key: cord-0708694-myj7dwef
authors: Hassan. E. Konozy, Emadeldin; El-fadil M. Osman, Makarim; Ibrahim Dirar, Amina
title: Plant Lectins as Potent Anti-coronaviruses, Anti-inflammatory, Antinociceptive and Antiulcer Agents
date: 2022-04-22
journal: Saudi J Biol Sci
DOI: 10.1016/j.sjbs.2022.103301
sha: 7d5bfc83115b0e749b59a219a0906fb96b2059a5
doc_id: 708694
cord_uid: myj7dwef

Lectins are defined as carbohydrate-binding proteins/glycoproteins of none immune origin, they are ubiquitous in nature, exist from bacteria to human cells. And due to their carbohydrate-binding recognition capacity, they have been a useful biological tool for the purification of glycoproteins and their subsequent characterization. Some plant lectins have also been revealed to own antinociceptive, antiulcer, and anti-inflammatory properties, where these features, in many instances, depending on the lectin carbohydrate-binding site. Coronavirus disease of 2019 (COVID-19) is a respiratory disease that struck the entire world leaving millions of people dead and more infected. Although COVID-19 vaccines have been made available, and quite a large number of world populations have already been immunized, the viral infection rates remained in acceleration, which continues to provoke major concern about the vaccines' efficacy. The belief in the ineffectiveness of the vaccine has been attributed in part to the recurrent mutations that occur in the epitope determinant fragments of the virus. Coronavirus envelope surface is extensively glycosylated being covered by more than sixty N-linked oligomannose, composite, and hybrid glycans with a core of Man3GlcNAc2Asn. In addition some O–linked glycans are also detected. Of these glyco-chains, many have also been exposed to several mutations, and a few remained conserved. Therefore, numerous plant lectins with a specificity directed towards these viral envelope sugars have been found to interact preferentially with them and are suggested to be scrutinized as a possible future tool to combat coronaviruses including severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) through blocking the viral attachment to the host cells. In this review, we will discuss the possible applications of plant lectins as anti-coronaviruses including SARS-CoV-2, antinociceptive, anti-inflammatory, and antiulcer agents with the proposed mechanism of their actions.

Emadeldin Hassan. E. Konozy (EHEK): Draft the idea and wrote the initial draft of the manuscript.

Makarim El-Fadil M. Osman (MEMO): Assisted in writing and editing the manuscript. Drew the figures 3.

Amina Ibrahim Dirar (AID): Assisted with drafting the basic idea and revised the manuscript.

Cell surface glycoconjugates coat play a crucial role in several biological paths, such as cell-cell adhesion, inflammatory relocation, host-pathogen interactions, immune response initiation, and cancer metastasis ( (Colgan et al. 1995; Gorelik et al. 2001; Hevey 2019; Nardy et al. 2016) , and references therein). These processes are collectively mediated through carbohydrate-binding proteins that span the surface of the opposing cells (Brandley and Schnaar 1986) .

Since ancient times, and to date plant parts such as leaves, barks, seeds, and roots have been used by human communities in the prevention and treatment of diseases, and for healthcare (Jones et al. 2018) . In Africa and Asia over 90% of folk medicine protocols are comprised of plants, and the majority of these remedies are used by villagers or nomads who are often delocalized, and away from governmental medical facilities access. These plant products have been routinely used for treatments of various ailments such as infectious diseases caused by bacteria, fungi, and viruses, as well as non-communicable diseases including heart disease, cancer, diabetes, chronic lung conditions, etc. (Sofowora et al. 2013) . It is generally thought that the presently in-use analgesiainducing medications such as opioids and Non-steroidal anti-inflammatory drugs (NSAIDs) are not suitable for many patients due to their side effects and low efficiency (Ahmadiani et al. 1998 ).

Consequently, the search for new drugs becomes necessary.

