Nanomedicine for Human and Veterinary Helminthiasis: Anthelmintic Delivery, Resistance, and Host–Parasite Interactions

Helminth infections are among the most prevalent neglected tropical diseases, affecting both humans and animals worldwide and posing a major public health and veterinary challenge. These infections are caused by parasitic worms (helminths), multicellular eukaryotic organisms that depend on their hosts for survival and nutrition, often resulting in chronic disease and significant health impairment.[1] Helminth infections are particularly common in tropical and subtropical regions and in low- and middle-income countries, where inadequate sanitation, contaminated food and water, and limited access to healthcare facilitate parasite transmission.[2] The major intestinal helminths include Ascaris spp., amphistomes, and strongyles, among others. Infection commonly leads to nausea, vomiting, diarrhea, abdominal pain, malnutrition, impaired physical growth, and reduced quality of life.[3] In addition to these clinical manifestations, several helminths are known to modulate the host immune system by expanding regulatory T cells (Tregs), thereby suppressing inflammatory responses.[4] Greenwood first reported an association between widespread helminth exposure and a lower prevalence of rheumatoid arthritis among African populations, suggesting that helminths may influence the development of inflammatory and autoimmune diseases.[5] Although these immunomodulatory properties have attracted considerable research interest, helminth infections remain a major global health concern. According to the U.S. Centers for Disease Control and Prevention (CDC), diarrheal diseases account for approximately one in nine child deaths worldwide, causing nearly 2,195 deaths each day—more than malaria, AIDS, and measles combined.[6] Soil-transmitted helminths alone infect more than 1.5 billion people globally, with preschool- and school-aged children and women of reproductive age bearing the greatest disease burden. [7] Approximately 267 million preschool-aged children and 568 million school-aged children live in regions where transmission remains intense, necessitating regular preventive interventions and effective treatment. The highest prevalence occurs in Sub-Saharan Africa, the Americas, China, and East Asia.[7] Beyond human health, helminth infections also have profound economic consequences for livestock production. Fasciolosis affects approximately 250 million sheep and 350 million cattle worldwide, placing more than 27 million people at risk.[8,9] Gastrointestinal helminths reduce animal growth, fertility, feed efficiency, milk production, and overall productivity, particularly under intensive farming conditions or in contaminated environments.[10] These losses are expected to become increasingly important as global meat consumption continues to rise, with per capita consumption projected to increase substantially over the coming decades.[11] The severity of infection depends on several factors, including parasite species, infective dose, host susceptibility, parasite reproductive capacity, and host–parasite interactions.[12] Consequently, helminth infections may cause weight loss, reduced fecundity, impaired growth, production losses, or go unnoticed clinically because of their chronic nature. Most helminths possess complex life cycles involving egg, larval, and adult stages. Depending on the parasite species, transmission occurs through ingestion of infective eggs or larvae in contaminated food, water, or fodder; consumption of infected intermediate hosts; skin penetration; or, less commonly, vertical transmission or arthropod vectors. Adult worms typically reside within the definitive host, whereas larval stages may develop in the environment or within intermediate hosts before completing their life cycle (Fig. 1). Control of helminth infections relies primarily on a limited number of anthelmintic drug classes, including benzimidazoles (albendazole and mebendazole), macrocyclic lactones (ivermectin and other avermectins), and praziquantel. Although these drugs have substantially reduced the burden of helminthiasis, their long-term effectiveness is increasingly threatened

Fig. 1:Taxonomic classification of helminths. There are three main groups of helminths, i.e., Trematodes (flukes), Cestodes (tapeworms), and Nematodes (roundworms). They are classified by their overall external shape and the host organ they inhabit. Helminths develop through egg, larval, and adult stages. The outer covering of the helminths is the cuticle or tegument.

Table 1: Current anthelminthic drugs and their mechanism

Name of the drug	Structure	Mechanism	Side effects
Fenbendazole		It binds to tubulin, disrupts the tubulin-microtubule equilibrium, and inhibits polymerization in helminths. It also induces apoptotic cell death of porcine trophectoderm and uterine cells following membrane disruption. It might lead to implantation failure in pigs during early pregnancy.[16,17]	Diarrhea, Loss of appetite, Lethargy
Piperazine		It is a selective GABA agonist that activates and gates GABA receptors on nematode muscle. It causes hyperpolarization of muscle cells and nerve membranes at the neuromuscular junction leading to parasite immobilization by flaccid paralysis and consequent removal by interfering with the energy metabolism of the nematode.[18]	Blurring vision, clumpiness, joint pain, skin rash, crawling or tingling feeling of the skin, skin rash, or itching.
Mebendazole		It inhibits the production of microtubules via binding to the colchicine binding site of β-tubulin and thereby blocking the polymerization of tubulin dimers in the intestinal cells of parasites. Hence, glucose uptake gets interrupted, resulting in immobilization and hindrance of egg production and death of helminth.[19]	Diarrhea, Stomach pain, swelling, nausea, vomiting, loss of appetite.
Albendazole		It binds to the tubulin dimer interface overlapping the  binding site of β-tubulin, thereby inhibiting  and disrupting microtubule networks.[20]	Headache, neck stiffness, abnormal liver functions, temporary hair loss.
Morantel[21]		It acts as an agonist of the nAChR subtype comprising ACR-26/ACR-27 subunits from H. contortus or Parascarisequorum expressed in X. laevis oocytes. In oocyte expression studies, morantel was seen to cause a non-competitive voltage-sensitive open channel block of the newly characterized A. suum ACR-16 receptor.[22-24]	Dyspnea, Tachypnea, Diarrhea, Tremor, Convulsions, Ataxia.
Pyrantel		It activates L-subtype nAChR in A. suum.It also causes an open channel block.[24,25]	Nausea, diarrhea, stomach cramps, trouble sleeping, loss of appetite.
Levamisole		Levamisole opens nematode AChRs that are ligand-gated, non-selective cationchannels.[25]	Agranulocytosis, skin rash, and febrile illness.
Ivermectin		Ivermectin increases the activity of γ-aminobutyric acid (GABA) receptors or glutamate-gated chloride ion channels (Glu-Cl), which blockades the signal between neurons and muscles.[21,26]	Dizziness, loss of appetite, nausea, diarrhea, weakness.

by poor bioavailability, limited drug targets, repeated treatment requirements, adverse effects, and, most importantly, the rapid emergence of anthelmintic resistance.[13,14] For example, monepantel, introduced for livestock treatment in 2009, experienced documented resistance in sheep and goats in New Zealand within only three years of its commercial release.[15] Such rapid development of resistance underscores the urgent need for innovative therapeutic approaches. The currently available anthelmintic drugs, their mechanisms of action, therapeutic applications, and major limitations are summarized in Table 1. Nanomedicine has emerged as a promising strategy to overcome many limitations associated with conventional anthelmintic therapy. Nanoparticle-based drug delivery systems can improve drug solubility, stability, and bioavailability, enable

Fig. 2:Life cycle of nematodes (A) direct life cycle (in monogenous nematodes), (B) indirect life cycle (in heterogenous nematodes), (C) indirect life cycle in the self-host (in auto-heterogenous nematodes). There are four stages in a nematode's life cycle: egg, four larval or juvenile stages, and adult. At each juvenile stage, the cuticle sheds, and the nematode increases in size. Nematode's growth depends on the conditions inside the host.

controlled release and targeted delivery, and potentially reduce toxicity and delay the emergence of drug resistance. Furthermore, several nanomaterials possess intrinsic antiparasitic activity and have demonstrated enhanced therapeutic efficacy when used alone or in combination with conventional anthelmintic drugs or herbal formulations. Consequently, nanotechnology represents a promising platform for developing next-generation therapeutics against parasitic worm infections. This review summarizes recent advances in the application of nanomedicine for the management of parasitic worm infections. We discuss different nanoparticle-based delivery systems, their mechanisms of action, therapeutic applications in human and veterinary helminthiasis, their advantages over conventional formulations, current limitations, and prospects for translating these technologies into clinical and veterinary practice.

