From structure to function: A comprehensive overview of polysaccharide roles and applications

Abstract

Polysaccharides, also known as glycans, are biological macromolecules consisting of many monosaccharide units. Alongside proteins, nucleic acids, and lipids, they constitute the four fundamental substances crucial for life activities and essential for the growth and development of living organisms. As natural products with inherent biological activity, polysaccharides are widely available, nontoxic, and possess numerous functional properties, holding immense potential for advancement in food, medicine, and cosmetics. Furthermore, the exploration of polysaccharide‐based drugs, as an alternative to conventional therapies, emerges as a promising avenue for addressing future disease challenges. This article comprehensively reviews the sources, structural characteristics, synthesis, degradation, functions, and applications of polysaccharides. The potential of polysaccharides for pharmacological applications, in antitumor, antiaging, antioxidant, hypoglycemic, anti‐inflammatory, antiviral, and anticoagulant properties, is summarized. Additionally, the role of polysaccharides in environmental protection is discussed. It is anticipated that this review will offer innovative strategic insights, serving as a theoretical foundation and inspiration for the subsequent research on polysaccharides in healthcare.

INTRODUCTION AND BACKGROUND

Polysaccharides exhibit numerous biological activities and hold significant medicinal value, being widely distributed in nature (Zhou & Huang, 2021). In comparison to proteins and nucleic acids, research on polysaccharides has developed more recently and has not received adequate attention. Traditionally, it has been simplistically considered a substance providing energy in living organisms and maintaining the structure of cellular organization (Zheng et al., 2015). However, a plethora of recent research has unveiled that polysaccharides possess a broad spectrum of physiological functions and intricate biological activities. They play pivotal roles in various biological processes that contribute significantly to human health, encompassing intercellular communication, embryonic development, responses to bacterial or viral infections, and modulation of humoral and cellular immunity (Chen et al., 2019; Gabius, 2018; Jin, Shi, et al., 2021; Kieninger & Maldener, 2021; Weng et al., 2018; Zhou & Huang, 2021; Zimmermann & Lepenies, 2015). Moreover, in contrast to many chemically synthesized medications currently employed in clinical settings, polysaccharides are more conducive to long-term use and exhibit fewer side effects due to their natural origin (Niu et al., 2021; Zhang, Wang, Tan, et al., 2022). Consequently, the full exploitation of the medicinal potential inherent in polysaccharides may positively impact the advancement of human health.

Currently, researchers have conducted extensive work on polysaccharides. First, the functional properties of polysaccharides are intricately linked to their structural composition, making characterization a vital aspect of their research. However, defining the structural characteristics of polysaccharides uniformly poses challenges due to irregular changes influenced by the extraction process (Nai et al., 2021). Consequently, methods for polysaccharide identification must meet criteria such as simplicity, usability, and reliability. Second, the synthesis and degradation of polysaccharides involve the coordinated action of numerous enzymes, representing crucial life activities in living organisms (Sun et al., 2020). To facilitate in-depth investigations into their diverse properties, the preparation of high-purity polysaccharides becomes imperative. However, naturally synthesized polysaccharides often fall short of meeting high purity standards, and the outcomes of artificially synthesized polysaccharides are not always satisfactory (Tanaka et al., 2012), highlighting persisting challenges in the purification of polysaccharides. Certainly, the research on polysaccharides focuses on their biological activity and various applications. Polysaccharides have evolved into a complete development system from function to application, finding widespread use in various fields such as medicine, food, and industrial production (Gong et al., 2022; Song et al., 2023; Zhang, Xu, et al., 2022).

In recent years, there has been a growing number of review articles focusing on polysaccharides due to increased attention. However, these reviews tend to favor research in singular fields and often lack a thorough investigation into the combined mechanisms of various polysaccharide functions (Gaikwad et al., 2024; Manzoor et al., 2022; Yu et al., 2018). In this context, this article provides a comprehensive review encompassing polysaccharide properties, synthesis, degradation, functional roles, and application prospects. Furthermore, it delves into the potential applications of polysaccharides in promoting green and healthy living. The aim is to offer researchers in the field with systematic theoretical information on the applications of polysaccharides. It is anticipated that this review will contribute to a deeper understanding of polysaccharide applications.

INTRODUCTION TO POLYSACCHARIDES

Definition and characteristics of polysaccharides

Polysaccharides represent a class of biomolecules comprised multiple monosaccharide molecules linked by glycosidic bonds, with the capacity to contain hundreds to thousands of monosaccharide residues. These molecules can adopt various structural forms, including linear, branched, or network configurations (Murphy et al., 2023). Linear polysaccharides consist of monosaccharides linked in a straight line in a specific order, whereas branched polysaccharides have additional monosaccharide molecules attached to the main chain through branching connections. Network polysaccharides are made up of different structures cross-linked to create a three-dimensional mesh (Nagae & Yamaguchi, 2014). Polysaccharides exhibit a wide range of functions, serving as vital sources for energy storage and supply and playing crucial roles in biological processes such as cell recognition, signaling, and immune responses (Elango et al., 2023; Murphy et al., 2023). Furthermore, they find applications as food additives, materials for tissue engineering, and components in drug delivery systems (Chandra et al., 2022; Fricain et al., 2013; Kakar et al., 2021). Due to their water-soluble nature and strong affinity for water molecules, polysaccharides can form gelatinous structures of varying viscosities in solution, making them excellent raw materials for the preparation of bioactive hydrogels (Nie et al., 2019). Additionally, polysaccharides offer extensive opportunities for both chemical and functional modification (Li et al., 2016). It is noteworthy that most polysaccharides exhibit high biodegradability and decompose in biological systems into low molecular weight substances and nontoxic metabolites through specific enzymes, promoting environment-friendly disposal (Li, Xiang, et al., 2022). This characteristic positions them favorably for applications in environmental protection. In conclusion, polysaccharides demonstrate diverse functions and application potentials, underscoring the importance of continued exploration and development for applications in both industrial and biological sciences.

Classification and sources of polysaccharides

Polysaccharides exhibit significant diversity and can be classified into three main categories based on their sources: plant, animal, and microbial (Song et al., 2023). Plant polysaccharides constitute for the majority of polysaccharide composition in nature, followed by microbial polysaccharides and animal polysaccharides. Plant polysaccharides are widely distributed in marine algae, food plants, and medicinal plants. Animal polysaccharides are commonly found in marine organisms, mollusks, mammals, birds, and insects. Microbial polysaccharides can be derived from different types of microorganisms, including bacteria, fungi, and algae. The classification of polysaccharides is summarized in Table 1.

TABLE 1 The classification of polysaccharides: exploring functions, features, and properties.

Species	Polysaccharides	Features	Functionalities	References
Plant	Astragali radix polysaccharide, Angelica sinensis polysaccharide, Poria cocos polysaccharide, Panax ginseng polysaccharide, Lycium barbarum polysaccharide, Ganoderma lucidum polysaccharide, Auricularia auricula polysaccharide, Bletilla striata polysaccharide, Polygonatum sibiricum polysaccharide, Hericium erinaceus polysaccharide, Achyranthans, cellulose, and starch	Very widely available; an important component in the functioning of herbal medicine; an energy storage carrier in plants; a major component of plant cell walls	Tissue support; nutrient storage; antitumor; anti-inflammatory and antiviral; antiaging; antioxidant; hypoglycemic; anticoagulant; immune enhancement	Gao et al. (2015), Wei et al. (2018), Weng et al. (2018), Jin, Oh et al. (2021), Mahajan et al. (2021), Peng, Zhao et al. (2022)
Animal	Heparin, hyaluronic acid, chondroitin sulfate, and chitin	Animal polysaccharides are mostly linked to proteins; good compatibility with human cells	Regulating blood lipids; unblocking blood vessels; enhancing lipase activity; anticoagulation; antithrombosis; liver protection; lubricating bones and joints; protecting cells; regulating blood lipids and cholesterol	Hao et al. (2019), Li et al. (2018), Satitsri and Muanprasat (2020), Vasvani et al. (2020)
Microbe	Xanthan gum, pullulan, gellan gum, dextran, curdlan, seaweed polysaccharide, hyaluronic acid, and bacterial cellulose	Predominantly extracellular polysaccharides; short production cycle; easy to isolate and purify	Drug release modifiers; organizational engineering; wastewater treatment; wound dressing; drug carrier; food additives; skin protection; bioactivity	Bayer (2020), Costa, Silva-Correia et al. (2018), Kumar et al. (2018), López-Hortas et al. (2021), Nakamura et al. (2023), Wang, Tavakoli et al. (2019), Wei et al. (2021)

Plant polysaccharides

The molecular weights of plant polysaccharides span from tens of thousands to millions (Wang, Zhou, et al., 2023). Plant polysaccharides can be categorized into three groups based on their location within the cell: intracellular, cell wall, and extracellular. Among these, cell wall polysaccharides have attracted attention due to their substantial medicinal value. Three primary types of polysaccharides constitute plant cell walls: cellulose, pectin, and hemicellulose (Ruprecht et al., 2022). Hemicellulose can further be categorized into xyloglucan, mannan, and xylan based on the sugar groups it contains (Pauly et al., 2013). Numerous plant sources, such as brown algae, tea, tartary buckwheat, echinacea (the medicinal tree of Mali), and mistletoe, serve as excellent raw materials for extracting polysaccharides. In addition, some polysaccharides with potential therapeutic effects against cancer, viruses, and immune and system modulation can be extracted from medicinal plants like Dendrobium officinale kimura et migo, Astragali radix, Morinda officinalis, Cordyceps, Achyranthis bidentatae radix root, Codonopsis pilosula, Glycyrrhiza (Licorice), and Asari radix et rhizoma (Ji et al., 2024; Liu & Huang, 2019; Zhang, Xu, et al., 2022). As a result, research on plant polysaccharides has seen widespread attention and study in recent years.