The fortuitous discovery made by the German doctoral student Peter Hermann Stillmark in 1888 that the castor bean (Ricinus communis) extract can agglutinate erythrocytes, a mechanism confirmed later to occur through erythrocytes' surface glycoconjugates, had signified the milestone for our existing understanding of plant carbohydrate-binding protein in particular, and its animal/microbial counterparts in general (Sharon and Lis 2004) . These glycan interacting proteins were then termed lectin, "Legere" in Latin which means to choose or pick up. In subsequent years research has confirmed the presence of such proteins not merely in plant and animal cells but also in bacteria, viruses, yeast, and parasites where they could facilitate the attachment processes of these microbes to the glycoproteins and glycolipids, and permeate the host cell surface (Sharon and Lis 1997) . By their feature or capacity of being able to interact with glycans, these proteins have been defined as any multivalent protein/glycoprotein that possesses at least a single non-catalytic domain that could interact reversibly with sugars or carbohydrates and thereby causes agglutination of the cells (Goldstein et al. 1980 ). Owing to their being furnished with such unique carbohydrate-binding site(s) lectins are assigned to perform many biological functions such as endocytosis, act as intracellular transport vehicles for glycoproteins, and regulate the protein content in the blood ((Dias et al. 2015) and the references cited therein).

Nevertheless, animal lectin was discovered before plant lectin in 1872 but they were not recognized as glycan-binding protein (Kilpatrick 2002) . Although animal and plant lectins share no apparent primary structure resemblances, they share the aptitude to interact and recognize specific glycan receptors emphasizing the role of these proteins in molecular recognition (Reyes-Montaño and Vega-Castro 2018). Animal and microbial lectins are found in comparatively lesser amounts as compared to the plant lectins, the latter being detected almost in every part of the plant; seeds, leaves, bark, stem, flower, roots, etc (Mishra et al. 2019) . Although not all plants contain lectins, however, when present they may account for up to 10-12% of the total seeds soluble protein (Roopashree et al. 2006; Sathe and Deshpande 2003) . The abundance of plant lectin (Spilatro et al. 1996) , their ease to isolate (Mishra et al. 2019) , and the accelerated advancement in preparation of affinity chromatographies that facilitated the purification of plant lectin in a single or two steps (Freeze 1995; O'Connor et al. 2017) , have assisted in performing deep studies to solve their structures' ambiguity, possible biological effects, and clinical applications. While many plant lectins are sharing primary and secondary structures, they exhibit different biological effects which are probably ascribed to their varying glycan recognition specificities. Moreover, scientific proofs that present some plant lectins with antinociceptive, anti-inflammatory, antioxidant, and gastroprotective properties are also accumulating. While others are recognized to inhibit many microbes like viruses, parasites, nematodes, and bacteria (Breitenbach Barroso Coelho et al. 2018; Gaofu et al. 2008; Lusvarghi and Bewley 2016; Pinto et al. 2019; Vanderlei et al. 2010) . Coronavirus is the main causative of COVID-19 the acute respiratory syndrome which originates from Wuhan China and caused an outbreak worldwide leaving million dead and infected (Hu et al. 2021 ). Many vaccines have been manufactured and approved worldwide to curb the disease's rapid mortality and morbidity rates. However, due to the accelerated frequencies of the virus spike protein mutations, the efficacy of these vaccines becomes questionable (Baraniuk 2021; Hayawi et al. 2021; Khan et al. 2021) . However, the fact that the various N-linked glycosylation points of coronavirus-2 protein, which play a major role in the viral virulence are largely conserved, opened widely the window for the conceivable use of carbohydrate-binding agents such as lectins for targeting this virus' glycans, and thereby interfering with its initial binding stage to the host cell surface receptors (Ahmed et al. 2022; Barre et al. 2021; Martinez et al. 2021) . In this review, we aim to discuss, in-depth, the potential applications of plant lectins to combat coronavirus diseases as well as their uses as, antiulcer, antiinflammatory, and antinociceptive agents.

The coronavirus comprises four main structural proteins: spike (S), membrane (M), envelope (E), and nucleocapsid (N) proteins. Spike protein (S) is the main structural protein antigen of SARS-CoV-2, which mediates the fusion of the coronaviruses into susceptible human cells through their interactions with the angiotensin-converting enzyme-2 (ACE-2) receptor (Hsieh et al. 2005 ).