2.1 Helminthic zoonoses

Helminthic zoonoses are parasitic worm infections that are naturally transmitted between animals and humans, representing a major public health and veterinary concern worldwide.[27] Zoonotic diseases account for more than 60% of all human infectious diseases, with helminth parasites contributing substantially to this global burden. Transmission to humans occurs through multiple routes, most commonly via the accidental ingestion of infective eggs or larvae present in contaminated food, water, or soil. For example, Ascaris spp. and Trichuris spp. are primarily transmitted through the fecal–oral route, whereas infective larvae of hookworms and Schistosoma spp. can actively penetrate intact skin. Several helminths possess complex life cycles involving intermediate and definitive hosts, with humans serving as either intermediate or definitive hosts depending on the parasite species (Fig. 2). In contrast, some nematodes, such as roundworms and whipworms, readily infect both humans and animals. Environmental, socioeconomic, and behavioral factors, including sanitation, hygiene, climatic conditions, food preparation practices, dietary habits, and exposure to infected animals or vectors, strongly influence the prevalence and transmission of helminthic zoonoses. Soil-transmitted helminths (STHs) are the most widespread human helminth infections, collectively affecting approximately one-quarter of the global population.[28] The major STHs include Ascaris lumbricoides, Strongyloidesstercoralis, Trichuris trichiura, and the hookworms Ancylostoma duodenale and Necator americanus.[29] In addition to STHs, several foodborne and waterborne helminth infections—including echinococcosis, fascioliasis, taeniasis, cysticercosis, diphyllobothriasis, capillariasis, and ascariasis—remain important but often under-recognized causes of human disease. Their incidence has increased in recent years due to international travel, globalization of the food supply, changing dietary preferences, and the growing consumption of raw or undercooked foods, underscoring the continued need for effective surveillance, prevention, and control strategies.[30]

2.2 Helminth infections and host immune regulation

Helminths have evolved sophisticated mechanisms to evade and manipulate host immune responses, enabling long-term survival within their hosts while minimizing excessive tissue damage. Through millions of years of coevolution, these parasites have established a finely balanced relationship with their hosts by modulating both innate and adaptive immunity, thereby promoting chronic infection and limiting harmful inflammatory responses.[31] A key strategy employed by helminths is the secretion of excretory/secretory (ES) products, including extracellular vesicles (EVs). These vesicles transport a diverse range of bioactive molecules, including proteins, lipids, messenger RNAs, and microRNAs, that regulate host immune signaling pathways and suppress protective immune responses.[32] Growing evidence suggests that EV-mediated communication plays a central role in host–parasite interactions and offers promising targets for the development of novel therapeutics and vaccines.[33] Nevertheless, further studies are required to elucidate the specific functions of individual EV cargo molecules and their interactions with different immune cell populations. Among helminths, Schistosoma spp. provide one of the best-characterized examples of parasite-mediated immune regulation. In addition to causing chronic hepatic, intestinal, urinary, and renal diseases, schistosomes can modulate host immunity in ways that reduce the severity of certain inflammatory and autoimmune disorders. Experimental studies have demonstrated that Schistosoma mansoni infection alleviates allergic airway inflammation, suppresses autoimmune diabetes in mice, and delays disease progression in experimental models of multiple sclerosis.[5] The eggs of S. mansoni not only contribute to disease pathology but also secrete immunomodulatory molecules, including the glycoprotein omega-1, which plays a pivotal role in directing host immune responses toward a regulatory phenotype.[34] Such immune modulation limits excessive inflammation, thereby enhancing host survival while facilitating long-term parasite persistence. Helminth infections are also the leading cause of eosinophilia among travellers, accounting for approximately 30–60% of reported cases, with schistosomiasis representing the predominant diagnosis.[35] Although substantial progress has been made in schistosomiasis control through preventive chemotherapy and public health interventions, implementation remains uneven across endemic regions due to limited healthcare infrastructure and incomplete treatment coverage. Antibody-based assays remain useful for diagnosing travellers and individuals living in low-endemic or post-elimination settings. In contrast, antigen detection and parasite DNA-based assays are emerging as promising tools for field surveillance and disease monitoring.[36] Human hookworms, including Ancylostoma duodenale and Necator americanus, are major causes of iron-deficiency anemia and protein malnutrition, particularly among children. Interestingly, increasing evidence indicates that hookworm infections also possess immunomodulatory properties. Infection with N. americanus has been associated with reduced dust mite sensitization and decreased asthma-related wheezing in children. [37] Likewise, excretory/secretory products from the canine hookworm Ancylostoma caninum suppress allergic airway inflammation in murine models by inducing Th2-associated cytokines, including interleukin-5 (IL-5) and interleukin-13 (IL-13).[38] Furthermore, experimental infection with N. americanus has been shown to reduce intestinal expression of the pro-inflammatory cytokine interferon-γ in patients with autoimmune disorders such as celiac disease.[5] Collectively, these findings demonstrate that, despite their pathogenicity, helminths possess remarkable immunomodulatory properties that may inspire the development of novel immunotherapeutic strategies.

2.3 Current anthelmintic therapy and drug resistance

Despite the enormous global burden of helminth infections, therapeutic options remain limited to only a few classes of anthelmintic drugs. Benzimidazoles (albendazole, mebendazole, and triclabendazole), praziquantel, and macrocyclic lactones such as ivermectin constitute the cornerstone of helminth control in both human and veterinary medicine. The World Health Organization (WHO) recommends albendazole and praziquantel for the treatment of several intestinal helminth infections and for preventive chemotherapy programs, particularly among school-aged children, who represent the principal target population for mass drug administration.[39,40] Each class of anthelmintic drugs exhibits a distinct mechanism of action. Benzimidazoles selectively bind to β-tubulin, disrupting microtubule polymerization, impairing glucose uptake, and ultimately causing energy depletion and the parasite's death.[41] Praziquantel increases the permeability of the parasite's tegument to calcium ions, resulting in sustained muscle contraction, paralysis, and parasite elimination.[42] In contrast, ivermectin targets glutamate-gated chloride ion channels located in the nerve and muscle cells of nematodes, producing membrane hyperpolarization, paralysis, and eventual death of the parasite.[43] Although these drugs have substantially reduced the global burden of helminthiasis, they have several limitations, including limited activity against certain parasite species or developmental stages, poor bioavailability of some compounds (e.g., albendazole), the need for repeated treatment, and the increasing emergence of drug resistance. Treatment failures have now been reported for several human and veterinary helminths, particularly schistosomes and gastrointestinal nematodes, raising concerns regarding the long-term sustainability of existing control programs.[44] Anthelmintic resistance has become especially widespread in livestock, with prevalence exceeding 50% in many regions worldwide, resulting in significant economic losses, reduced animal productivity, and compromised animal welfare.[12] An overview of the major helminth infections affecting humans and animals, together with their first-line therapies and reported resistance, is presented in Table 2. Drug resistance arises through multiple molecular and genetic mechanisms. These include alterations in drug targets that reduce binding affinity; modifications in drug metabolism that enhance detoxification or decrease drug activation; alterations in drug transport, including increased efflux that reduces intracellular drug accumulation; and amplification or overexpression of target genes that reduce drug susceptibility.[12] Additional mechanisms, such as mutations in receptor genes, altered gene expression, and enhanced xenobiotic detoxification pathways, further contribute to the evolution of resistant parasite populations. Recognizing the growing threat of anthelmintic resistance, a Biotechnology and Biological

Table 2: Statistics of parasitic helminth infections in humans and animals and their clinical treatment concerning resistance to anthelmintic drugs

*Note: Helminths get transferred to animals and humans through the soil, food, and water. Generally, flu-like symptoms are common, including high temperatures and muscle aches. Symptoms may also include skin rashes, abdominal pain, diarrhea, and cough. Also, inflammation occurs in areas of the body where eggs have lodged, i.e., intestine, eggs, liver, lungs, and even brain. This table includes the activity of antiparasitic drugs, which outlines indications, dosage, adverse effects, and resistance towards these drugs.

NR = Not reported.

Table 3: Potential herbal medicines with anthelmintic properties

*Note: This table discusses the herbal plants used against different helminth infections, along with their active components, structure, and mode of action.

Sciences Research Council (BBSRC)-sponsored workshop identified four major research priorities: the development of improved diagnostic tools for resistance detection, a better understanding of drug mechanisms and resistance pathways, deeper insights into parasite biology, and the establishment of predictive decision-support systems for sustainable parasite control. These priorities emphasize the urgent need for integrated management strategies that extend beyond reliance on conventional chemotherapy. Among livestock parasites, gastrointestinal nematodes (GINs) remain one of the greatest constraints on profitable animal production worldwide. The widespread emergence of resistant parasite populations, together with the limited number of available drug classes and the high cost of developing new anthelmintics, has stimulated interest in alternative control strategies. Plant-derived anthelmintics, widely used in ethnoveterinary medicine, have demonstrated promising antiparasitic activity in numerous experimental studies (Table 3). However, their efficacy is often influenced by variability in phytochemical composition, dosage, host nutrition, and potential anti-nutritional effects. Consequently, standardized evaluation methods and integrated approaches are essential for maximizing their therapeutic benefits while minimizing adverse effects. These limitations have accelerated interest in nanotechnology-based drug delivery systems, which offer new opportunities to improve the efficacy, bioavailability, and targeted delivery of existing anthelmintic agents.

