Abstract
Dietary compounds significantly affected starch enzymatic digestion. However, effects of dietary compounds on starch digestion and their underlying mechanisms have been not systematically discussed yet. This review summarized the effects of dietary compounds including cell walls, proteins, lipids, non-starchy polysaccharides, and polyphenols on starch enzymatic digestion. Cell walls, proteins, and non-starchy polysaccharides restricted starch disruption during hydrothermal treatment and the retained ordered structures limited enzymatic binding. Moreover, they encapsulated starch granules and formed physical barriers for enzyme accessibility. Proteins, non-starchy polysaccharides along with lipids and polyphenols interacted with starch and formed ordered assemblies. Furthermore, non-starchy polysaccharides and polyphenols showed robust abilities to reduce activities of α-amylase and α-glucosidase. Accordingly, it can be concluded that dietary compounds lowered starch digestion mainly by three modes: (i) prevented ordered structures from disruption and formed ordered assemblies chaperoned with these dietary compounds; (ii) formed physical barriers and prevented enzymes from accessing/binding to starch; (iii) reduced enzymes activities. Dietary compounds showed great potentials in lowering starch enzymatic digestion, thereby modulating postprandial glucose response to food and preventing or treating type II diabetes disease.
Introduction
Starch which is a polysaccharide composed of linear chains (amylose) or branched chains (amylopectin) is a major source of energy in the human diet. Starch digestion is accomplished by two type of enzymes in human gastrointestinal tract (GIT): (i) salivary and pancreatic α-amylases and (ii) intestinal brush border glucoamylases, maltase-glucoamylase, and sucrase-isomaltase (). Amylase digests amylose into maltose subunits (disaccharide) and amylopectin into branched chains (i.e., dextrins). Both maltose and dextrins are digested by enzymes located in intestinal brush border, which in turn produced glucose. The glucose released from starch is subsequently absorbed in the intestine and hydrolyzed to produce adenosine triphosphate or stored in animals as the polysaccharide glycogen. Accordingly, starch-based diets are commonly the main foods that provide the necessary energy. However, rapid digestion of starch contributes to postprandial hyperglycaemia, which in turn possibly results in an impaired insulin secretion and the incidence of chronic diseases such as obesity and type II diabetes (, ). Slowing starch enzymatic digestion in the GIT is of great interest in preventing the incidence of chronic diseases.
Enzymatic reactions consist of three steps: diffusion of enzymes to the solid surfaces, absorption/binding, and catalysis (). Regarding to starch enzymatic digestion, there are two factors influencing the extent and rate of starch digestion: (i) barriers that slow down or prevent digestive enzymes from accessing/binding to starch and (ii) starch structural features that limit enzyme action after initial binding (). Starch structuration on fine structure, helical structure, crystalline structure, lamellar structure, short-range ordered structure, and nanoscale aggregate structure significantly slowed enzymes binding with starch and reduced enzymes catalyzation toward starch (–). It has been summarized that slowly digestible starch (SDS) was the fraction with high α-1,6 linkages, short branch chains [degree of polymerization (DP) <13], long chains with DP 25–36, or imperfect helical and crystalline structures, while the resistant starch (RS) was the fraction rich in high amylose content, double helix-promoting chains with DP ca. 12–24 and DP ≥ 37, along with some chains with DP 25–36, perfectly-packed double helices and crystalline structures, V-type crystals, or densely-packed crystalline lamellae and more ordered reassembled aggregate structures (). According to previous studies (–), starch digestion was affected not only by its intrinsic structures but also by the interactions between starch and dietary non-starchy compounds and between digestive enzymes and non-starchy foods. At present, many reviews have indicated dietary compounds such as polyphenols, lipids, and non-starchy polysaccharides significantly affected starch digestion (–). However, the effects of dietary compounds such as cell walls, protein, lipids, non-starchy polysaccharides, and polyphenols on starch digestion and their underlying mechanisms have been not systematically summarized yet.
Therefore, this review provided a survey of the latest developments on dietary strategies for slowing starch enzymatic digestion, with a particular focus on the mechanisms underlying the modulation of starch digestion. Future perspectives regarding the dietary strategies for the control of starch digestion will be proposed. This review can provide better insights into the modulation of starch enzymatic digestion through complexation with dietary compounds.