However, unlike other SARS-CoV-2 viral proteins, it is accountable for triggering the host immune response, and the antibodies directed towards the S protein can bring about protective immunity against subsequent infections (Boechat et al. 2021 ). The S protein (180-200 kDa) is a homotrimer of two subunits S1 and S2, linked through a membrane-embedded serine 2 protease. S1 contains the receptor-binding site, while S 2 is devoted to the viral fusion of the host cells.

Structural investigations of the S protein by cryo-EM and mass spectrometry revealed that S protein to be extensively glycosylated with as many as 66 N-glycosylation points (22 per monomer), covering the surface of the protein and assisting partly in mediating the virulence of the pathogen, and at the same time shielding the susceptible viral receptor-binding domain (RBD) against the neutralizing human antibodies (Huang et al. 2021; Shajahan et al. 2021) Figure 1 . Of distinct interest, the glycosylation points as N165 and N234 (mannose-rich glycans) are located at the vicinity of the ACE-2 RBD, they have been revealed by all-atom molecular dynamic simulation (MD) to play a role in the ACE2-S protein interaction. Point mutation of N165A and N234A, which resulted in glycosylation depletion at these sites, reduced effectively the receptorglycan interaction, though didn't utterly nullify it. These results underlined the possible importance of these glycosylation points in suitably orienting the conformation of the receptorbinding domain of the virus ( (Zhao et al. 2021 ) and references cited therein). Through using different expression vectors, several workers have reported recurrent mutations at N-glycans (Shajahan et al. 2021) . Nonetheless, 19 glycosylation sites were proven to remain conserved. O-glycosylation has also been shown to occur, however, to a limited extent (Hayawi et al. 2021 ).

Expression of SARS-CoV-2 genome on human cell line HEK-293 offered varied complex glycosylation patterns, habitually of the mannose-rich type glycan.

In the search for appropriate and effective treatment for SARS-CoV-2 infection, drug repurposing has already been underway. Inclusion of the antimalarial drugs such as chloroquine and hydroxychloroquine in COVID-19 treatment protocol has made a considerable debate before they were withdrawn after proven futile (Altulea et al. 2021; Ferner and Aronson 2020) . Anticancer drugs have also been proposed to be promising against viral replication however, higher toxicity described with the administration of such drugs is criticized (El Bairi et al. 2020) . As stated the heavily glycosylated SARS-CoV-2 S protein surface made it a tempting choice for glycan-binding proteins (lectins) especially those of plausible interaction with these glycans. It is largely believed that they will likely hinder the attachment of the virus to the host cells by inducing a conformational change that would favour uncovering the epitope recognition site of the virus, thereby neutralizing the virulence effect by the triggered immune response (Figure 2 ). At the vicinity to the RBD of the spike protein, the presence of two glycosylation sequences N165 and N234 which mainly comprised of Man3 GlcNAc core could represent an excellent target for plant lectins with complex-type biantennary oligo-mannosyl saccharides. Almost four decades ago, Greig and Bouillant had revealed extensive binding of Concanavalin A (ConA); the lectin from the Canavalia ensiformis to encephalomyelitis virus, a Coronavirus. Removal of the viral surface glycans by snake venom phospholipase A abolished ConA-virus interactions, emphasizing the importance of the sugar chains in the viral attachment process (Greig and Bouillant 1977) .

On the other hand, Urtica dioica agglutinin (UDA); the lectin purified from the Nettle (Urtica dioica L.), is a small 8.7 kDa protein with a specificity directed towards N, N′, N″-tri-acetylchitotriose (polymer of acetylated acetylglucosamines). This peptide lectin besides its sugar-binding site also had a hydrophobic binding region adjacent to its carbohydrate-binding site (Shibuya et al. 1986 ).