The rapid emergence of anthelmintic resistance, coupled with the limited number of available drug classes, has accelerated the search for innovative therapeutic strategies. Nanotechnology has emerged as a promising platform for improving the diagnosis, prevention, and treatment of parasitic diseases by enhancing drug delivery and therapeutic efficacy.[180] Nanomaterials, typically ranging from 1 to 100 nm in size, are composed of metallic, non-metallic, or hybrid materials whose physicochemical properties can be tailored through controlled synthesis and surface functionalization.[181] Polymer coatings are commonly employed to improve biocompatibility, prolong circulation time, and enable selective targeting of biological molecules.[182] The size, morphology, and surface characteristics of nanoparticles are influenced by synthesis parameters, including precursor composition, surfactants, reaction temperature, solvent system, and reactant concentrations (Table 4).[183] Conventional anthelmintic drugs primarily target parasite neuromuscular function or essential metabolic pathways, leading to paralysis, starvation, and eventual elimination of the parasite.[184] However, their effectiveness is often compromised by poor aqueous solubility, limited bioavailability, nonspecific tissue distribution, rapid clearance, and the increasing prevalence of drug-resistant parasite populations.[185] Nanotechnology offers an attractive solution to these limitations by improving drug stability, controlled release, cellular uptake, and site-specific delivery while reducing systemic toxicity. Consequently, nanoparticle-based drug delivery systems have gained considerable attention for enhancing the efficacy of existing anthelmintic agents rather than relying solely on the discovery of entirely new drugs.

An additional advantage of nanoformulation is its potential to mitigate transporter-mediated mechanisms of anthelmintic resistance. In resistant parasites, increased expression or activity of drug-efflux transporters can reduce the intracellular accumulation of anthelmintic compounds, thereby decreasing their access to molecular targets. Encapsulation within polymeric or lipid-based nanocarriers, such as PLGA nanoparticles, solid lipid nanoparticles (SLNs), and liposomes, can alter the physicochemical form and cellular trafficking of the drug, reducing its immediate exposure to membrane-associated efflux systems. Following uptake of the nanocarrier via endocytic or other membrane-associated processes, the encapsulated drug can be released intracellularly in a controlled manner, providing sustained drug exposure and potentially increasing the fraction of drug that reaches its intracellular or membrane-associated targets. In addition, modulation of particle size, surface charge, lipid composition, and surface functionalization can influence membrane interaction, cellular uptake, and intracellular trafficking, thereby providing delivery routes that differ from those of the freely dissolved drug. Thus, nano-delivery may partially circumvent rapid efflux by modifying the route, kinetics, and intracellular availability of the anthelmintic rather than directly inhibiting the resistance mechanism itself. This distinction is important because a nanocarrier's ability to overcome efflux-mediated resistance likely depends on the physicochemical properties of the carrier, the anthelmintic compound, the relevant transporter, and the parasite species. Consequently, nanocarriers may complement conventional resistance-management strategies by improving drug exposure at the parasite target while reducing the extent to which transporter-mediated drug extrusion limits therapeutic efficacy. Nanomedicine applies engineered nanomaterials for disease diagnosis, prevention, and therapy.[186-189] Although initially developed for applications such as oncology and infectious diseases, advances in nanomedicine have provided valuable strategies to improve antiparasitic chemotherapy. Compared with conventional formulations, nanocarriers can increase the solubility of hydrophobic drugs, protect active compounds from premature degradation, prolong circulation time, and facilitate targeted delivery to infected tissues. These advantages may reduce dosing frequency, improve therapeutic outcomes, and minimize adverse effects, making nanomedicine an attractive approach for both human and veterinary helminthiasis.[190]

A wide variety of nanocarriers have been investigated for anthelmintic drug delivery, including polymeric nanoparticles, lipid-based nanoparticles, dendrimers, metallic nanoparticles, micelles, and liposomes (Table 4).

Table 4. Anthelmintic activity of various nanoparticles and their mode of action

*Note: In this table, we have discussed multiple nanoparticles tested against different helminths and their mechanisms. The most-reported methods of action of nanoparticles are antimicrobial activity, ROS-induced cytotoxicity, genotoxicity, plant growth promotion, etc. It has been successfully demonstrated that the actions of nanoparticles are governed by their size, shape, dose, and concentration.

NR = Not reported.

Fig. 3:Graphical representation of Anthelmintic activity of various nanoparticles and their mode of action.

Poly(lactic-co-glycolic acid) (PLGA) remains one of the most extensively studied biodegradable polymers because of its excellent biocompatibility and controlled drug-release properties.[229] Encapsulation of praziquantel within PLGA nanoparticles has been shown to improve drug stability and modify release kinetics. In contrast, solid lipid nanoparticles (SLNs) have demonstrated enhanced oral bioavailability of poorly water-soluble drugs.[230] Likewise, liposomal formulations improve drug encapsulation efficiency, prolong systemic circulation, and promote targeted drug delivery, thereby increasing therapeutic efficacy while reducing systemic toxicity.[231] In addition to functioning as drug carriers, several nanoparticles exhibit intrinsic antiparasitic activity. Metallic nanoparticles, polymeric nanomaterials, and certain hybrid nanostructures have been reported to induce reactive oxygen species (ROS)-mediated oxidative stress, disrupt parasite membranes, impair reproductive capacity, and cause genotoxic damage, ultimately leading to parasite death (Fig. 3). When combined with conventional anthelmintic drugs, these nanomaterials often produce synergistic effects, enabling lower therapeutic doses and improving treatment efficacy against resistant parasites. However, the promising activity reported in experimental studies does not necessarily translate into effective in vivo therapy. Metallic nanoparticles may exert strong antiparasitic effects via oxidative stress, membrane disruption, or genotoxicity, but these mechanisms can also cause nonspecific toxicity in host tissues, thereby narrowing the therapeutic window. Moreover, their biological activity is strongly influenced by particle size, surface characteristics, dose, and aggregation state, meaning that effects observed at relatively high concentrations in isolated parasites may not be reproducible following administration in vivo. The studies summarized in Table 4 also predominantly involve experimental models such as Caenorhabditis elegans, Pheretima posthuma, or isolated parasite stages, which limits direct extrapolation to infected mammalian or livestock hosts. Polymeric and lipid-based nanocarriers may provide greater opportunities for controlled drug delivery, but they also face important barriers to in vivo translation. Although PLGA nanoparticles, SLNs, and chitosan-based systems can improve drug solubility, stability, and release characteristics, their in vivo performance may be compromised by protein corona formation, aggregation, uptake by the mononuclear phagocyte system, premature clearance, and inadequate delivery to the parasite site. Thus, the promising activity of the PLGA-, SLN-, and chitosan-based formulations summarized in Table 4 should be interpreted alongside the need for pharmacokinetic, biodistribution, toxicity, and infected-host studies before therapeutic efficacy can be established.

Despite these promising advances, several challenges continue to limit the clinical translation of nanomedicine. Following administration, nanoparticles encounter numerous biological barriers that influence their absorption, biodistribution, metabolism, and excretion (ADME). Interactions with plasma proteins lead to the formation of a protein corona, which alters nanoparticle stability, circulation time, and tissue distribution. Most nanoparticles are ultimately cleared by the mononuclear phagocyte system, particularly in the liver and spleen, thereby limiting their accumulation at target sites.[232,233] Consequently, optimizing nanoparticle physicochemical properties remains a major focus of current research. These ADME challenges are particularly important for veterinary applications involving food-producing animals. Unlike conventional small-molecule anthelmintics, nanoformulations may exhibit altered tissue distribution, prolonged retention, and formulation-dependent drug release, which can influence the depletion of active compounds and nanoparticle-associated components from edible tissues. Consequently, nanoparticles or their associated drug residues may potentially persist in tissues such as muscle, liver, kidney, and fat and, in lactating animals, may also be transferred into milk, raising concerns regarding consumer exposure. However, the extent of such accumulation and persistence is highly dependent on nanoparticle composition, particle size, surface characteristics, route of administration, dose, and the physicochemical properties of the encapsulated drug. Therefore, residue depletion cannot be reliably extrapolated from conventional formulations of the same anthelmintic. Dedicated pharmacokinetic and tissue-residue studies in relevant livestock species are required to characterize absorption, distribution, transformation, and elimination and to determine the persistence of both the active pharmaceutical ingredient and relevant nanoparticle-derived components. Such data are essential for establishing scientifically justified maximum residue limits (MRLs) and appropriate withdrawal periods for meat, milk, and other edible products before nano-anthelmintic formulations can be routinely used in food-producing animals. Regulatory evaluation should therefore consider formulation-specific pharmacokinetic, residue-depletion, toxicological, and consumer-safety data, particularly where repeated administration or prolonged tissue retention is anticipated.

