. Fructose, however, behaves differently because of its
that fructose is a single cause of non-alcoholic fatty liver disease conversion to other metabolites and because of its very low
driven by its dietary intake cannot be justified either based on insulinemic response (47). Although glucose and fructose are
data from excessive consumption (37). very often compared as monomers in metabolic studies, it
The view that sugars added to beverages are a causal factor needs to be addressed that humans usually do not consume
for obesity and diabetes is well-documented, because they cause fructose in isolation but almost always in combination with
a positive energy balance. However, in the case of sugars added glucose, as it is present in sucrose- and HFCS-sweetened
to solid foods such as confectionery, this causal association beverages, fruit juices, fruit syrups (see Figure 5), and fruits.
has not been shown (2, 38–40). Data showing that two thirds Accordingly, the interpretation of data derived from studies
of added sugars are being consumed in solid foods and only in which fructose was supplied as monomer in high amounts
one third in beverages (41) raise questions about other factors should be seen in the light that this does not represent
that may play a role in addition to sugar (2, 3). From the to normal human consumption situation. Concerns that all
above, it is clear that a focus on single CHO types, single CHO fructose from consumed SSBs and fruit juices goes straight
characteristics, or consumption in isolation as a single supply to the liver where it is all converted to lipid are not
source has limited generalizability, especially when one wants supported by evidence. In contrast, most fructose is converted to
to understand the overall effects of the diet containing these non-lipid substrates.
CHOs on postprandial appetite regulation, glycemia, lipidemia, Recently Jang et al. (48) (Figure 4) performed double labeling
low-grade inflammatory potential and possible health outcomes studies allowing for quantitatively tracing the metabolic fate of
(42, 43). Moreover, the physiological status of the person in fructose vs. glucose after supply to the mice. These researchers
question also plays a significant role in how the human body gave fructose together with glucose at 1:1 ratio, as normally is the
manages the metabolism of saccharides. Elite endurance athletes case in human consumption of fructose containing saccharide
such as professional cyclists ingest large amounts of refined sources. It needs to be noticed though, that for this work in
carbohydrates, to a large extent in beverages, to maintain a mice, oral gavage by which the test dose was directly given
high glucose availability for the benefit of delaying fatigue into the stomach, was used. Using this procedure a large
and maintaining a high-performance capacity. They burn the amount of fructose reaches the small intestine with much faster
calories ingested, even when these exceed 6,500 kcal/day for 21 kinetics than typical human fructose consumption. However,
days (44), and accordingly do not become develop overweight. while mouse metabolism is ∼10× faster than humans, rendering
Based on these and other observations, their metabolism of the the faster fructose dose to metabolic rate ratio similar between
carbohydrates and the interrelationship with lipid metabolism the species (Jang, 2020 personal communication). Using this
will be quite different from that of inactive overweight procedure, it was shown that a large fraction of the fructose
individuals who are insulin-resistant or who suffer from type 2 absorbed in the small intestine is converted to glucose and
diabetes when they consume large quantities of sugar-sweetened organic acids within the enterocytes to such an extent that only
beverages [e.g., (45, 46)]. In this respect, it is obvious that very little fructose spills over to the liver. Thus, instead of the
specific food-based dietary guidelines are required for certain common perception that the liver is the prime fructose clearing-
population subgroups. organ, it appears that small intestine fulfills this role. In case
an acute high-dose of fructose saturates intestinal absorption
and metabolic conversion capacity, a fraction on non-absorbed
LEGAL AND WHO DEFINITION OF “ADDED fructose partly passes from the small intestine to the colon,
SUGARS” AND “FREE SUGARS” to be subsequently fermented by the microbiota giving rise to
short chain fatty acids, mostly acetate, which will be absorbed
With respect to the classification and labeling of food and and passed on the liver. The fraction of fructose that escapes
beverages, one should note that the term “sugars” on the food metabolic conversion by the enterocytes also passes on the
label generally stands for “monosaccharides and disaccharides.” liver. Both acetate and fructose entering the liver can serve
In this respect, glucose and fructose are both simple sugars, but as a substrate for de novo triacylglycerol synthesis. The latter,
they behave very differently with regard to their metabolic effects. however, remains relatively small, even in a situation of acute
The hormonal responses that they induce (glucose is a significant very high doses of fructose. Studies using stable isotopes in
driver of glycemia and insulin secretion, while fructose only has humans (1, 52) showed that the 3–6 h after ingestion high doses of
very minor effects on glycemia and insulin) and their metabolic fructose only a small percentage (<1%- max 3%) was converted
fate, which includes the conversion to other intermediates such as to fatty acids. Thus, previous human work is in line with the
organic acids (in particular lactic acid) as well as fatty acids, their new insights obtained by Jang et al. (48). Future studies in
use as fuel and their possible storage as glycogen or lipids differ. humans need to verify how much fructose, at real-life intake