It inhibits the SARS-CoV replication cycle by interfering with the viral attachment to the host cell probably through binding to spike protein N-acetylglucosamine (GlcNAc) units (Kumaki et al. 2011; Saul et al. 2000) . Investigating the effect of the mannose-binding lectin griffithsin against MERS-CoV infection revealed that griffithsin even though displaying no apparent cytotoxicity, offered a strong inhibitory effect on MERS-CoV infection through binding to the mannose-rich viral spike protein (Millet et al. 2016) . The same lectin, in several other publications, was demonstrated to exhibit a broad antiviral spectrum. HIV-1 N-linked glycosylation sites on the gp-120 were affirmed to be recognized by griffithsin, the binding of lectin would lead to a change in the structure of gp-120, which would result in exposing the virus CD4 binding site (Alexandre et al. 2011; Fischer et al. 2019) . Similar results were obtained for banana (Musa acuminata) lectin which could inhibit HIV-1 by recognizing the mannose-rich gp-120 viral outer layer glycoprotein in a range of picomole quantities, hence interfering with the viral adherence to the human cell (Swanson et al. 2010) . Testing a collection of 33 plant lectins with different specificities indicated that the strongest antiviral activity was principally confined among lectins with mannose-binding specificity (Keyaerts et al. 2007 ). Using Vero B4 cells, the Wheat Germ Agglutinin (WGA) purified from Triticum vulgaris exhibited antiviral activity not only against the initially emerged SARS-CoV-2, but also its recent major two variants Alpha and Beta were found effective with IC 50 of <10 ng/mL at both the pre-incubation period with the virus or during the viral infection.

Interestingly, this lectin had a narrow specificity that was merely directed towards coronaviruses as it had no effect with other non-coronaviruses causing respiratory tract distresses (Auth et al. * Genetically engineered lectin with preserved antiviral activity however with a reduced mitogenic capacity (Swanson et al. 2010) 

Apart from the booming development that has arisen in recent years accompanied by studious progress in pharmaceutical biotechnology and the development of drugs manufacturing, the need for effective and powerful analgesics is in continuous demand (Khan et al. 2020) . Moreover, the lectin was successful in inhibiting neutrophils migration. This lectin had also shown a 72% diminution in the mice's abdominal writhing when the pain is induced by acetic acid and therefore is considered to possess both anti-inflammatory and antinociceptive characteristics (Leite et al. 2012) . Several algal lectins were also characterized by their analgesic and sometimes anti-inflammatory properties such as Caulerpa cupressoides lectin, this protein could decrease the effect of the writhing induced by acetic acid to up to 86%, however, it was not being able to produce significant antinociceptive effects in the hot plate experiment, indicating the sole involvement of the peripheral rather than central acting mechanism (Vanderlei et al. 2010 ). The antinociceptive properties of lectins have been, in many instances, attributed to the probable inhibition of inflammation producing molecules such as bradykinin, prostaglandins, substance P, and some cytokines, such as IL-1β and TNFα which will lead to activation of chemosensitive nociceptors and hence induction of pain (Vanderlei et al. 2010) . Similar observations were also noticed with the marine red algae Pterocladiella capillacea lectin (Silva et al 2010) . The writhing effect induced by acetic acid was significantly reduced when Amansia multifida Lamouroux lectin was administered along with the glycoprotein avidin at 1mg/kg. The authors ascribed the occurred attenuation in the antinociceptive property to the blockage of the lectin's active site by the inhibitory glycoprotein avidin, therefore concluding the involvement of the lectin sugar-binding site in the obtained analgesic effect (Neves et al. 2007 ). To examine the involvement of the opioid system, the analgesic morphine was used, and to block the thermal stimuli, the morphine antagonist naloxone was administered which resulted in complete reserve of the morphine effect indicating the involvement of the opioid receptor in the pain sensation (Neves et al. 2007 ). The red seaweed Bryothamnion triquetrum lectin was found potent at 10 mg/kg in inhibiting paw oedema induced by both carrageenan and dextran. This anti-inflammatory response was linked by inhibiting the neutrophil migration towards the peritoneal fluid (Fontenelle et al. 2018) . A mannose-specific lectin isolated from the seeds of Andira anthelmia expressed a potent antinociceptive property; it inhibited mice writhings caused by parenteral administration of 0.7% acetic acid by about 68% in as low as 10 µg/kg dose, this effect was neutralized in presence of mannose (Nascimento et al. 2016) . To wrap up, many legume lectins discussed in this review were proven to attenuate the inflammatory response including the lectin isolated from Canavalia boliviana seeds in a reaction mediated by the lectin binding site (Figueiredo et al. 2009 ), however, astonishingly, Canavalia virosa seed lectin which was purified by affinity interaction to Sephadex ® -G50 (polymer of glucose units), was rather found to trigger inflammation when injected subcutaneously into mice paw. The noticed paw oedema produced by lectin administration was dramatically reverted when the lectin was initially incubated with its haptenic sugar glucose, indicating the role of sugar affinity site in the observed inflammatory reaction (Osterne et al. 2017) . Another interesting work on Lonchocarpus araripensis seed lectin published by a Brazilian group, where the N-acetyl-D-glucosamine (GlcNAc) specific lectin demonstrated antinociceptive effect through a unique mechanism involving the nitric oxide pathway, additionally the same protein was also characterized by GlcNAc specific antiinflammatory property, inhibiting the neutrophil migration to the intraperitoneal cavity of the experimental animal. The administration of the anti-steroidal drug niflumic acid, instead of initiating an analgesic effect, aggravates the pain sensation. Since this drug is used as a calciumactivated chloride channel blocker, the involvement of the calcium channel, in this case, as a firstever report, is apparent (Assreuy et al. 2020; De Freitas Pires et al. 2019) . Similarly, the pretreatment of the animal with Cannabinoid receptor 1 (CB-1) and receptor 2 (CB-2) antagonist AM251 and AM630 respectively overturned the antinociceptive effect of Lonchocarpus araripensis seed lectin, emphasizing the contribution of endocannabinoid receptor (Amorim et al. 2021) . inhibiting the cytokines and chemokines production related to white blood cells migration (Alencar et al. 1999; Jandú et al. 2017 ). 