The safety of nanomedicine also requires careful evaluation. Parameters such as biocompatibility, biodegradability, pharmacokinetics, nanotoxicology, large-scale manufacturing, and regulatory compliance must be thoroughly assessed before clinical application.[234] Certain metallic nanoparticles have been reported to accumulate in the liver and other reticuloendothelial tissues, resulting in elevated liver enzyme levels and tissue damage following prolonged exposure.[235] Furthermore, the toxicity of metallic nanoparticles often depends on their composition, particle size, surface chemistry, and solubility, with highly soluble metal nanoparticles generally exhibiting greater cytotoxicity in vitro.[236] Therefore, balancing therapeutic efficacy with long-term safety remains a critical challenge for future nano-anthelmintic development. Overall, nanotechnology provides a versatile platform for overcoming many of the limitations associated with conventional anthelmintic therapy. Nanoformulations can enhance drug solubility, stability, bioavailability, and targeted delivery while minimizing systemic exposure and adverse effects. By increasing drug accumulation at parasite-infected tissues, including the gastrointestinal tract, liver, lungs, and skin, nanoparticle-based formulations have the potential to improve treatment outcomes, reduce the emergence of resistance, and revitalize the therapeutic efficacy of existing anthelmintic drugs.[144] Continued advances in nanocarrier design, safety evaluation, and clinical translation are expected to establish nanomedicine as an important component of future strategies for managing helminth infections.

Efficient drug delivery remains one of the major challenges in the treatment of helminth infections. Conventional anthelmintic drugs often exhibit poor aqueous solubility, limited bioavailability, rapid systemic clearance, and nonspecific tissue distribution, reducing their therapeutic efficacy. Consequently, considerable attention has been directed toward vesicular drug delivery systems, including extracellular vesicles (EVs), liposomes, lipid nanoparticles (LNPs), and solid lipid nanoparticles (SLNs), which offer improved drug stability, controlled release, targeted delivery, and reduced systemic toxicity. Extracellular vesicles are naturally occurring nanosized membrane-bound particles released by both prokaryotic and eukaryotic cells. Helminths also secrete EVs as components of their excretory/secretory products, thereby enabling them to communicate with host cells and modulate immune responses by transferring proteins, lipids, messenger RNAs, and microRNAs.[237,238] Beyond their biological role in host–parasite interactions, EVs have attracted considerable interest as naturally derived drug-delivery vehicles owing to their excellent biocompatibility, low immunogenicity, and inherent ability to transport bioactive molecules across biological barriers. These characteristics make EVs promising candidates for therapeutic delivery, vaccine development, and diagnostic applications in parasitic diseases. Among lipid-based nanocarriers, liposomes remain one of the most extensively investigated drug delivery systems. Liposomes are biodegradable phospholipid bilayer vesicles capable of encapsulating both hydrophilic and hydrophobic therapeutic agents, thereby protecting drugs from enzymatic degradation and improving their pharmacokinetic properties.[239,240] Their structural similarity to biological membranes facilitates cellular uptake and prolongs systemic circulation, while surface modification with antibodies, peptides, or receptor-specific ligands enables targeted drug delivery to diseased tissues. Consequently, liposomal formulations have been widely investigated for improving the efficacy and safety of anticancer, antiviral, antibacterial, and antiparasitic therapies.

In helminth infections, liposomal drug delivery has shown considerable promise for enhancing the therapeutic performance of existing anthelmintic agents. Encapsulation of praziquantel and ivermectin within liposomes has been reported to improve drug delivery, prolong drug release, and increase deworming efficacy compared with conventional formulations.[237] Interestingly, phospholipid-rich liposomes themselves may also exert direct effects on parasites by altering the motility and morphology of Schistosoma mansoni. At the same time, liposomal formulations have been reported to reduce the incidence of monensin resistance.[241] These findings suggest that liposomes can function both as efficient drug carriers and as adjunctive therapeutic platforms. Solid lipid nanoparticles (SLNs) represent another important class of lipid-based nanocarriers. They combine the advantages of polymeric nanoparticles and liposomes while offering improved physical stability, biodegradability, ease of large-scale production, and sustained drug release. Because of their favorable safety profile and high drug-loading capacity, SLNs have emerged as attractive carriers for poorly water-soluble anthelmintic drugs, improving oral bioavailability and therapeutic efficacy. Recent advances have also demonstrated the potential of biomimetic nanoparticle systems for targeted parasite control. Sharma et al. (2021) developed lipid-based nanoparticles using compounds naturally consumed by helminths, achieving an albendazole loading efficiency of 83.3 ± 6.5 mg/g and sustained drug release, with approximately 86% of the encapsulated drug released within 24 h. Uptake of Rhodamine B-loaded nanoparticles by Haemanthus contortus confirmed efficient particle ingestion and gradual dissemination throughout the parasite.[221] Notably, nanoparticle encapsulation enhanced albendazole potency by nearly 50-fold, highlighting the considerable potential of lipid-based nanocarriers to improve drug bioavailability, reduce therapeutic doses, and minimize systemic toxicity.[221]

Beyond nanoparticles, novel bioinspired drug delivery devices are also being explored. Theragrippers are miniature self-actuating devices inspired by the attachment mechanisms of gastrointestinal parasites. Upon exposure to physiological temperature, these microdevices anchor firmly to the intestinal mucosa and provide sustained local drug release. In preclinical studies, theragrippers remained attached to the gastrointestinal tract of rats for at least 24 hours, demonstrating their potential for prolonged intestinal drug delivery and improved therapeutic retention.[242] Collectively, extracellular vesicles and lipid-based nanocarriers represent highly promising platforms for next-generation anthelmintic therapy. Their ability to enhance drug stability, prolong circulation, improve target-site accumulation, and reduce systemic toxicity offers significant advantages over conventional formulations. Continued optimization of these delivery systems, together with comprehensive evaluations of their safety, pharmacokinetics, and large-scale manufacturability, will be essential for their successful translation into clinical and veterinary practice.

Despite the promising anthelmintic potential of nanoparticle-based approaches, several challenges must be addressed before their translation into practical applications in human and veterinary helminthiasis. Most available studies remain at the in vitro or preliminary experimental stage, with relatively limited validation in infected animal models and clinical settings. Differences in nanoparticle size, morphology, surface charge, composition, and preparation methods can substantially influence biological activity, cellular uptake, biodistribution, and toxicity, making comparison between studies difficult. In addition, metallic nanoparticles may induce nonspecific oxidative stress and tissue damage, raising concerns regarding their therapeutic window and long-term safety. Polymeric and lipid-based nanocarriers can improve drug solubility, stability, and controlled release; however, protein-corona formation, aggregation, rapid clearance, and inadequate delivery to parasite-associated sites may limit their effectiveness in vivo. These factors highlight the need for standardized nanoparticle characterization and systematic evaluation of pharmacokinetics, biodistribution, toxicity, and therapeutic efficacy in relevant host–parasite models. Future research should therefore focus on the development of biodegradable and biocompatible nanocarriers capable of delivering anthelmintic agents selectively to parasites while minimizing exposure to host tissues. Targeted and controlled-release systems may improve drug accumulation at the site of infection and potentially reduce the dose and frequency of administration. Nanocarriers could also facilitate combination therapy by co-delivering multiple anthelmintic agents or integrating conventional drugs with bioactive compounds, which may be particularly valuable in managing anthelmintic resistance. For veterinary applications, formulation-specific residue depletion studies will be essential to establish appropriate withdrawal periods and ensure the safety of meat, milk, and other edible products. Greater emphasis should also be placed on scalable manufacturing, batch-to-batch reproducibility, cost-effectiveness, and regulatory requirements. Ultimately, integrating nanotechnology with conventional anthelmintic therapy, resistance-management strategies, and One Health approaches could support the development of safer, more effective, and sustainable interventions against helminth infections.