Cell walls slow starch digestion
The basic architecture of plant cell wall is shown in Figure 1. Plant cell walls are cellulose-based assemblies containing cellulose and non-cellulosic polysaccharides (e.g., pectin, xyloglucans, heteroxylans, and β-glucans), lignin and some proteins (, ). Cellulose fibrils assembled and served as scaffold filling with amorphous non-starchy polysaccharides (). While the filling non-starchy polysaccharides prevented aggregation and collapse of the cellulose/hemicellulose network, the interactions of the non-starchy polysaccharides significantly contributed to the density and porosity of cell walls and in turn determined the permeability of hydrolases through the cell walls (, ).
Figure 1
Effects of cell walls on starch gelatinization and enzyme-starch interaction are shown in Figure 1 (
Li et al. indicated that enzymatic digestion of starch granules entrapped within cell walls in pulses depended on both the intactness of cell walls as well as the ordered structures of pulses after food processing (
In addition to the roles of cell walls on starch ordered structures, cell walls also bound with enzymes and affected hydrolyzation activities of the enzymes (
Proteins slow starch digestion
Protein is one of the most important compounds in foods systems. It has been summarized that proteins could non-covalently interact with starch through hydrogen bonding, hydrophobic interactions, electrostatic forces, ionic interactions, and van der Waals force (
Although the effects of protein structures on starch digestion in foods systems has not been resolved yet, it can be preliminarily concluded that dietary protein has a strong ability to mitigate starch enzymatic digestion. Dietary proteins affected starch enzymatic digestion via different pathways: (i) proteins acted as physical barriers and restricted the interaction of enzymes with starch molecules (
Lipids slow starch digestion
Lipids are the important hydrophobic dietary compounds in foods. In starch-containing foods systems, lipids tended to interact with starch through hydrophobic interaction and formed starch-lipid inclusion complexes or starch-lipid-protein complexes (
Starch structures, lipid type and structures, and the preparation conditions significantly affected the formation of starch-lipid inclusion complexes. Amylose is much easier to interact with lipids compared with amylopectin. Starches with higher amylose content formed more starch-lipids complexes compared with less amylose-containing starches (
Increasing lipids concentration favored the interaction of lipids with starch, thereby lowering starch digestion in a higher magnitude (
Proteins in food systems affected the formation of starch-lipid inclusion complexes. β-lactoglobulin favored lipids (e.g., fatty acids and monoglyceride) dissolution in water and promoted lipid-starch entanglement (
During the preparation of starch-lipid inclusion complexes, the temperature, complexation time and modes, pH, NaCl, and cooling rate significantly affected the formation of the complexes (
Non-starchy polysaccharides slow starch digestion
Non-starchy polysaccharides affected starch gelatinization, the viscosity of starch paste, starch reorganization, and enzymes activities, which affected starch digestion by different modes. Polysaccharides such as chitosan, guar gum, and xanthan interacted with starch granules and lowered starch swelling and amylose leaching during hydrothermal treatment (
Due to the interaction between starch and non-starchy polysaccharides, the viscosity of starch suspension which complexed with xanthan gum, guar gum, konjac glucomannan, pectin, and chitosan significantly increased (
The interaction between starch and non-starchy polysaccharides also significantly affected properties of matrix structures formed by starch and non-starchy polysaccharides (
Starch digestion was controlled not only by starch ordered structures and food viscosity, but also by the activities of enzymes. Pectin bound with pancreatic amylase to reduce amylase activity, resulting in slower starch enzymatic digestion (
Accordingly, non-starchy polysaccharides lowered starch digestion rate and extent by four different ways: (i) interacted with starch granules and restricted starch disruption during food processing (
Polyphenols slow starch digestion
Effects of polyphenols on starch digestion are schematically shown in Figure 2. α-amylase and α-glucosidase are two key enzymes for starch digestion. Accordingly, starch digestion could be significantly lowered through reducing activities of α-amylase and α-glucosidase. Tea polyphenols, flavonoids, phenolic acids, and tannins significantly reduced activities of α-amylase and α-glucosidase, which showed great potentials in mitigating starch digestion as summarized in previous reviews (