An ulcer is defined as a pain that originates from damage that occurs in the internal coating of the stomach or small intestine, when the problem is associated with stomach mucosal lesions it is termed gastric ulcer which is caused by acid secretion or pepsin (Sverdén et al. 2019) . In Nigeria, natives routinely use Carica papaya seeds flour for the treatment of peptic ulcers, whereas in Sudan Acacia Senegal and Aerva javanica are known for their healing effect of peptic ulcers (Karar and Kuhnert 2017) . In the experimental models, many gastric-lesion inducers have been employed such as ethanol, Indomethacin and Aspirin. A homotetrameric galactose-binding lectin purified from Artocarpus incise conventionally named frutalin was successful in providing considerable protection against both ethanol and indomethacin gastric injury in mice, however in a doseunrelated manner where a lectin at a concentration as low as 500 μg/kg was able to produce potent protection. Yet,the pre-treatment with the α2-receptor antagonist Yohimbine didn't revert or attenuate frutalin protection against ethanol lesions which indicated no involvement of α2receptor in the caused action of the lectin. Simultaneously administration of glibenclamide, a K +ATP channel inhibitor, resulted in a partial however significant reduction of the action frutalin, demonstrating the influence of K +ATP channel in shielding the stomach lining against the external mucosal attacker (De Vasconcellos Abdon et al. 2012) . A rabbit erythrocytes specific seeds lectin purified from the Brazilian plant Mucuna pruriens (L.) DC (MpLec), was also examined for its protective effect against ethanol-induced gastro-damage in mice. The lectin was able to provide a significant reduction in stomach lesions in comparison to control, however, pre-treatment with Yohimbine abolished the MpLec protective effect, which emphasize the role of α2 adrenoceptors in the attained defensive mechanism (Pinto et al. 2019 ). Another interesting highly stable GlcNAc specific seeds lectin, which agglutinates only the human A-blood group was capable of reducing the ethanol damage by up to 63%. The lectin was administered at three different doses 10, 100 and 1000 µg/kg, even though all of them lead to a significant outcome, major protection was offered by the 1000 µg/kg dose.