Nanotechnology has emerged as a promising strategy to address many of the limitations associated with conventional anthelmintic therapy. As highlighted throughout this review, nanoparticle-based drug delivery systems can enhance the solubility, stability, bioavailability, and targeted delivery of antiparasitic agents while enabling sustained drug release and reducing systemic toxicity. These advantages have the potential to improve therapeutic efficacy, lower required drug doses, and extend the clinical usefulness of existing anthelmintic drugs, particularly in the face of increasing drug resistance. The rapid emergence of anthelmintic resistance has become a major challenge in both human and veterinary medicine, threatening disease control programs, livestock productivity, and global food security. Although nanoparticle-based formulations have shown encouraging results in improving drug delivery and overcoming some pharmacokinetic limitations, their ability to prevent or reverse established drug resistance remains incompletely understood. Future studies should therefore focus not only on developing more effective nanoformulations but also on elucidating the molecular mechanisms underlying parasite resistance and on identifying nanoparticle-based strategies to restore drug susceptibility. Successful clinical translation of nanomedicine will require overcoming several scientific, technical, and regulatory challenges. The design of multifunctional nanocarriers capable of achieving efficient drug loading, prolonged circulation, controlled release, selective targeting, and minimal toxicity remains an important research priority. In addition, standardized methods for evaluating nanoparticle pharmacokinetics, biodistribution, biocompatibility, and long-term safety are needed to facilitate comparison among studies and accelerate regulatory approval. The development of reliable diagnostic tools for the early detection and surveillance of anthelmintic resistance will further strengthen integrated parasite management strategies and support the rational use of existing therapeutics. Overall, the evidence summarized in this review demonstrates that nanomedicine represents a promising complementary approach rather than a replacement for conventional anthelmintic therapy. Continued interdisciplinary research integrating parasitology, nanotechnology, materials science, pharmacology, and veterinary medicine will be essential for translating these promising laboratory findings into safe, effective, and economically viable clinical and field applications. Such advances have the potential to improve the management of helminth infections in both humans and animals while contributing to sustainable parasite control and global health.

Helminth infections continue to pose a major challenge to global public health and livestock production, particularly in regions with limited healthcare resources and inadequate sanitation. Although conventional anthelmintic drugs have substantially reduced the burden of these infections, their long-term effectiveness is increasingly threatened by poor bioavailability, limited therapeutic options, and the widespread emergence of drug resistance. Consequently, there is an urgent need for innovative therapeutic strategies to improve treatment efficacy and extend the lifespan of existing drugs. This review highlights the growing potential of nanomedicine as an advanced platform for the management of helminth infections. Nanoparticle-based drug delivery systems, including polymeric nanoparticles, lipid-based nanocarriers, extracellular vesicles, and other biomimetic delivery platforms, offer significant advantages by enhancing drug solubility, stability, controlled release, targeted delivery, and therapeutic efficacy while minimizing systemic toxicity. In addition to improving the performance of existing anthelmintic drugs, these technologies may provide new opportunities to overcome pharmacokinetic limitations and help address the growing challenge of anthelmintic resistance. Despite these promising advances, several barriers remain before nano-anthelmintic formulations can be widely implemented in clinical and veterinary practice. Comprehensive evaluation of long-term safety, pharmacokinetics, large-scale manufacturing, regulatory approval, and cost-effectiveness will be essential for successful clinical translation. Continued interdisciplinary collaboration among parasitologists, nanotechnologists, pharmacologists, veterinarians, and clinicians will accelerate the development of safe and effective nanoformulations. Overall, nanomedicine represents a promising complement to conventional anthelmintic therapy rather than a replacement for it. Continued research and technological innovation are expected to facilitate the development of next-generation therapeutics that improve parasite control, mitigate the impact of drug resistance, and enhance the health and productivity of both humans and animals.

None

Harmanjot Kaur: Conceptualization, Literature review, Analysis, Visualization, Writing – Original Draft, Writing – Review and Editing. Lachhman Das Singla: Conceptualization, Supervision, Writing – Review and Editing. Diptiman Choudhury: Conceptualization, Supervision, Writing – Review and Editing. All authors have read and approved the final version of the manuscript for publication and agree to be accountable for all aspects of the work, ensuring that questions related to the accuracy or integrity of any part of the work are appropriately investigated and resolved.

This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.

Not applicable.

Not applicable.

Not applicable.

No new datasets were generated or analyzed in this review. All information discussed in this manuscript is derived from published literature cited within the article.

Lachhman Das Singla serves as the Editor-in-Chief and is a co-author of this manuscript. To ensure a rigorous and unbiased peer-review process, he was not involved in any stage start from editorial evaluation, peer review process and final publication decision. The handling of this manuscript was managed independently by another editorial board member. The other authors declare no competing interests.

The authors declare that artificial intelligence (AI)-assisted tools were used only for language refinement, grammar improvement, and manuscript structuring purposes during the preparation of this work. All technical content, experimental implementation, results, and interpretations were independently developed and verified by the authors.

Not applicable.