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FIGURE 4 | Metabolic fate of oral fructose. When ingesting small doses of fructose (F) and glucose together, as in human nutrition, most absorbed F is converted to
glucose, lactic acid and other organic acids within the enterocytes, which appear in the portal vein supplying the liver. The amount of F passing to the liver after small
oral doses is negligible. Glucose largely passes the liver and enters the blood circulation to be available to all tissues. Lactate will favorably be converted into liver
glycogen. Non-converted lactate will pass on to the blood circulation. After ingesting acute large doses (≥1 g/kg body weight, equivalent to >1 liter of
sugar-sweetened beverage/juice), F partly escapes its own slow absorption process and will pass on to the colon, where it may cause osmotic fluid shifts potentially
leading to laxation and will be fermented by the microbiota leading to the formation of short-chain fatty acids, mostly acetate, which will be absorbed and pass on to
the liver with portal blood. In this situation, the absorbed but non-converted fraction of F will serve as substrate for de novo fatty acids synthesis, along with the
acetate coming from the colon. As a result of the above, F enters the circulation only in very small quantities. (Based on data from (12, 48–53)]. Figure based on data
from Jang et al. (48) and Zhao et al. (53).

levels (concerning dose-time interrelationships generally much acetate, propionate, and butyrate, generally in a molar range
lower than experimental supply levels), really passes on the liver of 70:20:10%, respectively. Individuals who consume relatively
and the colon and what the conversion rate is to liver fat. To high amounts of dietary fiber such as fruit fibers and fructans
put this in perspective, early human studies, using the ileostomy (inulin) generally suffer less from being overweight. Why would
model or breath hydrogen as marker of malabsorption, showed fiber derived acetate, compared to fructose derived acetate, not or
very clearly that fructose ingested as monomer at doses of >25 g differently contribute to fatty liver? Is there a protecting role from
induces malabsorption. However, when co-ingested with glucose propionate? (60). And, Why do physically active lean individuals,
(such as isomaltulose or sucrose)— even up to acute doses of up who consume substantial amounts of sugar, not suffer from an
to 100 g sucrose (equivalent to about 1 l of SSB or fruit juice)— overweight and fatty liver, whereas most overweight individuals
this is not the case (54–59). Since humans seldom consume do? Is excess calories/positive energy balance the prime
fructose in isolation, this is an important point to consider. driving factor?
In addition, it needs to be addressed what other factors, apart
from fructose contribute to the novo lipogenesis. In very recent Natural and Refined Sugars: Do They
work, it was shown that fructose fermentation derived acetate Differ?
contributes to liver lipogenesis (53). Concerning the latter, a The metabolism of isolated monosaccharides and disaccharides
range of well-fermentable dietary fibers give rise to a significant (glucose, fructose, and sucrose/table sugar) is basically similar to
amount of SCFA the cecum and colon, most importantly that present in natural sources which contain mixtures of these

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FIGURE 5 | Sugars in syrups. The sugar monomer content of sucrose (sucrose water content is subtracted from the total mass and this value is set at 100%) is
compared with high-fructose corn syrup (HFCS, containing either 42 or 55% fructose) and other types of syrups. Maple syrup consists almost entirely of sucrose
[source: Andrea et al. (61)].