Animal polysaccharides

Polysaccharides identified in animal tissues and organs are termed animal polysaccharides. Naturally occurring animals or animal parts rich in polysaccharides in nature include sea cucumbers, loaches, shrimps, and velvet antlers (Gong et al., 2022; Liu, Ding, et al., 2023; Pal et al., 2021; Zhang & Huang, 2005). Presently, the fully utilized animal polysaccharides mainly encompass heparin, hyaluronic acid, chondroitin sulfate, and chitin (Zhao et al., 2015). Heparin, a sulfur-containing mucopolysaccharide, is widely distributed in the animal blood, thymus, muscles, and five internal organs (heart, liver, spleen, lungs, and kidneys) (Hao et al., 2019), exhibiting multiple pharmacological effects such as regulating blood lipids and unblocking blood vessels. Chondroitin sulfate is mainly found in tissues such as cartilage, nasal bones, and tendons and is an acidic mucopolysaccharide extracted from fresh or dried cartilage (Li et al., 2018). It possesses functions such as enhancing lipase activity, anticoagulation, antithrombosis, and liver protection. Hyaluronic acid, an acidic polysaccharide, is mainly found in the heads, umbilical cords, skin, and eyeballs of animals, composed of glucuronic acid and N-acetyl-d-glucosamine linked by glycosidic bonds (Vasvani et al., 2020). Hyaluronic acid, a potent active ingredient, demonstrates positive effects in lubricating bones and joints and protecting cells, making it suitable for the development of healthy foods and cosmetics. Animal polysaccharide chitin, found in shrimps, crabs, shells, and the cell walls of some higher plants (Satitsri & Muanprasat, 2020), regulates blood lipids and cholesterol levels, presenting promising prospects in medicine. In summary, animal polysaccharides exhibit a wide range of potential applications with significant medicinal and pharmaceutical value.

Microbial polysaccharides

Similar to plant polysaccharides, microbial polysaccharides can be categorized based on their structural components into intracellular, extracellular, and cell wall polysaccharides (Song et al., 2023). Research on microbial polysaccharides originated in the mid-20th century. The extracellular polysaccharide xanthan gum was first successfully extracted and studied in 1952 from Xanthomonas black rot in cabbage, which attracted continuous attention (Kumar et al., 2018). In recent years, functional research on microbial polysaccharides has gradually improved due to the development of related technologies, emerging as a research hotspot in various fields, including biology, immunology, and medicine. Microbial polysaccharides with diverse active functions, such as xanthan gum, pullulan, gellan gum, dextran, curdlan, seaweed polysaccharide, hyaluronic acid, and bacterial cellulose, have been discovered and investigated (Song et al., 2023; Wang, Tavakoli, et al., 2019; Wei et al., 2021). Additionally, due to the abundance of marine microbial resources and intense interspecies competition, many marine microbes produce polysaccharides through metabolism that possess unique functions. Consequently, marine microbial polysaccharides are increasingly gaining attention. Most of these polysaccharides are extracellular, holding significant scientific significance and commercial value (Xu et al., 2017).

Structure of polysaccharides

The structural complexity of polysaccharides surpasses that of proteins and DNA, presenting a significant challenge for their structural analysis. The classification of polysaccharide structures follows a pattern similar to that of proteins and DNA (Diener et al., 2019; Zdunek et al., 2021). The primary structure serves as the foundation for the structural analysis of polysaccharides. The second, third, and fourth levels are collectively known as high-level structures. The secondary structure involves the regular conformation formed between the main chains, with hydrogen bonds being the primary secondary bonds. The tertiary structure of a polysaccharide chain refers to the polysaccharide chain structure formed by noncovalent interactions among the hydroxyl, carboxyl, amino, and sulfate groups on the polysaccharide units, building upon the secondary structure. Finally, the aggregates formed by noncovalent bonds between polysaccharide chains are termed the quaternary structure of polysaccharides. Recently, the analysis of polysaccharide structures is primarily carried out using methods such as nuclear magnetic resonance, X-ray diffraction, mass spectrometry, infrared spectroscopy, and scanning electron microscopy (Chakraborty et al., 2022; Guo et al., 2023; Li, Zhao, et al., 2022; Wang, Zhao, et al., 2023; Zhao, Yang, et al., 2023), as depicted in schematic shown as Figure 1.

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SYNTHESIS, MODIFICATION, AND DEGRADATION OF POLYSACCHARIDES

Synthesis of polysaccharides

Polysaccharides can be synthesized through two methods: natural and chemical. Natural synthesis takes place within living organisms and relies on enzymatic catalysis. This method offers efficiency, specificity, and mild reaction conditions, but its application is constrained by the organism's metabolic pathways and enzyme specificity (Zhao, Ma, et al., 2021). On the other hand, chemical synthesis is conducted in a laboratory setting. This approach involves designing synthetic pathways and regulating reaction conditions to construct polysaccharide structures. However, the synthesis process is relatively complex (Li, Yu, et al., 2020). Figure 2 provides a summary of the similarities and differences between the natural and chemical synthesis pathways of polysaccharides.

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Natural synthesis pathways of polysaccharides

The natural synthesis pathways of polysaccharides encompass several key routes, including the nucleotide sugar, glycogen, exopolysaccharide synthesis, cleavage and modification, and the polymerase (Bar-Peled & O'neill, 2011; Ellingwood & Cheng, 2018; Islam & Lam, 2014; Thompson et al., 2012; Yang et al., 2020).

The nucleotide sugar pathway, a pivotal route for polysaccharide synthesis, sugar nucleic acids combine with other molecules to form polysaccharides through phosphorylation and glycosyltransferase. Nag et al. (2012) found that plant polysaccharides are produced from nucleotide sugars, wherein nucleoside diphosphate (NDP) forms sugar complexes through chemically activated phosphoester bonds. Glycosyltransferases then catalyze the incorporation of specific monomers from NDP sugars into the extending polysaccharide chain, ultimately forming complete polysaccharides. The glycogen pathway, another significant synthesis route, involves the activation of glucose into uridine diphosphate glucose (UDPG). UDPG is then transferred to the nonreducing end of the glycogen primer and connected with α-1,4 glycosidic bonds to form the main chain under the action of glycogen synthase, leading to the further extension and formation of glycogen (Ellingwood & Cheng, 2018). The exopolysaccharide synthesis pathway parallels the glyconucleic acid pathway, generating polysaccharides through phosphorylation and glycosyltransferase (Yang et al., 2020). Extracellular polysaccharide synthesis involves monosaccharide binding to nucleotides, forming precursor nucleotide-activated sugars. Under enzyme catalysis, these precursors are converted into starting units, elongated through conversion and polymerization reactions to create repeating units. Finally, these repeating units are sequentially linked and transported out of the cell via lipid carriers. Cleavage and modification pathways, vital for polysaccharide synthesis, encompass enzymatic catalysis and chemical modification, playing a crucial role in shaping different sugar chain structures (Thompson et al., 2020). The polymerase pathway involves the connection of monomeric sugars to form polysaccharide chains through the action of polymerase. Wiseman et al. (2021) discovered that polysaccharide copolymerase Wzz regulates the length of growing polysaccharide chains, while polymerase Wzy polymerzes oligosaccharides into longer polysaccharide chains. Further research has highlighted the importance of close contact between polymerase Wzy and copolymerase Wzz for the proper adjustment of polysaccharide chain length (Islam et al., 2013). These natural synthesis pathways play a vital role in organisms, ensuring the normal function of polysaccharides.