1. [01] G. A. Castro, Helminths: structure, classification, growth, and development. In: Medical Microbiology, 4th ed., University of Texas Medical Branch at Galveston, Galveston (TX), 2011.
2. [02] F. Varyani, J. O. Fleming, R. M. Maizels, Helminths in the gastrointestinal tract as modulators of immunity and pathology, American Journal of Physiology-Gastrointestinal and Liver Physiology, 2017, 312, G537-G549, doi: 10.1152/ajpgi.00024.2017
3. [03] Z. Greenwood, J. Black, L. Weld, D. O’Brien, K. Leder, F. Von Sonnenburg, P. Pandey, E. Schwartz, B. A. Brown, D. O. Freedman, Gastrointestinal infection among international travelers globally, Journal of travel medicine, 2008, 15, 221-228, doi: 10.1111/j.1708-8305.2008.00203.x
4. [04] R. M, Maizels, H. J. McSorley, Regulation of the host immune system by helminth parasites, Journal of Allergy and Clinical Immunology, 2016, 138, 666-675, doi: 10.1016/j.jaci.2016.07.007
5. [05] B. M. Greenwood, Autoimmune disease and parasitic infections in Nigerians. The Lancet, 1968, 292, 380-382, doi: 10.1016/S0140-6736(68)90595-3
6. [06] R. M. Humphries, A. J. Linscott, Practical guidance for clinical microbiology laboratories: diagnosis of bacterial gastroenteritis, Clinical microbiology reviews, 2015, 28, 3-31, doi: 10.1128/cmr.00073-14
7. [07] N. J. Beesley, D. J. Williams, S. Paterson, J. Hodgkinson, Fasciola hepatica demonstrates high levels of genetic diversity, a lack of population structure and high gene flow: possible implications for drug resistance, International journal for parasitology, 2017, 47, 11-20, doi: 10.1016/j.ijpara.2016.09.007
8. [08] R. Sanabria, Nanotechnological improvement of veterinary anthelmintics, Pharmaceutical Nanotechnology, 2021, 9, 5-14, doi: 10.2174/2211738508666200524233724
9. [09] M. Werkman, J. E. Wright, J. E. Truscott, W. E. Oswald, K. E. Halliday, M. Papaiakovou, S. H. Farrell, R. L. Pullan, R. M. Anderson, The impact of community-wide, mass drug administration on aggregation of soil-transmitted helminth infection in human host populations, Parasites & Vectors, 2020, 13, 290, doi: 10.1186/s13071-020-04149-4
10. [10] W. Grünberg, MSD Manual – Vetrinary Manual, https://www.msdvetmanual.com/authors/gruenberg-walter Accessed 28 July 2026.
11. [11] P. K. Thornton, Livestock production: recent trends, future prospects, Philosophical Transactions of the Royal Society B: Biological Sciences, 2010, 365, 2853, doi: 10.1098/rstb.2010.0134
12. [12] A. J. Wolstenholme, I. Fairweather, R. Prichard, G. Von Samson-Himmelstjerna, N. C. Sangster, Drug resistance in veterinary helminths, Trends in parasitology, 2004, 20, 469–476, doi: 10.1016/j.pt.2004.07.010
13. [13] C. Lanusse, C. Canton, G. Virkel, L. Alvarez, L. Costa-Junior, A. Lifschitz, Strategies to optimize the efficacy of anthelmintic drugs in ruminants, Trends in parasitology, 2018, 34, 664-682, doi: 10.1016/j.pt.2018.05.005
14. [14] B. H. B. Baiak, C. R. Lehnen, R. A. da Rocha, Anthelmintic resistance in cattle: A systematic review and meta-analysis, Livestock Science, 2018, 217, 127-135, doi: 10.1016/j.livsci.2018.09.022
15. [15] I. Scott, W. E. Pomroy, P. R. Kenyon, G. Smith, B. Adlington, A. Moss, Lack of efficacy of monepantel against Teladorsagia circumcincta and Trichostrongylus colubriformis, Veterinary parasitology, 2012, 198, 166-171, doi: 10.1016/j.vetpar.2013.07.037
16. [16] Microtubule/Tubulin Inhibitor, MedChemExpress https://www.medchemexpress.com/Targets/Microtubule_Tubulin/effect/inhibitor.html?locale=ko-KR&page=9 Accessed 20 July 2026.
17. [17] H. Park, W. Lim, S. You, G. Song, Fenbendazole induces apoptosis of porcine uterine luminal epithelial and trophoblast cells during early pregnancy. Science of The Total Environment, 2019, 681, 28-38, doi: 10.1016/j.scitotenv.2019.05.116
18. [18] T. Feyera, I. Ruhnke, B. Sharpe, T. Elliott, A. Shifaw, S. W. Walkden-Brown, Comparative therapeutic efficacies of oral and in-water administered levamisole, piperazine and fenbendazole against experimental Ascaridia galli infection in chickens, Veterinary Parasitology, 2021, 298, 109514, doi: 10.1016/j.vetpar.2021.109514
19. [19] R. K. Thakur, S. P. Patel, Mebendazole. In StatPearls [Internet]. StatPearls Publishing, 2023, https://www.ncbi.nlm.nih.gov/books/NBK557705/ Accessed 29 July 2026.
20. [20] S. J. Emery-Corbin, Q. Su, S. Tichkule, L. Baker, E. Lacey, A. R. Jex, In vitro selection of Giardia duodenalis for Albendazole resistance identifies a β-tubulin mutation at amino acid E198K, International Journal for Parasitology: Drugs and Drug Resistance, 2021, 16, 162-173, doi: 10.1016/j.ijpddr.2021.05.003
21. [21] Y. Zhang, J. Wu, W. Xu, J. Gao, H. Cao, M. Yang, B. Wang, Y. hao, L. Tao, Cytotoxic effects of Avermectin on human HepG2 cells in vitro bioassays, Environmental Pollution, 2017, 220, 1127-1137, doi: 10.1016/j.envpol.2016.11.022
22. [22] E. Courtot, C. L. Charvet, R. N. Beech, A. Harmache, A. J. Wolstenholme, L. Holden-Dye, C. Neveu, Functional characterization of a novel class of morantel-sensitive acetylcholine receptors in nematodes, PLoS Pathogens, 2015, 11, e1005267, doi: 10.1371/journal.ppat.1005267
23. [23] M. Abongwa, K. E. Baber, R. J. Martin, A. P. Robertson, The cholinomimetic morantel as an open channel blocker of the Ascaris suum ACR-16 nAChR, Invertebrate Neuroscience, 2016, 16, 10, doi: 10.1007/s10158-016-0193-4
24. [24] M. Abongwa, R. J. Martin, A. P. Robertson, A brief review on the mode of action of antinematodal drugs, Acta veterinaria, 2017, 67, 137-152, doi: 10.1515/acve-2017-0013
25. [25] R. J. Martin, S. Verma, M. Levandoski, C. L. Clark, H. Qian, M. Stewart, A. P. Robertson, Drug resistance and neurotransmitter receptors of nematodes: recent studies on the mode of action of levamisole, Parasitology, 2005, 131, S71-S84, doi: 10.1017/S0031182005008668
26. [26] J. Liu, K. Zhang, L. Cheng, H. Zhu, T. Xu, Progress in understanding the molecular mechanisms underlying the antitumour effects of ivermectin, Drug design, development and therapy, 2020, 285-296, doi: 10.2147/DDDT.S237393
27. [27] M. O. Harhay, J. Horton, P. L. Olliaro, Epidemiology and control of human gastrointestinal parasites in children, Expert review of anti-infective therapy, 2010, 8, 219-234, doi: 10.1586/eri.09.119
28. [28] M. E. Scott, K. G. Koski, Soil-transmitted helminths: Does nutrition make a difference?, Nutrition and Infectious Diseases: Shifting the Clinical Paradigm, Cham: Springer International Publishing, 2020, 325-364, doi: 10.1007/978-3-030-56913-6_12
29. [29] P. M. Jourdan, P. H. Lamberton, A. Fenwick, D. G. Addiss, Strongyloides stercoralis: the need for accurate information–Authors' reply, The Lancet, 2018, 391, 2323, doi: 10.1016/S0140-6736(18)30858-4
30. [30] P. Dorny, N. Praet, N. Deckers, S. Gabriël, Emerging food-borne parasites, Veterinary parasitology, 2009, 163, 196-206, doi: 10.1016/j.vetpar.2009.05.026
31. [31] B. Douglas, O. Oyesola, M. M. Cooper, A. Possey, E. Tait Wojno, P. R. Giacomin, D. B. R. Herbert, Immune system investigation using parasitic helminths, Annual review of immunology, 2021, 39, 639-665, doi: 10.1146/annurev-immunol-093019-122827
32. [32] S. Bartlett, R. M. Eichenberger, R. J. Nevagi, K. A. Ghaffar, N. Marasini, Y. Dai, A. Loukas, I. Toth, M. Skwarczynski, Lipopeptide-based oral vaccine against hookworm infection, The Journal of Infectious Diseases, 2020, 221, 934-942, doi: 10.1093/infdis/jiz528
33. [33] C. Drurey, R. M. Maizels, Helminth extracellular vesicles: Interactions with the host immune system, Molecular Immunology, 2021, 137, 124-133, doi: 10.1016/j.molimm.2021.06.017