sugars, such as in fruits or fruit-derived syrups. Because of their TABLE 4 | Definitions of “added sugars” and their use in governmental reports
molecular similarity and related physiological responses, sugars [Source: Buyken et al. (63)].

naturally present in honey, fruit-derived syrups (Figure 5) and “Sugars” are generally defined as “mono- and disaccharides.” Accordingly,
fruit juices have recently been proposed by the WHO (62) to be “added sugars” is mostly considered to be “added mono- and disaccharides.”
of similar nature as “commonly added sugars”. This approach • WHO report (62): introduced the term “free sugars” as “all monosaccharides
has led to a new, mutually inclusive category of “free sugars” and di-saccharides added to foods by the manufacturer, cook, or consumer,
plus sugars naturally present in honey, syrups, and fruit juices.”
and to questions about the scientific basis of the term “free
• US: United States Food and Drug Administration (US-FDA)-(64) and
sugar”. For example, why are sugars in 100% fruit juice “free United States Department of Agriculture: Added Sugars are all sugars that are
sugar” and the same sugars naturally present in the fruit not? either added during the processing of foods, or are packaged as such, and
Why is milk sugar naturally present in milk not considered to these include sugars (free, mono- and disaccharides), syrups, naturally occurring
be a free sugar? In this respect, fruit juices have been classified sugars that are isolated from a whole food and concentrated so that sugar is the
primary component (e.g., fruit juice concentrates), and other caloric sweeteners.
in many epidemiological studies together with sugar-sweetened
• UK: SACN report (39) adopted the term “free sugars from WHO,” which now
beverages (SSBs) as one category of “sugar-sweetened beverages”. replaces the terms “added sugars” and “non- milk extrinsic sugars” (NMES) used
Such a pooling of beverages and related observational data has previously. “Free sugars’ comprises all monosaccharides* and disaccharides*
resulted in the conclusion that fruit juices, similar to sucrose- added to foods by the manufacturer, cook or consumer, plus sugars naturally
sweetened drinks, are a cause of obesity. This outcome has led to present in honey, syrups and unsweetened fruit juices. Under this definition,
it includes lactose (the sugar in milk), when naturally present in milk and
international recommendations for reducing the consumption of milk products, and the sugars contained within the cellular structure of foods
“free sugar.” Table 4 gives an overview of definitions for “added (particularly fruits and vegetables) are excluded.”
sugars” used by various health authorities, as recently reviewed • EU: EFSA report (65): added sugars are “mono- and disaccharides and
by Buyken et al. (63). starch hydrolysates (e.g., glucose syrup, fructose syrup, maltodextrins) added
during food preparation and manufacturing.”
Based on the molecular similarity of sugars, the pooling
of juices and SSBs is understandable. However, data from
intervention studies do not support this assumption. Murphy
et al. (66) evaluated the effects of 100% fruit juice and measures
of glucose control as well as insulin sensitivity in a systematic endpoint values with error terms. It was concluded that the
review and meta-analysis of randomized controlled trials. In repeated intake of 100% fruit juice does not have a significant
this research, clinical trials were eligible for inclusion if the effect on glycemic control or measures of insulin resistance,
following criteria were met: [1] the trial was randomized and which is consistent with findings from some observational studies
conducted in human subjects; [2] the trial was a controlled in which the consumption of 100% fruit juice was studied
intervention providing 100% fruit juice and a control beverage separately from SSBs and in which lifestyle factors were also
(e.g., sugar/carbohydrate or energy-matched beverage, water or taken into account (2, 67). One reason may be that juice
no beverage); [3] the fruit juice consumed was identified as 100% contains a wide variety of micronutrients and plant-bioactive
fruit juice; [4] subjects consumed 100% fruit juice for a minimum substances from the original fruit, which may be “protective”
of 2 weeks; [5] outcome data for at least one measure of glucose (68). Another reason may be that individuals who decide to
control or insulin sensitivity were reported; and [6] reported consume 100% juice instead of SSBs also make other healthy
outcomes included change from baseline values or baseline and lifestyle decisions. As a result, 100% juice consumers usually have