Chemical synthesis pathway of polysaccharides

Due to their complex structures and microscopic heterogeneity, natural polysaccharides pose a challenge in terms of separation and purification to obtain sufficiently pure polysaccharide samples. Unlike proteins or nucleic acids, polysaccharides cannot be mass-produced using PCR technology because there is no set template for their biosynthesis. Consequently, achieving polysaccharides with a consistent structure is partially attainable through chemical synthesis (Li & Ye, 2020). However, chemical methods typically only yield short polysaccharide fragments due to the complexity of polysaccharide structures and limitations in synthetic technology. Moreover, replicating the intricate effects of natural polysaccharides in the molecular size-dependent biorecognition process for short sugar chains is often challenging (Tanaka et al., 2012). A study found that chemically synthesized small molecule heparin inhibits thrombin activity. However, when compared with natural heparin, the inhibitory effect of synthetic pentadecaccharide remains weak (Petitou et al., 1999). Furthermore, synthetic short sugar chains may elicit lower titer antibody responses, whereas bacterial antigenic polysaccharide components frequently induce strong immune responses (Akhmatova et al., 2016). Therefore, researchers must endeavor to synthesize glycan compounds that are similar in size and complexity to natural polysaccharides to gain a deeper understanding of the biological functions and structure–activity relationships of polysaccharides.

The glycosylation reaction is one of the most important reactions in the chemical synthesis of polysaccharides, involving coupling reactions between sugar units. Glycosylation reactions are categorized as oxygen, nitrogen, and carbon glycosylation reactions based on the nucleophilic reactive group of the glycosyl acceptor. The corresponding reaction products are termed oxygen, nitrogen, and carbon glycosides (Inuki et al., 2022). Another critical issue in polysaccharide synthesis is the assembly of polysaccharides, which entails the effective assembly of sugar units into complex polysaccharide structures through the utilization of existing glycosylation methods. Nowadays, linear and convergent syntheses are the two main approaches for assembling polysaccharides (Liu, Qin, et al., 2021). Regardless of the strategy employed, it typically requires the adjustment of end-group substituents or the removal of temporary protecting groups on the sugar units. These procedures contribute to the complexity and time-consuming nature of polysaccharide synthesis, making it challenging to isolate and purify intermediates and, consequently, reducing the efficiency of polysaccharide synthesis. Over the past three decades, researchers have proposed some new strategies, including automated solid-phase synthesis, to address these challenges (Liu, Qin, et al., 2021). However, the chemical synthesis of complexly structured long-chain polysaccharides remains a daunting task. Currently, only a few syntheses of complex long-chain polysaccharides have been accomplished. Therefore, further exploration of efficient polysaccharide assembly strategies remains an urgent problem in the field of glycochemistry.

Chemical modification of polysaccharides

The application of polysaccharides depends on their origin and structure. Modification techniques can be used to selectively alter the structure of polysaccharides to meet specific functional requirements. Physical, chemical, and biological modifications are commonly employed to enhance the functional activity of polysaccharides. Studies have demonstrated that chemical modifications significantly enhance the structural and functional properties of polysaccharides (Ma et al., 2012). Chemical modification methods, such as polysaccharide chelation, carboxymethylation, sulfation, phosphorylation, and selenization, are effective approaches for safely improving polysaccharides (Gao et al., 2015; Guo, Chen, et al., 2020; Zhou & Huang, 2021). These methods can replace or introduce functional groups into polysaccharides under specific conditions, thereby changing the biological activity of polysaccharides.

Polysaccharide chelates are complexes formed by polysaccharide molecules with metal ions or other compounds. In such complexes, polysaccharides create a stable structure through the coordination of their functional groups (e.g., hydroxyl and amine) with metal ions or other compounds (Zhang, Chen, et al., 2019). The properties of polysaccharide chelates can be regulated by modulating the structural and functional groups of polysaccharides (Yin et al., 2012). Therefore, polysaccharide chelates hold a wide range of application prospects and research value. Sulfation modification is a chemical method wherein polysaccharide polymer chains are modified by adding sulfate groups (Chen et al., 2019). This process can be achieved by dissolving a known amount of polysaccharide in a solvent and treating it with a specific reagent under certain conditions to replace the hydroxyl group with a sulfate group. With its benefits of simplicity, nontoxicity, and affordability, it is a popular chemical modification technique for polysaccharides (An et al., 2022). Selenization modification is another common method for polysaccharide modification, based on introducing selenium into the polymer chain of polysaccharides (Gao et al., 2020). Polysaccharides contain various groups, such as free hydroxyl, aldehyde, and carbonyl groups, which can combine with inorganic selenium to form selenopolysaccharides. Numerous studies have demonstrated that selenopolysaccharides possess better antioxidant, antitumor, immunomodulatory, hypoglycemic, and heavy metal scavenging properties compared to inorganic selenium, polysaccharides, or mixtures of inorganic selenium and polysaccharides (Huang et al., 2020). Additionally, selenopolysaccharides exhibit lower toxicity and higher bioavailability, providing them an advantage over traditional selenium supplementation. Phosphorylation modification is one of the most important techniques for the structural modification of polysaccharides. The principle is to introduce a phosphate group to replace the hydroxyl group in the polysaccharide polymerization chain. The biological activity and properties of polysaccharides can be effectively altered by phosphorylation modification (Zhang, Su, et al., 2017). Acetylation modification is another common method of polysaccharide modification, based on the conversion of hydroxyl groups in polysaccharides to acetyl groups through esterification reactions (Ren et al., 2020). Acetyl groups can stretch polysaccharide branches and change the spatial arrangement of polysaccharide chains, exposing more polysaccharide hydroxyl groups. Thus, acetylation modifications can alter the water solubility and hydrophobicity of polysaccharides.

In addition to the aforementioned techniques, polysaccharide modification can also be carried out by methylation, iodination, sulfonylation, deacetylation, and desulfation (Staudacher, 2012). Although these techniques are not yet extensively employed, they provide significant insights into the pharmacological, chemical, and physical characteristics of polysaccharides. To enhance the structural and functional properties of polysaccharides, future research on polysaccharides needs to be intensified, and new modification methods also need to be continuously developed.

Degradation of polysaccharides

Due to their high molecular weight and difficulty in penetrating cell membranes, the majority of polysaccharides are not readily absorbed and utilized by the gastrointestinal system (Li et al., 2017; Xia et al., 2021). Consequently, they must undergo a series of degradation processes before being absorbed and performing various functions. The process in which enzymes or other degrading substances break down polysaccharide molecules into smaller molecules or monosaccharides is known as polysaccharide degradation. Currently, microbial transformation and enzymatic degradation are the primary methods.

Enzymatic degradation

Enzymatic degradation involves the use of specific biological enzymes to break down polysaccharides (Wu et al., 2020). The main processes include as follows: (1) The enzyme binds noncovalently to the polysaccharide substrate, forming an enzyme–substrate complex. (2) The enzyme hydrolyzes the substrate via a specific active site. (3) After the enzymatic degradation of the substrate is completed, the product is released, and the enzyme is then available to accept a new substrate for the reaction. The degradation process of polysaccharides by different enzymes may vary, and the specific mechanism depends on the structure of the enzyme and the chemical properties of the substrate. The study of enzymatic degradation of polysaccharides is widely reported for its extraordinary importance. For instance, Zeng et al. (2022) discovered that glycohydrolase and fungal galactosidase produced by Aspergillus flavus can catalyze the degradation of peach gum polysaccharides. In addition, the polysaccharide exhibited higher antioxidant activity after enzymatic degradation. Berlemont and Martiny (2016) found that the degradation of polysaccharides by glycoside hydrolases, lytic polysaccharide monooxygenases, and other polysaccharide-activating enzymes plays a crucial role in natural nutrient cycling and nutrient utilization in animals. Furthermore, Kobayashi et al. (2018) identified a range of polysaccharide hydrolyzing enzymes in some marine species. This adaptation allows them to obtain nutrients from plant debris and carry out enzymatic reactions at low temperatures, enabling them to thrive in the nutrient-poor deep-sea environment.

Microbial transformation

Microbial transformation is another important pathway for polysaccharide degradation. Some polysaccharides are challenging to break down and utilize due to their complex and specialized structures. Nonetheless, certain microorganisms, such as gut microbiota, can break down these polysaccharides into smaller monosaccharide units by secreting specific enzymes that assist the host in absorbing and utilizing them (Mckee et al., 2021). Yeager et al. (2017) found that bacteria belonging to the order Actinomycetes exhibit efficient degradation of complex polysaccharides in soil. La Reau and Suen (2018) found that Ruminococcus spp. can act on polysaccharides that are challenging to degrade in ruminants, converting them into various nutrients that can be utilized by the host. Additionally, this capacity for degradation serves an essential ecological role in the natural world by facilitating the recycling and breakdown of organic materials (Gardner, 2016).