34. [34] E. E. Mouser, G. Pollakis, H. H. Smits, J. Thomas, Y. Yazdanbakhsh, E. C. de Jong, W. A. Paxton, Schistosoma mansoni soluble egg antigen (SEA) and recombinant Omega-1 modulate induced CD4+ T-lymphocyte responses and HIV-1 infection in vitro, PLoS pathogens, 2019, 15, e1007924, doi: 10.1371/journal.ppat.1007924
35. [35] R. E. Wiegand, F. M. Fleming, A. Straily, S. P. Montgomery, S. J. de Vlas, J. Utzinger, P. Vounatsou, W. E. Secor, Urogenital schistosomiasis infection prevalence targets to determine elimination as a public health problem based on microhematuria prevalence in school-age children, PLoS neglected tropical diseases, 2021, 15, e0009451, doi: 10.1371/journal.pntd.0009451
36. [36] A. Olorunlana, Dancing in a cycle: Global Health Agenda and Schistosomiasis control in Africa, In Parasitic Helminths and Zoonoses-From Basic to Applied Research. IntechOpen, 2022.
37. [37] M. Arrais, O. Lulua, F. Quifica, J. Rosado-Pinto, J. M. Gama, P. J. Cooper, L. Taborda-Barata, M. Brito, Lack of consistent association between asthma, allergic diseases, and intestinal helminth infection in school-aged children in the province of bengo, angola, International Journal of Environmental Research and Public Health, 2021, 18, 6156, doi: 10.3390/ijerph18116156
38. [38] G. A. Rook, 99th Dahlem conference on infection, inflammation and chronic inflammatory disorders: Darwinian medicine and the ‘hygiene’or ‘old friends’ hypothesis, Clinical & Experimental Immunology, 2010, 160, 70-79, doi: 10.1111/j.1365-2249.2010.04133.x
39. [39] P. Blair, D. Diemert, Update on prevention and treatment of intestinal helminth infections, Current infectious disease reports, 2015, 17, 12, doi: 10.1007/s11908-015-0465-x
40. [40] S. T. Hong, Albendazole and praziquantel: review and safety monitoring in Korea, Infection & chemotherapy, 2018, 50, 1, doi: 10.3947/ic.2018.50.1.1
41. [41] E. Lacey, Mode of action of benzimidazoles, Parasitology Today, 1990, 6, 112-115, doi: 10.1016/0169-4758(90)90227-U
42. [42] C. M. Thomas, D. J. Timson, The mechanism of action of praziquantel: can new drugs exploit similar mechanisms?, Current Medicinal Chemistry, 2020, 27, 676-696, doi: 10.2174/0929867325666180926145537
43. [43] R. Laing, V. Gillan, E. Devaney, Ivermectin–old drug, new tricks?, Trends in Parasitology, 2017, 33, 463-472, doi: 10.1016/j.pt.2017.02.004
44. [44] S. Geerts, B. Gryseels, Drug resistance in human helminths: current situation and lessons from livestock, Clinical Microbiology Reviews, 2000, 13, 207-222, doi: 10.1128/cmr.13.2.207
45. [45] Q. Y. Li, D. H. Zhao, H. Y. Qu, C. N. Zhou, Life-threatening complications of ascariasis in trauma patients: a review of the literature, World Journal of Emergency Medicine, 2014, 5, 165-170, doi: 10.5847/wjem.j.issn.1920-8642.2014.03.001
46. [46] J. Krücken, K. Fraundorfer, J. C. Mugisha, S. Ramünke, K. C. Sifft, D. Geus, F. Habarugira, J. Ndoli, A. Sendegeya, C. Mukampunga, C. Bayingana, Reduced efficacy of albendazole against Ascaris lumbricoides in Rwandan schoolchildren, International Journal for Parasitology: Drugs and Drug Resistance, 2017, 7, 262-271, doi: 10.1016/j.ijpddr.2017.06.001
47. [47] C. R. Caffrey, Parasitic Helminths: Targets, Screens, Drugs and Vaccines, John Wiley & Sons, 2012, doi: 10.1002/9783527652969
48. [48] A. A. Adegnika, F. Lötsch, R. M. O. Mba, M. Ramharter, Update on treatment and resistance of human trichuriasis, Current Tropical Medicine Reports, 2015, 2, 218-223, doi: 10.1007/s40475-015-0061-z
49. [49] A. Viswanath, S. N. S. Yarrarapu, M. Williams, Trichuris trichiura Infection, StatPearls, StatPearls-NCBI Publishing, 2023.
50. [50] D. Wimmersberger, J. T. Coulibaly, J. D. Schulz, M. Puchkow, J. Huwyler, Y. N’Gbesso, J. Hattendorf, J. Keiser, Efficacy and safety of ivermectin against Trichuris trichiura in preschool-aged and school-aged children: a randomized controlled dose-finding trial, Clinical Infectious Diseases, 2018, 67, 1247-1255, doi: 10.1093/cid/ciy246
51. [51] D. Humphries, S. Nguyen, S. Kumar, J. E. Quagraine, J. Otchere, L. M. Harrison, M. Cappello, Effectiveness of albendazole for hookworm varies widely by community and correlates with nutritional factors: a cross-sectional study of school-age children in Ghana, The American journal of tropical medicine and hygiene, 2017, 96, 347, doi: 10.4269/ajtmh.16-0682
52. [52] A. Loukas, P. J. Hotez, D. Diemert, M. Yazdanbakhsh, J. S. McCarthy, R. Correa-Oliveira, J. Croese, J. M. Bethony, Hookworm infection, Nature reviews Disease primers, 2016, 2, 16088, doi: 10.1038/nrdp.2016.88
53. [53] L. George, M. Harrison, F. P. Dornas, B. Evans, A. Caccone, D. Humphries, M. D. Wilson, M. Cappello, Genetic Markers of Benzimidazole Resistance among Human Hookworms (Necator americanus) in Kintampo North Municipality, Ghana, The American Journal of Tropical Medicine and Hygiene, 2019, 100, 351-356, doi: 10.4269/ajtmh.18-0727
54. [54] L. F. V. Furtado, P. H. Nascimento de Aguiar, L. W. Zuccherato, T. T. G. Teixeira, W. P. Alves, V. Jordania da Silva, R. B Gasser, É. M. L. Rabelo, Albendazole resistance induced in Ancylostoma ceylanicum is not due to single-nucleotide polymorphisms (SNPs) at codons 167, 198, or 200 of the beta-tubulin gene, indicating another resistance mechanism, Parasitology research, 2019, 118, 837-849, doi: 10.1007/s00436-019-06218-9
55. [55] N. Sandoval, M. Siles-Lucas, J. L. Perez-Arellano, C. Carranza, S. Puente, J. López-Abán, A. Muro, A new PCR-based approach for the specific amplification of DNA from different Schistosoma species applicable to human urine samples, Parasitology, 2006, 133, 581-587, doi: 10.1017/S0031182006000898
56. [56] C. H. King, Toward the elimination of schistosomiasis, 2009, New England Journal of Medicine, 360, 106-109, doi: 10.1056/NEJMp0808041
57. [57] A. Tesfie, G. Getnet, A. Abere, G. Yihenew, Y. Belete, M. Kassa, A. Gize, Praziquantel is an effective drug for the treatment of Schistosoma Mansoni infection among school-aged children in Northwest Ethiopia, Tropical medicine and health, 2020, 48, 28, doi: 10.1186/s41182-020-00204-z
58. [58] S. Lustigman, B. L. Makepeace, T. R. Klei, S. A. Babayan, P. Hotez, D. Abraham, M. E. Bottazzi, Onchocerca volvulus: the road from basic biology to a vaccine, Trends in parasitology, 2018, 34, 64-79, doi: 10.1016/j.pt.2017.08.011
59. [59] S. Wendt, H. Trawinski, S. Schubert, A. C. Rodloff, J. Mössner, C. Lübbert, The Diagnosis and Treatment of Pinworm Infection, Deutsches Arzteblatt International, 2019, 116, 213-219, doi: 10.3238/arztebl.2019.0213
60. [60] B. Barda, S. Sayasone, K. Phongluxa, S. Xayavong, K. Keoduangsy, P. Odermatt, M. Puchkov, J. Huwyler, J. Hattendorf, J. Keiser, Efficacy of Moxidectin Versus Ivermectin Against Strongyloides stercoralis Infections: A Randomized, Controlled Noninferiority Trial, Clinical Infectious Diseases, 2017, 65, 276–281, doi: 10.1093/cid/cix278
61. [61] A. Echazú, M. Juarez, P. A. Vargas, S. P. Cajal, R. O. Cimino, V. Heredia, S. Caropresi, G. Paredes, L. M. Arias, M. Abril, S. Gold, P. Lammie, A. J. Krolewiecki, Albendazole and ivermectin for the control of soil-transmitted helminths in an area with high prevalence of Strongyloides stercoralis and hookworm in northwestern Argentina: A community-based pragmatic study, PLOS Neglected Tropical Diseases, 2017, 11, e0006003, doi: 10.1371/journal.pntd.0006003
62. [62] F. Heidary, R. Gharebaghi, Ivermectin: a systematic review from antiviral effects to COVID-19 complementary regimen, The Journal of Antibiotics, 2020, 73, 593–602, doi: 10.1038/s41429-020-0336-z
63. [63] D. Buonfrate, D. Bisanzio, G. Giorli, P. Odermatt, T. Fürst, C. Greenaway, M. French, R. Reithinger, F. Gobbi, A. Montresor, Z. Bisoffi, The Global Prevalence of Strongyloides stercoralis Infection, Pathogens, 2020, 9, 468, doi: 10.3390/pathogens9060468