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TABLE 5 | Some physicochemical, technological, and functional characteristics TABLE 6 | Relative sweetness of sugars.
that are important for food design and food processing.
Sugar Relative sweetness
• Sweetness
• Solubility Monosaccharides
• Viscosity Fructose 115–180*
• Reducing power
Glucose 50–70*
• Crystallinity
• Glass transition temperature Galactose 54
• Cooling effect (mouth) Disaccharides
• Melting temperature Sucrose (gluc+fruc) 100
• Freezing behavior
Maltose (gluc+gluc) 30–50*
Lactose (gluc+galac) 15–40*
Isomaltulose (gluc+fruc) 50

a more favorable body mass index [(69, 70); BMI], while the Trehalose (gluc+gluc) 45

quality of the daily diet also appears to be better, as has been HFCS–(gluc + 55% fruc) >100

observed in both children and adults (68, 71). Very recently, HFCS–(gluc + 42% fruc) 100

Khan et al. (4) challenged the classification of juices in the *Degree of sweetness is influenced by concentration and higher at higher concentrations.
same box as soda, since their consumption is associated with Gluc, glucose; fruc, fructose; galac, galactose [Source: Clemens (73)].
different health effects. This example also shows that looking
at a single sugar type or sugar characteristic in isolation is not consequence, the beverage will contain more calories! To
meaningful and may lead to wrong interpretations with respect replace 100 g of sucrose in a beverage with fructose (relative
to health. sweetness of fructose at higher concentrations ∼ = 150), 33%
Although the metabolism of CHO molecules naturally present less sugar is required. As a consequence, the drink will
in food or isolated (such as plain table sugar) is basically identical, contain less calories but also a high level of fructose, which
it is important to understand as well that the food matrix can play may cause gastrointestinal distress/diarrhea and unfavorable
a significant role in the rate of intake, digestion and absorption. metabolic effects.
The effects of sucrose added to a beverage (rapid gastric emptying B) Glucose, fructose, galactose, lactose and maltose are
and small intestinal absorption) will lead to a rapid increase reducing sugars. Sucrose and trehalose are non-reducing
in blood glucose and insulin, which differs from effects in a sugars. During the cooking/baking/roasting of food at high
solid matrix such as confectionery (lower rate of digestion and temperatures, reducing sugars react with amino acids in a
absorption as well as a less rapid increase in blood glucose Maillard reaction. This “browning reaction,” such as when
and insulin). As a consequence of its rapid gastrointestinal baking meat or bread or roasting coffee, affects the taste and
transit, sucrose in a beverage induces less satiation compared flavor. For this reason, selective use can be made of reducing
with sucrose in a solid food. This “incomplete sensing” drives sugars to obtain the desired browning and flavor (74).
“unnoticed” calorie intake, a positive energy balance and obesity, There is a wide range of Maillard reaction products (MRPs)
when happening frequently (72). known to influence digestive physiology, gut microbiota
and metabolism, which are also suspected of triggering an
PHYSICOCHEMICAL, TECHNOLOGICAL, immune reaction to and the allergenic potential of proteins
AND FUNCTIONAL CHARACTERISTICS (75). Overheating leads to the formation of advanced
glycosylation end products (AGEs), which are thought to
INFLUENCE SUGARS METABOLISM influence inflammation and possibly insulin resistance,
Sugars can also be listed according to their physicochemical, whereas acrylamide (a product resulting from a reaction of a
technological, and functional characteristics, which are reducing sugar with the amino acid asparagine) is a known
important for food design and food processing (see Table 5). carcinogen. This information has prompted strategies to
These characteristics can also affect the responses in our body. limit the formation of harmful MRPs. For example, limiting
Two examples will be given here: sugars as well as the asparagine content of potato and cereal
products before thermal processing by measures such as
A) The relative sweetness of sugars (Table 6) plays an essential selecting potato varieties with a low content of reducing
role when sweetening foods and beverages. The lowest sugars may help reduce acrylamide. Targeted potato storage
amount of a sugar needed to realize a certain sweetness temperatures such as storage below 8◦ C causes an increase in
is determined by the highest relative sweetness. Most used reducing sugar content and higher amounts of acrylamide.
for sweetening is sucrose, the reason why the sweetness Modifying heat-processing conditions (time, temperature)
of sucrose is set at 100%. To replace sucrose (sweetness and applying appropriate preheating treatments, such as
= 100) in a drink with glucose (in a concentration of soaking or blanching, can also help impact on the level
10% of its relative sweetness ∼
= 70), 30% more glucose is of reducing sugars and thereby reduce the formation of
required to achieve the same degree of sweetness. As a MRPs (76, 77).