Other

Polysaccharides can be degraded by chemical and physical methods in industrial production and environmental protection. However, some hazardous or hard-to-degrade chemical reagents used in chemical methods might cause contamination during polysaccharide degradation. In contrast, biodegradation has garnered a lot of attention due to its many advantages, including a high extraction rate, reproducibility, mild conditions, rapid reaction, environmental friendliness, and simple operation. In addition, glycosyl transfer has been extensively studied in the polysaccharide synthesis pathway but is mentioned less frequently in the degradation process of polysaccharides. Nevertheless, catalyzed by glycosyltransferases, polysaccharides can combine with other molecules to form new compounds that are more easily degraded or utilized. Therefore, glycosyl transfer may be crucial to the degradation of polysaccharides. Dura et al. (2017) found that starch modified by cyclodextrin glycosyltransferase (CGTase) exhibits higher digestibility in the presence of digestive enzymes. The modification of starch by CGTase can occur by altering the amorphous portion of the starch granules. This leads to the appearance of small pores on the surface, along with the release of oligosaccharides and the production of cyclodextrins, resulting in higher digestibility of the starch granules.

THE FUNCTIONS OF POLYSACCHARIDES IN LIVING ORGANISMS

Functions of polysaccharides in energy storage

Energy storage is a crucial physiological function evolved by organisms through natural selection (Cifuente et al., 2019). It enables the preservation of excess nutrients when available and their release when physiological needs arise in the future. Bioenergy is stored in long-term forms, including polysaccharides, proteins, and fats. Polysaccharides, in particular, play a vital role in energy storage across various forms in animals, plants, and microorganisms.

Among the polysaccharides, glycogen serves as a key energy storage molecule for certain microorganisms and animals. In animals, glycogen is predominantly present in the liver and muscles (Ellingwood & Cheng, 2018). Research has identified that glycogen, a macromolecular branched-chain water-soluble α-glucose unit polymer formed by glucose molecules connected through glycogen synthase, is a crucial player in maintaining energy metabolism homeostasis (Wang, Liu, et al., 2019). Starch, another energy-storing polysaccharide, is typically produced as semi-crystalline insoluble particles in the roots, stems, leaves, and seeds of plants (Mahajan et al., 2021). It can be categorized into two types based on its structural composition: amylose and amylopectin, with amylopectin being the primary polymerization state in starch (Seung, 2020). As a nutrient and energy reserve carrier in plants, starch plays a pivotal role in the biological world. Cellulose, mainly found in plant cell walls, is a significant polysaccharide involved in energy storage (Bhat et al., 2019). Although its molecular structure resembles that of starch, cellulose's glucose molecules are linked by β-glucose. Humans cannot digest cellulose, but it serves as a beneficial dietary fiber, contributing to digestive tract health when included in food. Furthermore, the exploration of the ocean has revealed that certain marine plants or microorganisms possess energy-storage polysaccharides with specific functions. For instance, the alginate polysaccharides extracted from seaweed not only store energy but also possess various biological activities, such as antiviral and antitumor properties. (Guo, Wang, et al., 2020).

Functions of polysaccharides in cellular structural support and protection

The stability of tissue cell structure is crucial for maintaining the health of organisms, with polysaccharides playing a significant role in this regard. First, they are integral components of the “cell wall,” the primary protective structure in plants. The cell wall's structural components include polysaccharides (cellulose, hemicellulose, and pectin), lignin, and proteins. Furthermore, polysaccharides are vital for bone development, providing strength and elasticity. Bones, crucial supporting structures in animals, rely on polysaccharides for the synthesis of organic components (Salbach et al., 2012). The integrity of bones not only determines an animal's ability to carry out normal life activities but also contributes to their potential for bone repair. Studies by Aguilar et al. (2019) demonstrated that combining chitosan (CS)-based scaffolds with other polymer biomaterials promotes osteogenesis, accelerating bone regeneration and accelerating the formation of new blood vessels. Tang et al. (2020) also found that combining CS-based hydrogels with bioactive molecules can promote tissue angiogenesis and bone repair in the application progress of CS-based injectable hydrogels. In addition to bone support, polysaccharides are essential for protecting soft tissue. Mucopolysaccharides, for instance, form mucus, envelop and shield soft tissues, reducing external stimulation and some inflammatory damage (Guzmán-Mejía et al., 2021). It is noteworthy that some arthropods, like insects and crustaceans, rely on polysaccharide exoskeletons, particularly chitin, for body support, protection, and mobility in their environment and while hunting for food (Satitsri & Muanprasat, 2020).

Functions of polysaccharides in cellular recognition and signaling

Understanding the role of polysaccharides in cellular recognition and signaling is crucial for unraveling the complexity of cellular interactions and life. Polysaccharides play a vital role in cell recognition by serving as cell surface markers, aiding the identification and differentiation of various cell types. For instance, blood group antigens, which are polysaccharides on the surface of red blood cells, play a crucial role in blood group matching and transfusions (Dotz & Wuhrer, 2016). Additionally, proteins, cells, or other organisms can discern specific properties from polysaccharides present on cell surfaces (Grondin et al., 2022). Polysaccharides also contribute significantly to cellular signaling. Peng, Tang et al. (2022) demonstrated that Angelica polysaccharide can reduce the occurrence of oxidative stress and delay cell aging by activating the silent information regulator 1 (Sirt1)/forkhead box O1 (FoxO1) signaling pathway. Similarly, Lycii fructus polysaccharide, as shown by Peng, Zhao et al. (2022), improves exercise-related fatigue by inhibiting oxidative stress through the activation of the nuclear factor erythroid-related factor 2 (Nrf2)/heme oxygenase-1 (HO-1) signaling pathway. Moreover, polysaccharides can convey cytopathic information. In the context of biosensors for tumor cell identification, Zhang, Wang et al. (2017) discovered abnormal expression of polysaccharides on the surface of cell lesions. Recognizing these abnormal states facilitates a sensitive and rapid disease diagnosis. Polysaccharides can also serve as signal amplifiers in clinical disease detection systems. According to Zhao et al. (2020), a polysaccharide-based electrochemical biosensor exhibited remarkable sensitivity in identifying nonsmall cell lung cancer biomarkers, significantly amplifying the electrochemical signal of the biomarker for improved system detection.

Although extensive literature investigates the function of polysaccharides in cell recognition and information transmission, there is a dearth of applied research in this area. Polysaccharides, often associated with cell pathologies and undergoing varying degrees of changes, hold significant potential as disease indicators. Furthermore, polysaccharides can mitigate various detrimental response processes, including oxidative stress and inflammation. Considering these properties, polysaccharides could be explored for the development of specific drugs that prevent the activation of these detrimental pathways, potentially offering more effective therapeutic effects.

Bioactivity of polysaccharides

The biological activity of polysaccharides encompasses numerous health-promoting functions for organisms, making it a prominent topic in various scientific fields globally. Currently, researchers have investigated several important biological properties of polysaccharides, including antitumor, antiaging, antioxidant, hypoglycemic, anti-inflammatory, antibacterial, and immunoregulatory effects. These bioactive effects position polysaccharides as potential therapeutic substances for diseases that lack treatment options (schematically shown in Figure 3).

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Antitumor activity of polysaccharides

In contemporary medicine, tumor diseases present one of the most challenging issues. Studies have demonstrated that certain polysaccharides, with low toxicity and minimal side effects, can inhibit the growth and spread of tumor cells through various mechanisms (Tian et al., 2016; Zhang, Liu, et al., 2021). These mechanisms include autophagy, induction of tumor cell apoptosis, immune regulation, antiangiogenesis, and antitumor cell metastasis.

Currently, the autophagy-mediated antitumor activity of polysaccharides is emerging as a key mechanism in regulating tumorigenesis and therapy (Hanyu et al., 2020; Zhang, Liu, et al., 2021). For instance, Zhang, Zhou et al. (2021) found that polysaccharides from D. officinale can activate the AMP-responsive protein kinase (AMPK)/mammalian target of rapamycin signaling pathway, promoting excessive autophagy and tumor cell death in CT26 cells. Additionally, polysaccharides can induce apoptosis in tumor cells by regulating apoptosis-related signaling pathways, thereby exerting antitumor activity (Mei et al., 2015). Tian et al. (2016) discovered that a novel neutral polysaccharide isolated from Lentinus edodes induces intrinsic mitochondria apoptosis in HepG2 hepatocellular carcinoma cells through various pathways, including the Bax/Bcl-2 ratio and regulating the phosphatidylinositol 3-kinase (PI3K)/threonine protein kinase (AKT) signaling pathway by inhibiting Akt phosphorylation. Certain polysaccharides, such as those found in A. radix, Ganoderma lucidum, and L. fructus, play a crucial role in inhibiting the occurrence and development of tumor cells by enhancing the immune system through the immunomodulatory mechanisms. These mechanisms include increasing the killing capacity of macrophages and natural killer (NK) cells (Li, Hu, et al., 2020; Wang et al., 2021; Yu et al., 2021). In addition, reports on the antiangiogenic properties of polysaccharides, as a mechanism of their antitumor activity, are gradually increasing. Polysaccharides, during tumor control, can suppress the process of angiogenesis of tumor cells, leading to a lack of nutrients and oxygen, ultimately resulting in tumor cell death (Varghese et al., 2017; Zhou et al., 2018). Additionally, polysaccharides can prevent tumor metastasis and recurrence by precisely blocking the mechanism of tumor cell metastasis to other tissues and organs (Ahmad et al., 2012; Zhang, Zhang, et al., 2022). Lee et al. (2014) found that Inonotus obliquus polysaccharide inhibits the MAPK, PI3K/AKT, and nuclear factor-κB (NF-κB) signaling pathways, reducing the expression of matrix metalloproteinases associated with tumor cell invasion and migration.