The information on the great potency of plant lectins on microorganisms such as bacteria, viruses, parasites, and fungi as well as their reported action as antioxidants, antinociceptive, antitumors and antiulcer have been accumulating. And due to their peculiar sugar recognition site that compliments in minute details with the acknowledged tumor cellular glycan changes, clinical trials on their possible application as a drug shuttle for cancer treatment are underway (Wijetunge et al. 2018; Wode et al. 2020 ). Some of their unfavourable characteristics such as large molecular weight which will likely induce immunogenicity and toxicity may limit the widespread use of these interesting proteins in drug applications. Therefore the suggestion of the implementation of small molecular weight lectins is promising and may pave the way to overcome the immunogenic hurdle. While there is still much to be investigated and disclosed about the in-vivo A) The homotrimer structure of the S-protein showing the N-glycans (Presented in red). B) The monomer of the S-protein A chain is highlighted in blue, of the 22 N-glycans, 11 were presented in 3 different types (hybrid, complex, and high mannose glycans), O-glycans are not shown. The 3D structures were visualized and edited using UCSF-Chimera 1.8v software. 

Antinociceptive and Anti-inflammatory Activities of the Lectin from Marine Red Alga Solieria filiformis

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An anti-inflammatory lectin from Luetzelburgia auriculata seeds inhibits adhesion and rolling of leukocytes and modulates histamine and PGE2 action in acute inflammation models

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Anti-inflammatory effect of glucose-mannose binding lectins isolated from Brazilian beans

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Lectin from Triticum vulgaris (WGA) Inhibits Infection with SARS-CoV-2 and Its Variants of Concern Alpha and Beta

Mannose-specific plant lectins from the Amaryllidaceae family qualify as efficient microbicides for prevention of human immunodeficiency virus infection

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Lectin from green speckled lentil seeds (Lens culinaris) triggered apoptosis in nasopharyngeal carcinoma cell lines

Isolation and Characterization of Myrianthus holstii Lectin, a Potent HIV-1 Inhibitory Protein from the Plant Myrianthus holstii

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Receptors involved in carbohydrate binding modulate intestinal epithelialneutrophil interactions

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Lectin purified from Lonchocarpus campestris seeds inhibits inflammatory nociception

Is the orofacial antinociceptive effect of lectins intrinsically related to their specificity to monosaccharides?

Schinus terebinthifolia leaf lectin (SteLL) has anti-infective action and modulates the response of Staphylococcus aureus-infected macrophages

Gastroprotective potential of frutalin, a d-galactose binding lectin, against ethanol-induced gastric lesions

Insights into animal and plant lectins with antimicrobial activities

The Anti-Inflammatory Effect of Andira Anthelmia Lectin In Rats Involves Inhibition of The Prostanoid Pathway

Repurposing anticancer drugs for the management of COVID-19

Chloroquine and hydroxychloroquine in covid-19

Antinociceptive activity and toxicology of the lectin from Canavalia boliviana seeds in mice

Griffithsin Retains Anti-HIV-1 Potency with Changes in gp120 Glycosylation and Complements Broadly Neutralizing Antibodies PGT121 and PGT126

Lectin obtained from the red seaweed Bryothamnion triquetrum: Secondary structure and anti-inflammatory activity in mice

Lectin Affinity Chromatography

In vitro assessment of plant lectins with anti-pinwood nematode activity

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On the role of cell surface carbohydrates and their binding proteins (lectins) in tumor metastasis

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Antinociceptive activity of lectins from Diocleinae seeds on acetic acid-induced writhing test in mice

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Plant lectins are potent inhibitors of coronaviruses by interfering with two targets in the viral replication cycle

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Lectin isolated from Brazilian seeds of velvet bean (Mucuna pruriens (L) DC.) presents analgesic, anti-inflammatory and antihemolytic action

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The Use of Lectins as Tools to Combat SARS-CoV-2

Middle East respiratory syndrome coronavirus infection is inhibited by griffithsin

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Purification and characterization of Dolichos lablab lectin

A Comparative Analysis of the Chemical Composition, Anti-Inflammatory, and Antinociceptive Effects of the Essential Oils from Three Species of Mentha Cultivated in Romania

Isolation and characterization of a lectin from the seeds of Dioclea grandiflora (Mart.)