Figure 2

Mechanism scheme for effects of polyphenols on starch digestion. Figure was adapted from Sun and Miao (
Figure 3

Flavonoids key sites that affecting activities of α-glucosidase. The up arrows and down arrows represent increasing and reducing the inhibition activity, respectively. Figure was collected from Xiao et al. (
Polyphenols could interact with starch and form ordered starch structures, and in turn, lowering starch digestion as shown in Figure 2. It has been reported that polyphenols could interact with starch and promote the formation of starch ordered structures (
Other dietary compounds
In addition to the compounds discussed above, other dietary compounds such as NaCl and phytosterols also affected starch digestibility (
Concluding remarks and future directions
This review summarized effects of dietary compounds including cell walls, proteins, lipids, non-starchy polysaccharides, and polyphenols on starch enzymatic digestion and their underlying mechanisms were discussed. Dietary compounds lowered starch digestion through three pathways: (i) retained starch ordered structures or formed ordered assemblies chaperoned with these dietary compounds; (ii) formed physical barriers and prevented enzymes from accessing/binding to starch; (iii) reduced enzymes activities. Cell walls, proteins, and non-starchy polysaccharides restricted starch disruption during hydrothermal treatment and the retained ordered structures limited enzymatic binding. In addition, they encapsulated starch granules and formed physical barriers for enzymes accessing. Proteins, non-starchy polysaccharides along with lipids and polyphenols interacted with starch and formed ordered assemblies. Non-starchy polysaccharides and polyphenols showed robust ability to reduce activities of α-amylase and α-glucosidase. Comparing with cell walls, protein, and non-starchy polysaccharides, lipids and polyphenols had stronger ability to slow starch digestion.
Food systems are relative complex with cell walls, proteins, lipids, non-starchy polysaccharides, polyphenols, vitamin, minerals, sugar, salts, etc. Dietary compounds might interact with each other and affect starch digestion in complicated pathways. How the complex dietary compounds affected starch digestion in real foods systems must be further investigated. Currently, effects of dietary compounds on starch digestion were interrogated in vitro. Dietary compounds would be digested and absorbed in the gastrointestinal tract and in turn affected starch digestibility. Starch in vitro digestibility may quite different to in vivo digestibility. Accordingly, in vivo glycemic response is the most important property of starchy foods. Further studies are needed to investigate the roles of dietary compounds on starch in vivo glycemic response. In addition, different groups of people such as children, athletes, middle- and old-aged human have different requirements for starch digestion rate and extent. Targeted structuring food structures and starch digestion behaviors via complexation with dietary compounds remains of interest.
Funding
The research was financially supported from project of the Scientific Research Innovation Program Xiyuanjiang River Scholarship of College of Life Sciences, Fujian Normal University (22FSSK004).
Publisher's note
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
Statements
Author contributions
CC: conceptualization, literature analysis, reviewing, and editing. MS: reviewing and editing. YZ: supervision, writing, reviewing, and editing. BC: investigation, writing, and editing. YH: investigation and reviewing. MW: supervision, reviewing, and editing. All authors contributed to the article and approved the submitted version.
Conflict of interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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Summary
Keywords
starch digestion, dietary compounds, starch structure, enzyme activity, nutrition
Citation
Chi C, Shi M, Zhao Y, Chen B, He Y and Wang M (2022) Dietary compounds slow starch enzymatic digestion: A review. Front. Nutr. 9:1004966. doi: 10.3389/fnut.2022.1004966
Received
27 July 2022
Accepted
23 August 2022
Published
15 September 2022
Volume
9 - 2022
Edited by
Yanjun Zhang, Chinese Academy of Tropical Agricultural Sciences, China
Reviewed by
Jinhu Tian, Zhejiang University, China; Zhili Ji, Wuhan Polytechnic University, China; Xiaoyan Tan, Nanjing Tech University, China
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Copyright
© 2022 Chi, Shi, Zhao, Chen, He and Wang.
This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Chengdeng Chi c_cd@fjnu.edu.cnMeiying Wang mwang15@uoguelph.ca
This article was submitted to Food Chemistry, a section of the journal Frontiers in Nutrition
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