It is noteworthy that although many studies report the antioxidant activity of polysaccharides, few have directly linked this property to the antitumor mechanism of polysaccharides. However, combining extensive literature suggests that polysaccharides with antioxidant activity tend to exhibit antitumor effects (Guo, Wang, et al., 2020; He et al., 2019; Wu et al., 2018). This may be attributed to the strong ability of polysaccharides to scavenge free radicals, reducing oxidative stress-induced cellular damage, and enhancing the physiological functions and defense capabilities of normal cells against the invasion and harm from tumor cells. Polysaccharides not only combat tumors through their inherent biological activity but can also synergize with other therapeutic drugs. By enhancing efficacy and eliminating drug resistance, the therapeutic effect on tumor diseases can be further improved (Luo et al., 2022). Wu et al. (2018) found that Astragalus polysaccharides could enhance the antitumor effect of apatinib on gastric cancer cells by inhibiting the AKT signaling pathway. Furthermore, some studies indicate that polysaccharides can exert therapeutic effects on tumors by arresting the cell cycle. Low molecular weight polysaccharides extracted from apples, for instance, can prevent the development of colorectal cancer by affecting the cell cycle (Li et al., 2012). In conclusion, polysaccharides exhibit promising therapeutic potential by manifesting antitumor effects through multiple mechanisms.

Antiaging activity of polysaccharides

Cell cycle arrest stands as a crucial hallmark of cellular senescence, characterized by the progressive decline in cellular function and viability, coupled with the degradation and damage to cellular structures. Interestingly, certain polysaccharides, as natural macromolecules, demonstrate to exhibit antiaging properties primarily through antioxidation, immunomodulation, protection of cellular DNA, and promotion of cell proliferation and repair.

Cellular senescence is significantly influenced by oxidative stress, making the antioxidant mechanism of polysaccharides crucial. Polysaccharides scavenge free radicals and enhance the activity of antioxidant enzymes, thereby inhibiting the cellular aging process (Pu et al., 2015). According to Tong et al. (2018), sulfated modification of Bupleuri radix polysaccharides reduces cellular oxidative stress damage and significantly inhibits the expression of the p53-p21 and p16-pRb pathways, which are closely related to the cellular senescence process. Immunomodulation also plays a significant role in the antiaging activity of polysaccharides (Liu et al., 2019; Nigam et al., 2022; Zhang, Hu, et al., 2020). The decline in immunological function, a hallmark of aging, manifests in the senescence of immune cells with reduced proliferative activity and weakened clearance ability, resulting in the accumulation of senescent cells and triggering a decline in organism functions and inflammatory responses. Encouragingly, some polysaccharides can effectively enhance the proliferative vigor of immune cells by suppressing the activation of AMPK in these cells (Akbar et al., 2016; Zhao, Liu, et al., 2021). Furthermore, the protection of cellular DNA by inhibiting DNA oxidation and repairing DNA damage is a key mechanism for the antiaging activity of polysaccharides (Getachew & Chun, 2017). It is well known that DNA damage and mutations are known causes of cellular aging, and polysaccharides can help cells detect DNA damage and initiate DNA repair programs (D'adda Di Fagagna, 2008). Additionally, polysaccharides have been shown to promote cell proliferation and repair mechanisms, which firmly establishes their antiaging properties. Tang et al. (2022) have demonstrated that Desmodium styracifolium polysaccharide carboxymethylation modification possesses antiaging activities such as promoting cell proliferation.

Although reports on the antiaging activity of polysaccharides are mostly limited to the antioxidant mechanism, this focus may hinder the broader application of their antiaging efficacy. The potential connection of polysaccharide anti-inflammatory activity to the antiaging process, beyond the previously mentioned mechanisms, warrants further exploration. Oxidative stress and immunomodulatory pathways often involve the expression of inflammatory factors, suggesting the possible involvement of polysaccharide anti-inflammatory activity in the whole process.

Antioxidant activity of polysaccharides

Oxidative stress refers to the disruption of intracellular oxidative reactions, leading to the excessive production of intracellular oxidized substances, such as free radicals. This production surpasses the cellular antioxidant capacity, triggering a cascade of physiological and pathological responses (Zhu et al., 2018). Currently, polysaccharides are a focal point in antioxidant activity research. Related studies suggest that the antioxidant mechanisms of polysaccharides encompass scavenging free radicals, inhibiting the generation of oxidized free radicals, promoting the activity of antioxidant enzymes, and enhancing the stability of antioxidant substances.

The primary mechanism through which polysaccharides exert antioxidant effects is the scavenging of free radicals. Free radicals, unstable atomic groups with high activity values, cause various damaging behaviors in living organisms (Yu-Hao et al., 2021), making them a major contributor to oxidative damage. Polysaccharides can directly scavenge free radicals or indirectly regulate the antioxidant enzyme system, hereby exhibiting antioxidant effects (Zhu et al., 2018). For instance, Wang et al. (2018) found that Ginger polysaccharide has a strong scavenging effect on intracellularly generated 2, 2-diphenyl-1-picrylhydrazyl radicals, indicating high antioxidant activity. Additionally, some polysaccharides exert antioxidant activity by inhibiting the production of oxidizing free radicals (Farah et al., 2018; Sheu et al., 2018). Liu, Ye et al. (2021) demonstrated that Lignified okra polysaccharide protects cells by inhibiting mRNA expression of iNOS and reducing NO secretion. Polysaccharides can enhance the antioxidant capacity of organisms by promoting the activity of antioxidant enzymes, including superoxide dismutase, catalase, and glutathione peroxidase (Peng, Tang, et al., 2022). Liao et al. (2019) found that a novel polysaccharide isolated from okra could increase the levels of antioxidant enzymes and decrease malondialdehyde (lipid peroxidation marker), inhibiting free radical generation and exerting antioxidant activity. Additionally, polysaccharides can improve the stability of antioxidant substances, prolonging the duration of their antioxidant activity. For example, Guo et al. (2022) found that stabilizing complexes of soluble Soybean polysaccharides with wheat alcohol soluble protein nanoparticles, combined with curcumin, enhanced the stability and antioxidant properties of the latter.

Hypoglycemic activity of polysaccharides

Hyperglycemia has become a pressing issue, and maintaining blood glucose levels within a stable range is crucial for normal organ function. Polysaccharides contribute to lowering blood glucose levels through various mechanisms inhibiting glucose-related synthase activity, promoting glycogen synthesis, inhibiting gluconeogenesis, promoting glucose catabolism, and promoting insulin secretion. The main factor contributing to elevate blood glucose is the massive production of glucose and its absorption into the bloodstream. Thus, blocking the enzyme activity mechanism linked to glucose synthesis is one way to lower blood glucose. Zhang, Zhao et al. (2020) found that Red clover polysaccharides reduce postprandial blood glucose concentration by inhibiting α-amylase and α-glucosidase activities crucial to glucose synthesis. Similarly, polysaccharides from mulberry, Sargassum pallidum (Phaeophyceae), and licorice have shown similar effects (Chen et al., 2015; Pan et al., 2020; Ren et al., 2017). Furthermore, disrupting pathways like glycogenolysis and gluconeogenesis, which increase blood glucose levels, can lead to hypoglycemia. However, some polysaccharides have been found to stimulate the synthesis of glycogen and suppress glycogenolysis and gluconeogenesis. Liu et al. (2020) reported that D. officinale polysaccharides increase liver glycogen synthesis and reduce its decomposition, hindering gluconeogenesis by affecting the glucagon-mediated cAMP/protein kinase A (PKA) and Akt/FoxO1 signaling pathways. Cao, Li et al. (2019) demonstrated that S. pallidum polysaccharides reduce blood glucose concentration by promoting glycogen synthesis and increasing the activities of key enzymes, hexokinase, and pyruvate kinase in the glycolysis process. In a similar vein, Liu et al. (2017) found that sulfated Chlorophyta polysaccharides accelerate glucose metabolism by promoting the expression of AMPKα2, a major isoform of AMPK. Apart from various regulatory factors aiding in glucose metabolism, the most direct mechanistic pathway to rapid blood glucose depletion involves the promotion of insulin secretion. Yang et al. (2021) discovered that Bee pollen polysaccharides stimulate the proliferation of pancreatic β-cells by activating the MAPK and AKT pathways related to cell proliferation. Simultaneously, they regulate key transcription factors, Maf A and Pdx1, involved in the expression and secretion of the insulin gene in β-cells. This dual mechanism results in an increase in insulin secretion, ultimately manifesting the hypoglycemic activity of polysaccharides.