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Purification of a thermostable antinociceptive lectin isolated from Andira anthelmia

Antinociceptive properties in mice of a lectin isolated from the marine alga Amansia multifida Lamouroux Braz

Lectin extracted from Canavalia grandiflora seeds presents potential antiinflammatory and analgesic effects

Anti-nociceptive and anti-inflammatory activities of Tetracarpidium conophorum seed lectin

Purification and physicochemical characterization of a cotyledonary lectin from Luetzelburgia auriculata

Isolation and partial characterization of a lectin from Cratylia floribunda mart. seeds

Purification and characterization of a new antiviral protein from the leaves of Pandanus amaryllifolius (Pandanaceae)

Structural characterization of a lectin from Canavalia virosa seeds with inflammatory and cytotoxic activities

Clematis montana lectin, a novel mannose-binding lectin from traditional Chinese medicine with antiviral and apoptosis-inducing activities

Antiulcer and Antioxidant Activity of a Lectin from Mucuna pruriens Seeds on Ethanol-induced Gastropathy: Involvement of Alpha-2 Adrenoceptors and Prostaglandins

Traditional knowledge on medicinal plants used for the treatment of skin diseases in Bidar district

Antinociceptive activity of Schinus terebinthifolia leaf lectin (SteLL) in sarcoma 180-bearing mice

isolation and partial characterization of a lectin from seeds of Dioclea violacea

Plant Lectins with Insecticidal and Insectistatic Activities, Insecticides Agriculture and Toxicology

Anti-asthmatic potential of a D-galactose-binding lectin from Synadenium carinatum latex

Dual-function protein in plant defence: seed lectin from Dolichos biflorus (horse gram) exhibits lipoxygenase activity

Purification and biophysical characterization of a mannose/N-acetyl-dglucosamine-specific lectin from Machaerium acutifolium and its effect on inhibition of orofacial pain via TRPV1 receptor

BEANS. In: Caballero B (ed) Encyclopedia of Food Sciences and Nutrition

Crystal structure of Urtica dioica agglutinin, a superantigen presented by MHC molecules of class I and class II

Glycosylation of SARS-CoV-2: structural and functional insights

Purification and Characterization of a Lectin from Phaseolus vulgaris cv. (Anasazi Beans)

Microbial lectins and their glycoprotein receptors

History of lectins: from hemagglutinins to biological recognition molecules

Evidence that Maackia amurensis seed lectin (MASL) exerts pleiotropic actions on oral squamous cells to inhibit SARS-CoV-2 infection and COVID-19 disease progression

Carbohydrate binding properties of the stinging nettle (Urtica dioica) rhizome lectin

Purification and primary structure of a mannose/glucose-binding lectin from Parkia biglobosa Jacq. seeds with antinociceptive and anti-inflammatory properties

Antinociceptive and anti-inflammatory activities of lectin from marine red alga Pterocladiella capillacea

The role and place of medicinal plants in the strategies for disease prevention

Characterization of a new lectin of soybean vegetative tissues

Peptic ulcer disease

A Lectin Isolated from Bananas Is a Potent Inhibitor of HIV Replication*

Medicinal Plants of the Family Lamiaceae in Pain Therapy: A Review

Isolation, characterization and molecular cloning of the bark lectins from Maackia amurensis

Antinociceptive and anti-inflammatory activities of lectin from the marine green alga Caulerpa cupressoides

Purification and partial characterization of a lectin from the seeds of Dioclea guianensis

Lentil lectin derived from Lens culinaris exhibit broad antiviral activities against SARS-CoV-2 variants

Lectin-Conjugated Liposomes as Biocompatible, Bioadhesive Drug Carriers for the Management of Oral Ulcerative Lesions

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The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.Emadeldin H. E. Konozy (The corresponding author)