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Brouns Saccharide Characteristics and Health

SUGARS AND ORAL HEALTH will increase tooth surface contact exposure time, thus enhance
detrimental effects on tooth mineral. Sucrose is known to be most
Recently, the WHO (62) recommended to reduce the potent in causing cariogenicity, which raises questions; since the
consumption of free sugars to preferably below 5% of the effects appear to be more potent than the effects of its composing
total energy intake as a conditional ∗ recommendation for monomers glucose and fructose.
both adults and children, the most important reason being Recently, it was hypothesized that an oral microbiota
the detrimental effects on oral health, despite the fact that imbalance due to frequent sucrose exposure may be a causal
the evidence was judged to be of a very low quality (62). factor driving sucrose to be more harmful because sucrose
[∗ Conditional recommendations are made when there is less exposure disrupted the homeostasis between acid-producing
certainty “about the balance between the benefits and harms and alkali-producing bacteria (90). Because the oral microbial
or disadvantages of implementing a recommendation.” This composition and metabolism changed significantly with sucrose
means that “policy-making will require substantial debate and exposure, while no significant difference was detected after
involvement of various stakeholders” for translating them into lactose and glucose exposure, the authors claim that these
action (78)]. findings indicate that the cariogenicity of sugars is closely related
This evidence was based on data derived by experts as to their effects on the oral microecology.
published in various reviews (79–82). Detrimental effects of Acidified drinks containing substantial amounts of sucrose
sugars on oral health occur along two main routes: Firstly, are of particular concern (91, 92), because they do not only
this can be in the form of demineralization of enamel and cause caries but also dental erosion. Even acidified drinks with
dentine caused by acid, resulting from saccharolytic fermentation low sugar contents or without sugars making use of non-sugar
of sugars by oral microbiota; these monosaccharides and sweeteners are erosive because of the acids present therein result
disaccharides include glucose derived from starch degradation (93). Despite the primary focus on the role of sugars in causing
by salivary amylase; Secondly, detrimental effects can result from caries, it should be noted that the process of dental erosion and
exposure to food acids added to sugar-sweetened or light drinks, caries initiation is multifactorial (36, 94).
or acids naturally present such as in citrus juices, resulting in a In particular, the effects of sucrose appear to be of great
low drink PH. These food-acids will directly erode the enamel concern during childhood, given the fact that SSBs intake
and dentine without intervention of the oral microbiota In significantly increases the caries burden in 10-year-olds with
normal conditions, the acid present in the food/drink or formed attenuated effects in 15-year-olds-age groups that are known to
by the microbiota is buffered over time and hence neutralized be the highest consumers of free sugars. To prevent caries, SSBs
by saliva. In addition, saliva at neutral pH is supersaturated for consumption should therefore be reduced, especially in children
calcium and phosphate, enabling the repair of the acid-induced and adolescents (95). Of great concern are a simultaneous
demineralization (83). Acids derived from sugars can cause combination of high sipping frequency and low PH beverage
net demineralization when frequently taken and this is more and sugar concentration, especially in young children, leading
detrimental if salivary buffer capacity is exceeded when saliva to early childhood caries. Giving very young children sugary
production is low or absent. Examples of the latter are athletes drinks in a sipping bottle will lead to continuous small quantities
during intensive exercise when saliva production is inhibited and flushing especially of the front teeth. This process will be even
persons suffering from a low or absent salivary flow as a result more detrimental if the child falls asleep, resulting in a low
of cancer radiation treatment, autoimmune diseases, (multiple) salivary flow and the reduction of the salivary-buffering effect
medications or physiologically by ageing. (96, 97). There is no doubt about the fact that sugar and food
Many studies have been performed to define the in vivo acids are not the only factors of importance. Poor oral hygiene,
(in situ) cariogenic and erosive effects of sugars and acids on use of fluoride, appropriate salivary flow, presence of calcium
tooth mineral by the application of small intra-oral blocks of in the drink/food, type of food acid used (94), consumption
dentine or enamel or by using standardized solution enamel- pattern and bottle or breastfeeding (97) play a role in the etiology
rinsing essays in vitro. In the latter, the effects of remineralization of caries. In addition, the frequency of exposure may be more
can also be studied in detail. Depending on the frequency relevant than the quantity. Van Loveren (98) addressed the
and dose, sucrose, glucose, fructose, lactose or starch may all question of which sugar-reducing strategy is the best for caries
result in demineralization (84–88). It appears that the molecular prevention. To answer this question, the following aspects should
composition of sugars plays a role in the degree of fermentability be addressed: the shape of the dose-response association between
by oral microbiota. For example, sucrose is composed of sugar intake and caries, the influence of fluoridated toothpaste on
glucose and fructose, has an α-1,2 bond and is more rapidly the association of sugar intake and caries, as well as the relative
fermented, and this lead to a critical lowering of plaque pH contribution of frequency and amount of sugar intake to caries
than isomaltulose, which is composed of the same monomers levels. The author argues that when fluoride is appropriately used,
but which has an α-1,6 bond. Along similar lines, starch which the relation between sugar consumption and caries is very low
is rapidly degradable by amylase and which leads to a higher or absent. The high correlation between amount and frequency
glucose availability appears to be more cariogenic than slowly hampers the decision on which of the two is more important.
digestible starch which contains a higher fraction of amylose (89). Reducing the amount without reducing the frequency does not
It needs to be considered in this respect that a sticky food-matrix seem to be an effective approach to prevent caries.