In addition to these mechanisms, polysaccharides can form a gel in the stomach, reducing the rate of gastric emptying and delaying the absorption of simple sugars (Nagarwal & Pandit, 2008). They are also digested and absorbed relatively slowly, releasing blood sugar gradually and avoiding drastic fluctuations in blood glucose levels. The fiber in polysaccharides lowers the overall absorption of simple sugars, increasing gastrointestinal peristalsis and promoting food movement and excretion. Furthermore, some components in certain polysaccharides inhibit sugar absorption in the intestine, further reducing the rise in blood glucose levels (Garcia-Barrios et al., 2013).

Anti-inflammatory and antiviral activity of polysaccharides

Inflammatory response is a common physiological reaction of tissue damage or infection. Recent studies on polysaccharides with anti-inflammatory properties have focused on modulating pro- and anti-inflammatory mediators and cell signaling (Bai et al., 2020). The balance between pro- and anti-inflammatory mediators in tissue cells determines the course of inflammation. Polysaccharides possess the ability to either increase the production of anti-inflammatory mediators or inhibit pro-inflammatory mediators, thereby suppressing the inflammatory response and reducing tissue damage (Jen et al., 2021; Wang, Wang, et al., 2020). Shang et al. (2021) reported that Dendrobium huoshanense stem polysaccharide promoted the expression of IL-10 and TGF-β1, while significantly inhibiting pro-inflammatory factors such as IL-1β, IL-6, IL-17, TNF-α, and GM-CSF, slowing the progression of arthritis in mice. Numerous studies have demonstrated that polysaccharides exhibit anti-inflammatory properties by regulating cell signaling pathways and inhibiting the activation of inflammation-related signaling pathways, such as NF-κB, MAPK, PI3K/AKT, and Janus kinase 1/signal transducers and activators of transcription 3 (Fang et al., 2020; Jayawardena et al., 2020; Wang, Lai, et al., 2019; Wang et al., 2022; Xie et al., 2019). For instance, Wang, Lai et al. (2019) revealed that Phallus indusiatus (formerly Dictyophora indusiata) (Basidiomycota) polysaccharide exerts anti-inflammatory activity by inhibiting the toll-like receptor (TLR264)/NF-κB signaling pathway and blocking the activation of the dNLRP7 inflammasome. Jayawardena et al. (2020) pointed out that sulfation-modified Sargassum polysaccharides have anti-inflammatory effects in inflammation-related diseases by inhibiting TLR-mediated myeloid differentiation factor 88 and inhibitor of kappa B kinase complex expression, thereby lowering NF-κB and MAPK activation.

Viral infections pose significant risks to both human and animal health. Some polysaccharides have proven effective in combating viruses with minimal negative effects on the host cells. The antiviral mechanism of polysaccharides involves interference and inhibition of viral infection and replication processes, including destroying virus structure, inhibiting virus binding and invasion of host cells, inducing host cell antiviral defense, interfering with viral replication and transcription, and regulating the body's inflammatory response (Lin et al., 2016). Jones et al. (2020) found that cyclodextrins can induce deoxyribonuclease to degrade the herpes simplex virus genome, inactivating the virus. Chiu et al. (2012) reported that kappa carrageenan can bind to viruses, forming carrageenan-virus complexes and preventing virus attachment to host cells. Tian et al. (2017) demonstrated that Lnonotus obliquus (Basidiomycota) polysaccharides directly inhibit viral particles and may exert antiviral activity by affecting the process of viral binding and invasion of cells. Huan et al. (2022) found that Plantago asiatica polysaccharides block the adsorption of pseudorabies virus particles to the cell surface, reducing penetration into tissue cells and activating the host's antiviral defense mechanism. Terasawa et al. (2020) reported that rhamnan sulfate can inhibit the early steps of influenza virus infection and protect cells by triggering the generation of virus-specific antibodies. Feng et al. (2013) identified novel Radix cyathulae polysaccharides that significantly enhance the phagocytosis of organismal macrophages, increase the activity of NK cells and cytotoxic T lymphocytes, as well as the related antibody titer. Additionally, it has been reported that interfering with viral replication and transcription is an important mechanism by which polysaccharides exert antiviral activity, hindering the synthesis of viral genetic material and inhibiting the translation of viral proteins (Su et al., 2016). Chen et al. (2014) found that Bush sophora root polysaccharides could significantly reduce the virus level in the blood of diseased ducks by inhibiting the replication and release of duck hepatitis virus.

Additionally, polysaccharides can protect host tissues by modulating the inflammatory response of host cells and reducing inflammatory damage caused by viral infections. Wan et al. (2022) have confirmed that Inulin polysaccharides inhibit the expression of TLR3/4 mRNA, reduce pro-inflammatory factors TNF-α and TNF-β, and attenuating virus-induced inflammatory injury. It is important to note that the antiviral activity of polysaccharides may be influenced by factors such as polysaccharide structure, concentration, and host cell type. Therefore, the specific antiviral mechanisms of polysaccharides require further exploration.

Anticoagulant activity of polysaccharides

Excessive coagulation can lead to pathological disorders such as thrombosis, posing serious harm to the organisms. Polysaccharides have demonstrated anticoagulant effects through mechanisms like inhibition of anticoagulant factor activation, prevention of platelet aggregation, protection of vascular endothelial cells, and suppression of thrombus fibrin formation. For instance, heparin, a widely used anticoagulant drug, inhibits coagulation factor activation by binding to antithrombin III (AT-III) and promoting AT-III's binding to coagulation factors IIa and Xa (Ofosu, 1988). Shang et al. (2018) found that fucoidan extracted from sulfation-modified sea cucumber possessed potent anticoagulant capacity as it significantly inhibited the activation of endogenous coagulation factor Xa, showcasing potent anticoagulant capacity. Polysaccharides can mitigate thrombosis by reducing platelet adhesion and aggregation. Carboxymethylglucan has been shown to reduce platelet aggregation behavior and improve vascular function (Bezerra et al., 2022). A novel Auricularia auricula-judae (formerly Auricularia auricula) (Basidiomycota) polysaccharides demonstrated the potential to be an anticoagulant drug by promoting the expression of endothelial nitric oxide synthases and prostacyclin, while inhibiting procoagulant factors such as endothelin-1 and thromboxane B2 (Bian et al., 2022). Furthermore, polysaccharides protect vascular endothelial cells, vital for maintaining vascular function and blood flow. Astragalus polysaccharides induced macrophage polarization, alleviating vascular endothelial dysfunction (Bezerra et al., 2022). Sha et al. (2023) found that Astragalus polysaccharides induced macrophage polarization toward the M1 phenotype by modulating the Nrf2/HO-1 signaling pathway, thereby alleviating the problem of vascular endothelial dysfunction in the body. Parnigoni et al. (2022) found that hyaluronan enhances the ability of vascular endothelial cells to dedifferentiate, migrate, and proliferate, maintaining the protective effect of the vascular wall on blood flow. In addition, polysaccharides can interfere with thrombus formation by interfering with the polymerization and stabilization of thrombus fibrin, promoting the activation of the fibrinolytic system and accelerating thrombus dissolution (Cao, He, et al., 2019).

Ongoing exploration of polysaccharide anticoagulant activity suggests that inhibiting the inflammatory response could be a potential mechanism. Inflammatory response is a significant factor in thrombosis (Wojta, 2020), and polysaccharides with demonstrated anti-inflammatory properties may reduce the occurrence of the coagulation process by preventing the release of inflammatory factors and suppressing the activation of inflammatory cells.

Immunomodulatory activity of polysaccharides

Polysaccharides possess immunomodulatory properties, allowing them to regulate the immune system's function for disease treatment and prevention. The mechanism primarily involves the regulation of immune cell activation, proliferation, and the balance of immune responses, ultimately enhancing the immune system's function.

The role of polysaccharides in regulating immune cell activation is well documented. For instance, Astragalus polysaccharides have been shown to activate macrophages, induce apoptosis, and arrest the cell cycle in the G4 phase, thereby inhibiting the growth of cancer cells (Li, Hu, et al., 2020). Moreover, research on polysaccharides enhancing immune stimulation and promoting the body's immune response has been expanding. Codium fragile (Chlorophyta) polysaccharides, as investigated by Park et al. (2020), can directly activate NK cells or through cytokine-mediated means. This activation leads to the release of CD69, IFN-γ, and cytotoxic mediators, fostering the apoptosis of target cells and enhancing the immune response. Polysaccharides also exhibit immunosuppressive effects. G. lucidum (Basidiomycota) polysaccharides, according to Wei et al. (2018), mitigate acute colitis episodes in mice by suppressing the immune response. This includes inhibiting the secretion of pro-inflammatory cytokines and modulating B-cell, Th17 cell, NK cell, and NKT cell populations. Furthermore, the anti-inflammatory and antioxidant activities of polysaccharides, highlighted in previous studies, contribute significantly to their immunomodulatory functions. These findings underscore the substantial potential of polysaccharides in drug development (Liu, Dong, et al., 2023; Sun et al., 2023; Wang, Yan, et al., 2023).

Other biological activities of polysaccharides

In recent years, research on polysaccharides has made significant progress across various disciplines. Beyond the well-known biological activities, polysaccharides have emerged as effective prebiotics, influencing the intestinal microbiota, safeguarding the intestinal mucosa, and elevating short-chain fatty acid levels. Wang et al. (2021) demonstrated that L. fructus polysaccharides can strengthen the immunity system by increasing the diversity and relative abundance of intestinal microbiota. In a study by Fu et al. (2018), Codonopsis polysaccharides were found to modulate the gut microbiota in experimental mice, promoting the production of short-chain fatty acids. This modulation plays a crucial role in repairing immune damage to the membrane and inhibiting colonization of pathogenic bacteria. Ge et al. (2021) observed that polysaccharides influence the microbial population and metabolic activity in the intestinal tract by fostering the formation of short-chain fatty acids. This coordination supports the normal physiological functions of the colon, contributing to overall host health. Furthermore, some polysaccharides exhibit notable antiradiation properties. Yuan et al. (2020) discovered that Aloe vera polysaccharides mitigate radiation damage to the skin caused by UV rays through the Keap12/Nrf1/ARE signaling pathway. Wang, Xue et al. (2020) reported that polysaccharides from various sources can exert positive radiation protection effects on the human body.

APPLICATION OF POLYSACCHARIDES

Polysaccharides, owing to their intricate structure and multifunctionality, offer a broad spectrum of potential applications. Various industries, including medicine, the food industry, and industrial production, have embraced the utilization of polysaccharides due to ongoing exploration into their diverse properties, summarized in Table 2.

TABLE 2 Applications of polysaccharides in various fields.

Realm	Polysaccharides	Mechanism	Effects	References
Medicine	Chinese medicine polysaccharide; chitosan; polysaccharide-based hydrogels; vegetable collagen; sodium alginate; dextran	Mechanisms of synergistic regulation of multibiotic activities; alter structure and physical properties; promotes cell proliferation and differentiation; interacts with specific biomolecules or cells	Improves its ability to fight diseases; protects the drug and improves its stability; enabling targeted drug delivery and controlled release; promote tissue regeneration and repair; building biosensors and diagnostic reagents	Ali and Ahmed (2018), Baei et al. (2023), Bayer (2020), Kim et al. (2018), Mussi-Pinhata et al. (2022), Qasemi and Ghaemy (2020), Tan et al. (2022)
Food processing	Pectin; chitosan; dietary fiber; prebiotics	Increase the viscosity of the food; increase food stability; promote intestinal peristalsis and maintain intestinal flora balance	Improvement of food texture and taste; extending the shelf life of food; maintains intestinal flora balance and enhances immunity	Hu and Gänzle (2019), Kim et al. (2019), Ścieszka and Klewicka (2019, 2020), Thakur et al. (1997)
Textile industry and beauty industry	Mannan; chitosan; algal polysaccharide;	Forms a stable chemical bond with the fiber surface; absorbs and retains moisture in the skin; promotes collagen synthesis; inhibits melanin production	Improvement of color fastness of textiles; prevents dry skin; increase skin elasticity and firmness; brightens and evens out the skin	Costa, Silva et al. (2018), Feng et al. (2021), Hu et al. (2019), Juncan et al. (2021), Sharma and Baldi (2016)
Environmental protection	Chitosan; cyclodextrin; polysaccharide-based hydrogels; alginate; tapioca starch; hyaluronic acid; hemicellulose	By adsorption and precipitation; good water absorption; preparation of biodegradable plastics; fermentation produces bioenergy	Water purification; adsorption and storage of large quantities of water; protecting the environment; providing sustainable energy support	Cann et al. (2020), Liu, Du et al. (2021), Martínez-Cano et al. (2022), Matheus et al. (2023), Zhang, Wang, Fan et al. (2022), Zhao, Liu et al. (2023)

Application of polysaccharides in medicine

Polysaccharides serve as excellent raw materials in the pharmaceutical industry due to their biological activity, exhibiting positive therapeutic effects on specific diseases with minimal side effects. Existing drugs containing polysaccharides encompass a range of categories such as antitumor drugs, antidiabetic drugs, anticoagulants, anti-inflammatory drugs, antiviral drugs, and vaccines (He et al., 2019; Homaeigohar et al., 2023; Mussi-Pinhata et al., 2022; Onishi et al., 2016; Xiang et al., 2021; Zhang, Zhang, et al., 2019). Notably, most polysaccharides are not single-target therapeutic agents, and they often comprise a complex of multiple active components that collectively enhance the body's immunity, establishing a multisystem coregulatory mechanism. Therefore, drugs with polysaccharides typically have longer therapeutic periods and more intricate therapeutic pathways, making them less suitable for sudden onset diseases. Currently, most polysaccharide drugs in the market fall under the category of healthcare products.

Further development of polysaccharide drug production technology is still needed, as processing more polysaccharides into suitable drugs poses challenges due to purification difficulties and the requirement for stringent preservation methods. Research is ongoing to discover novel approaches, with some studies suggesting that traditional Chinese medicine decoction processes could serve as a valuable reference for processing polysaccharide-based medicines (Chen et al., 2020). The long-established history of producing decoctions from traditional Chinese medicine highlights the potential benefits of imitating relevant processing techniques to enhance the efficacy of polysaccharides (Cao et al., 2020). Additionally, polysaccharides show promise as carriers or coating materials for drug delivery (Nai et al., 2021). Their excellent biocompatibility ensures the protection and stability of drugs. Polysaccharides’ surface properties, rich in reactive groups and variable molecular weights, can be modified for targeted delivery and controlled release, influencing the rate and pathway of drug release (Bayer, 2020). Examples include polysaccharide-based drug delivery systems like hydrogels, microspheres, and nanoparticles. These systems protect drugs from degradation and excretion, with CS nanoparticles showcasing the potential for targeted drug delivery to tumor tissue (Ali & Ahmed, 2018). In recent advancements, researchers have developed drug carriers responding to pH, temperature, ionic strength, and magnetic fields through polysaccharide modifications (Lupu et al., 2023). Polysaccharides play a crucial role in extending the circulating half-life of drugs, reducing total doses, and enhancing drug bioavailability (Karmali et al., 2012). In addition, polysaccharides find extensive use in tissue engineering for constructing artificial tissues and organs. Polysaccharide scaffold materials provide the necessary support structure for cell adhesion, growth, and promote cell proliferation and differentiation (Baei et al., 2023). Commonly used materials like collagen and sodium alginate aid in the construction of artificial bone, cartilage, and skin, fostering tissue regeneration and repair due to their high bioactivity and biocompatibility (Kim et al., 2018). Biosensor technology, pivotal in clinical diagnostics, benefits from polysaccharides in the construction of biosensors and diagnostic reagents. Polysaccharides interact with specific biomolecules or cells, facilitating the detection and analysis of biomolecules or cells (Qasemi & Ghaemy, 2020). For instance, dextran interacts with specific components of bacterial cell walls for bacterial detection and identification (Nakamura et al., 2023). Polysaccharides also contribute to highly sensitive detection and quantitative analysis of specific molecules by modulating the surface properties and structure of nanoparticles (Deng et al., 2022). In conclusion, polysaccharides continue to demonstrate a wide array of applications in medicine, with ongoing innovations and advancements in their utilization.