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Brouns Saccharide Characteristics and Health

CONCLUDING REMARKS consumption (1, 3). It appears that mutual and interactive
effects exceed the sum of the individual characteristics, while
All rapid fermentable sugars give rise to acid production by they also determine the effects on health and disease. For this
microbiota present in the oral cavity which, dependent on reason, an increased focus on the overall effects and quality
frequency of exposure, salivary buffer capacity, presence of of carbohydrate sources and meals for food-based guidelines
calcium for remineralization and oral hygiene status will impact rather than individual component-based recommendations
on erosive potential and cariogenicity. All digestible CHOs is desired.
deliver “sugars” as monosaccharides to the gut epithelium for
absorption. Post-absorption, the metabolism of these monomers AUTHOR CONTRIBUTIONS
is basically identical and independent of the original source.
However, the way in which CHOs have been processed (natural, The author confirms being the sole contributor of this work and
low-processed vs. refined/highly processed, and heat-exposed), has approved it for publication.
the matrix in which these CHOs are present (e.g., liquid,
solid, viscous, and non-viscous), the co-presence of other FUNDING
nutrients (e.g., proteins, polyphenols, vitamins, minerals, and
plant-bioactive substances) in the natural CHO source/matrix Fund for open access publication: Maastricht University.
vs. their absence in refined CHOs and the dose ingested all
play a role in the overall effects in the human body. Looking ACKNOWLEDGMENTS
at one particular CHO characteristic will almost always lead
to a different conclusion, such as that fructose is toxic (99) I thank Prof. C. van Loveren, Academic Centre for Dentistry,
than evaluating from a “total perspective”; fructose is only Amsterdam for the critical reading and suggestions to the section
toxic at excessive exposure levels that do not mimic human Sugars and Oral Health.

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for developing goals for sugar intake. BMC Public Health. (2014) absence of any commercial or financial relationships that could be construed as a
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