Application of polysaccharides in food processing

The utilization of polysaccharides in food processing is an extensively researched field. Incorporating polysaccharides can alter the physicochemical properties of foods, including rheological properties, stability, texture, and nutritional value (Wang, Wang, et al., 2023), aligning with consumers’ expectations for diverse food products. One of the most common applications of polysaccharides in food is as thickeners and viscosity enhancers to improve the texture and mouthfeel (Bernaerts et al., 2019). For instance, in the production of jams and jellies, pectin is employed to improve viscosity and coagulability, resulting in a more appealing taste and texture (Thakur et al., 1997). Polysaccharides are also utilized to modulate the rheological properties of foods, such as viscosity, flowability, and rheological characteristics (Fan et al., 2022). Adjusting the type and quantity of added polysaccharides allows for control over food flow, cohesion, and stability, particularly in processing liquid foods, sauces, and soups. Furthermore, the addition of polysaccharides can enhance the stability of food products, preventing ingredient separation, precipitation, or segregation (Ścieszka & Klewicka, 2019). Polysaccharides create a colloidal structure that envelops and shields other ingredients from the external environment, thereby prolonging the shelf life of the food while preserving its original texture and taste (Bose et al., 2021; Zhang, Lan, et al., 2022). Notably, CS serves as a widely employed preservative, inhibiting microbial growth in foods and preserving freshness and taste, making it a common choice for products like dried fruits, candied fruits, and candies (Hu & Gänzle, 2019). Similarly, seed-derived polysaccharides function as preservatives, enhancing food safety and durability (Abbasi et al., 2023; Hajji et al., 2021). Moreover, the introduction of polysaccharides can effectively prevent the precipitation and sedimentation of proteins in food products, maintaining stability and quality (Li et al., 2023). This has positive implications for body weight control and reduction of chronic diseases such as obesity and diabetes. Additionally, the incorporation of polysaccharides enhances the taste and flavor of food. Polysaccharides contribute to increased nutritional value in foods, promoting digestive health. Specialized polysaccharides, such as fiber and prebiotics, play distinct roles—fiber increases the fiber content, promotes intestinal peristalsis, and helps prevent conditions like intestinal inflammation (Kim et al., 2020). Probiotics foster the growth of beneficial bacteria, maintain balance, and strengthen immunity (Kim et al., 2019). In conclusion, polysaccharides exhibit a wide array of applications in food processing. However, it is crucial to adjust the amount and method of their use based on the specific characteristics of foods and processing requirements to ensure food quality and safety. Future research can delve deeper into the application of polysaccharides in food processing, developing more functional polysaccharides that contribute to the ongoing evolution of the food in the industry.

Polysaccharide applications in the textile and cosmetic industries

In the textile industry, the primary focus of polysaccharide application lies in dyes and printing agents. Research has demonstrated that polysaccharides can create a stable chemical bond with the fiber surface, ensuring strong adherence of dyes to fibers and enhancing textile washability and color fastness (Sharma & Baldi, 2016). Mannan addition to dyes has proven effective in significantly increasing their fastness and adhesion to cotton fibers (Singh et al., 2018). Furthermore, polysaccharides play a role in regulating the permeability and diffusivity of dyes, ensuring even distribution within the fiber and thereby improving the overall dyeing effect (Bezerra et al., 2020). Beyond this, polysaccharides find application as wrinkle inhibitors and softeners for textiles (Özen et al., 2023). Recent findings also suggest that polysaccharides can contribute to textiles achieving antibacterial effects. As a crucial line of defense protecting the skin, textiles with added CS exhibit biological activity against various skin pathogens, potentially offering antibacterial benefits when incorporated into the textile process (Costa, Silva, et al., 2018).

In the beauty industry, polysaccharides are widely employed in skin care products and cosmetics. Their robust moisturizing properties play a key role in preventing skin dryness and water loss (Juncan et al., 2021). López-Hortas et al. (2021) noted a significant enhancement in skin water content and moisturizing capacity through the addition of seaweed polysaccharides to skincare products. Additionally, owing to their strong adhesive properties, polysaccharides serve as effective binders and stabilizers, ensuring even distribution of cosmetic ingredients and enhancing viscosity and stability (Shariatinia, 2018). Furthermore, studies indicate that polysaccharides contribute to collagen synthesis, thereby improving skin elasticity, firmness, and reducing the appearance of fine lines and wrinkles (Feng et al., 2021). Polysaccharides also exhibit antiacne and whitening properties. Research by Hu et al. (2019) demonstrated that polysaccharides inhibit melanin, resulting in brighter and more even skin tone by reducing discoloration and freckles. As research on polysaccharides progresses, their applications in the textile and cosmetic industries are anticipated to expand and evolve.

Application of polysaccharides in environmental protection

As a renewable and biodegradable natural material, polysaccharides play a crucial role in the realm of environmental protection. Existing studies showcase the remarkable efficacy of polysaccharides in sewage treatment and soil remediation. Polysaccharides effectively remove heavy metal ions from sewage and soil through adsorption and precipitation, contributing to the restoration of a healthy environmental state (Fenyvesi & Sohajda, 2022; Liu, Du, et al., 2021; Zhao, Liu, et al., 2023). Additionally, polysaccharides can be crafted into hydrogel materials with excellent water-absorbing properties, providing a promising avenue for water resource protection. These hydrogel materials can absorb and store significant amounts of water, curbing water resource wastage and holding significant application potential (Martínez-Cano et al., 2022).

In the realm of food packaging, the use of polysaccharides is a pivotal aspect of environmental protection. Traditional plastic packaging materials often pose environmental pollution risks, whereas biodegradable plastics crafted from polysaccharides, such as starch, offer an eco-friendly alternative. These biodegradable plastics can fully break down into environment-friendly substances, mitigating environmental pollution (Nešić et al., 2019; Zhu, 2021). For instance, the utilization of polysaccharides like starch in food packaging not only preserves food freshness and hygiene but also minimizes the environmental impact of plastic waste (Matheus et al., 2023). Furthermore, employing polysaccharides in personal care products aligns with environmental protection goals. Many conventional personal care products feature chemically synthesized ingredients, which can be detrimental to both human health and the environment. In contrast, the use of polysaccharides allows the formulation of natural personal care products, including shampoos, body washes, and face masks. These products boast good biocompatibility and biodegradability, presenting a sustainable and eco-friendly choice that avoids adverse effects on both humans and the environment. Additionally, natural sunscreens prepared with polysaccharides effectively shield against the damaging effects of ultraviolet rays, promoting skin health (Zhu, 2021; Zhu et al., 2018). In textiles and home furnishings, integrating polysaccharides is a vital part of environmental protection. Traditional textiles and home furnishings often incorporate chemically synthesized fibers and additives, posing risks to both human health and the environment. Polysaccharides, characterized by excellent biocompatibility and biodegradability, offer an alternative for crafting natural fibers and additives, such as natural cellulose fibers and natural dyes. This shift ensures products that do not adversely impact human health or the environment (Zhang, Wang, Fan, et al., 2022). Finally, polysaccharides play a pivotal role in the energy sector as a means of environmental protection (Pfrengle, 2017). Traditional energy sources reliant on fossil fuels contribute significantly to environmental pollution. In contrast, polysaccharides can be fermented to produce bioenergy, including bioethanol and biogas, offering environment-friendly alternatives to fossil fuels and reducing greenhouse gas emissions (Cann et al., 2020). Additionally, polysaccharides can be utilized in the fabrication of new energy devices like solar and fuel cells, providing sustainable energy solutions to support environmental protection (Lin et al., 2021). In summary, polysaccharides exhibit a diverse range of potential applications in environmental protection and contribute significantly to sustainable development.

CONCLUDING REMARKS AND OUTLOOK

The field of polysaccharide research has seen significant growth in recent years, with researchers exploring the potential of this natural, nontoxic substance across various domains of life. This article offers a comprehensive review delving into the fundamental characteristics, synthesis and degradation, biological functions, and potential applications of polysaccharides, as illustrated in Figure 4. Furthermore, it discusses the mechanisms behind the biological functions of polysaccharides, aiming to aid researchers in their initial understanding and provide a theoretical foundation for future exploration.

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Polysaccharides exhibit immense promise, particularly in pharmaceutical production—a primary focus of ongoing research. However, many types of polysaccharides remain underutilized in clinical production, with research reports often concentrated on basic theories. This trend guides the direction for future research endeavors. Moreover, it is crucial to acknowledge the significant role polysaccharides can play in environmental protection efforts. Some studies have successfully integrated polysaccharides into low-carbon living practices, showcasing valuable applications beyond their current utilization. The inherent biocompatibility and degradability of polysaccharides position them as valuable assets for environmental preservation.

Our research endeavors aim to contribute to the development of polysaccharide-based drugs and their applications in daily life. By harnessing the inherent properties of polysaccharides, we anticipate a meaningful impact on both pharmaceutical development and environmental sustainability.

AUTHOR CONTRIBUTIONS

Bo-wen Xu and Sai-sai Li: Data curation; writing—original draft preparation; investigation. Wen-li Ding: Resource; validation; methodology. Cai Zhang: Data curation; investigation. Mujeeb Ur Rehman: Investigation; validation. Muhammad Farooq Tareen: Resource; investigation. Li Wang: Resource; validation. Shu-cheng Huang: Supervision; writing review and editing; project administration; funding acquisition.

ACKNOWLEDGMENTS

This work was financially supported by the National Natural Science Foundation of China (No. 32202876), the China Postdoctoral Science Foundation (No. 2023T160198), and the Key Scientific and Technological Project of Henan Province Department of China (No. 232102111046).

CONFLICT OF INTEREST STATEMENT

The authors confirm that they have no conflicts of interest to declare for this publication.

DATA AVAILABILITY STATEMENT

No data was used for the research described in the article.

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