64. [64] B. G. Yangco, C. De Lerma, G. H. Lyman, D. L. Price, Clinical study evaluating efficacy of praziquantel in clonorchiasis, Antimicrobial agents and chemotherapy, 1987, 31, 135-138, doi: 10.3390/pathogens9060468
65. [65] S. Li, X. Chen, J. Zhou, Z. Xie, M. Shang, L. He, P. Liang, T. Chen, Q. Mao, C. Liang, X. Li, Amino acids serve as an important energy source for adult flukes of Clonorchis sinensis, PLoS Neglected Tropical Diseases, 2020, 14, e0008287, doi: 10.1371/journal.pntd.0008287
66. [66] P. J. Budge, C. Herbert, B. J. Andersen, G. J. Weil, Adverse events following single dose treatment of lymphatic filariasis: Observations from a review of the literature, PLoS neglected tropical diseases, 2018, 12, e0006454, doi: 10.1371/journal.pntd.0006454
67. [67] F. Cobo, Determinants of parasite drug resistance in human lymphatic filariasis, Revista Española de Quimioterapia, 2016, 29, 288, doi: 10.1371/journal.pntd.0006454
68. [68] C. Whittaker, C. B. Chesnais, S. D. S. Pion, J. Kamgno, M. Walker, M. G. Basáñez, M. Boussinesq, Factors associated with variation in single-dose albendazole pharmacokinetics: A systematic review and modelling analysis, PLoS neglected tropical diseases, 2022, 16, e0010497, doi: 10.1371/journal.pntd.0010497
69. [69] L. C. McDonald, , D. N. Gerding, S. Johnson, J. S. Bakken, K. C. Carroll, S. E. Coffin, E. R. Dubberke, K.W. Garey, C.V. Gould, C. Kelly, V. Loo, Clinical practice guidelines for Clostridium difficile infection in adults and children: 2017 update by the Infectious Diseases Society of America (IDSA) and Society for Healthcare Epidemiology of America (SHEA), Clinical infectious diseases, 2018, 66, e1-e48, doi: 10.1093/cid/cix1085
70. [70] I. Fairweather, G. P. Brennan, R. E. B. Hanna, M. W. Robinson, P. J. Skuce, Drug resistance in liver flukes, International Journal for Parasitology: drugs and drug resistance, 2020, 12, 39-59, doi: 10.1016/j.ijpddr.2019.11.003
71. [71] K. Thinkhamrop, N. Khuntikeo, P. Sithithaworn, W. Thinkhamrop, K. Wangdi, M. J. Kelly, A. T. Suwannatrai, D. J. Gray, Repeated praziquantel treatment and Opisthorchis viverrini infection: a population-based cross-sectional study in northeast Thailand, Infectious Diseases of Poverty, 2019, 8, 18, doi: 10.1186/s40249-019-0529-5
72. [72] M. Bhaibulaya, P. Punthuprapasa, Treatment of Opisthorchiasis viverrini in hamsters with albendazole, The Southeast Asian Journal of Tropical Medicine and Public Health, 1984, 15, 389-393.
73. [73] A. Suwannatrai, P. Saichua, M. Haswell, Epidemiology of Opisthorchis viverrini infection, Advances in Parasitology, 2018, 101, 41-67, doi: 10.1016/bs.apar.2018.05.002
74. [74] P. A. Soukhathammavong, S. Sayasone, K. Phongluxa, V. Xayaseng, J. Utzinger, P. Vounatsou, C. Hatz, K. Akkhavong, J. Keiser, P. Odermatt, Low efficacy of single-dose albendazole and mebendazole against hookworm and effect on concomitant helminth infection in Lao PDR, PLoS neglected tropical diseases, 2012, 6, e1417, doi: 10.1371/journal.pntd.0001417
75. [75] A. C. Kotze, R. K. Prichard, Anthelmintic resistance in Haemonchus contortus: history, mechanisms and diagnosis, Advances in parasitology, 2016, 93, 397-428, doi: 10.1016/bs.apar.2016.02.012
76. [76] S. Puspitasari, A. Farajallah, E. Sulistiawati, Muladno, Effectiveness of Ivermectin and Albendazole against Haemonchus contortus in Sheep in West Java, Indonesia, Tropical life sciences research, 2016, 27, 135–144.
77. [77] N. Dogra, A. Kumar, T. Mukhopadhyay, Fenbendazole acts as a moderate microtubule destabilizing agent and causes cancer cell death by modulating multiple cellular pathways, Scientific reports, 2018, 8, 11926, doi: 10.1038/s41598-018-30158-6
78. [78] V. Goel, L. D. Singla, D. Choudhury, Cuminaldehyde induces oxidative stress-mediated physical damage and death of Haemonchus contortus, Biomedicine & Pharmacotherapy, 2020, 130, 110411, doi: 10.1016/j.biopha.2020.110411
79. [79] N. O. Opoku, D. K. Bakajika, E. M. Kanza, H. Howard, G. L. Mambandu, A. Nyathirombo, M. M. Nigo, K. Kasonia, S. L. Masembe, M.. Mumbere, K. Kataliko, J. P. Larbelee, M. Kpawor, K. M. Bolay, F. Bolay, S. Asare, S. K. Attah, G. Olipoh, M. Vaillant, C. M. Halleux, A. C. Kuesel, Single dose moxidectin versus ivermectin for Onchocerca volvulus infection in Ghana, Liberia, and the Democratic Republic of the Congo: a randomised, controlled, double-blind phase 3 trial, The Lancet, 392, 1207-1216, doi: 10.1016/S0140-6736(17)32844-1
80. [80] P. Gandhi, E. K. Schmitt, C. Chen, S. Samantray, V. K. Venishetty, D. Hughes, Triclabendazole in the treatment of human fascioliasis: a review, Transactions of the Royal Society of Tropical Medicine and Hygiene, 2019, 113, 797-804, doi: 10.1093/trstmh/trz093
81. [81] D. S. Ashour, A. A. Othman, D. A. Radi, Insights into regulatory molecules of intestinal epithelial cell turnover during experimental infection by Heterophyes heterophyes, Experimental Parasitology, 2014, 143, 48-54, doi: 10.1016/j.exppara.2014.05.003
82. [82] C. R. Wang, J. F. Gao, X. Q. Zhu, Q. Zhao, Characterization of Bunostomum trigonocephalum and Bunostomum phlebotomum from sheep and cattle by internal transcribed spacers of nuclear ribosomal DNA, Research in Veterinary Science, 2012, 92, 99-102, doi: 10.1016/j.rvsc.2010.10.024
83. [83] A. Morrone, G. Franco, R. Fazio, M. Valenzano, R. Calcaterra, Bullous cutaneous larva migrans, Acta Dermatovenerologica Croatica, 2011, 19.
84. [84] M. Shaharyar, A. Mazumder, Benzimidazoles: A biologically active compounds, Arabian Journal of Chemistry, 2017, 10, S157-S173, doi: 10.1016/j.arabjc.2012.07.017
85. [85] Y. Li, Y. Chen, L. Wang, Y. Liu, W. Wang, X. Zhou, J. Yi, Z. Zuo, Y. Xie, The complete mitochondrial genome of the beef cattle hookworm Bunostomum phlebotomum (Nematoda: Bunostominae), Mitochondrial DNA Part B Resources, 2021, 6, 617–619, doi: 10.1080/23802359.2021.1875918
86. [86] K. Ashrafi, M. Sharifdini, Z. Heidari, B. Rahmati, E. B. Kia, Zoonotic transmission of Teladorsagia circumcincta and Trichostrongylus species in Guilan province, northern Iran: molecular and morphological characterizations, BMC Infectious Diseases, 2020, 20, 28, doi: 10.1186/s12879-020-4762-0
87. [87] M. A. Saeed, I. Beveridge, G. Abbas, A. Beasley, J. Bauquier, E. Wilkes, C. Jacobson, K. J. Hughes, C. El-Hage, R. O'Handley, J. Hurley, L. Cudmore, P. Carrigan, L. Walter, B. Tennent-Brown, M. K. Nielsen, A. Jabbar, Systematic review of gastrointestinal nematodes of horses from Australia, Parasites & Vectors, 2019, 12, 188, doi: 10.1186/s13071-019-3445-4
88. [88] C. Palcy, A. Silvestre, C. Sauve, J. Cortet, J. Cabaret, Benzimidazole resistance in Trichostrongylus axei in sheep: long-term monitoring of affected sheep and genotypic evaluation of the parasite, The Veterinary Journal, 2010, 183, 68-74, doi: 10.1016/j.tvjl.2008.09.012
89. [89] M. Schnyder, P. R. Torgerson, M. Schönmann, L. Kohler, H. Hertzberg, Multiple anthelmintic resistance in Haemonchus contortus isolated from South African Boer goats in Switzerland, Veterinary Parasitology, 2005, 128, 285-290, doi: 10.1016/j.vetpar.2004.12.010
90. [90] S. A. Seddiek, M. M. Ali, H. F. Khater, M. M. El-Shorbagy, Anthelmintic activity of the white wormwood, Artemisia herba-alba against Heterakis gallinarum infecting turkey poults, Journal of Medicinal Plants Research, 2011, 5, 3946-3957, doi: 10.5897/JMPR.9000085
91. [91] A. H. Ahmed, M. Ejo, T. Feyera, D. Regassa, B. Mummed, S. A. Huluka, In vitro anthelmintic activity of crude extracts of Artemisia herba-alba and Punica granatum against Haemonchus contortus, Journal of Parasitology Research, 2020, 2020, 4950196, doi: 10.1155/2020/4950196
92. [92] M. Lateef, Z. Iqbal, M. S. Sajid, R. Z. Abbas, Z. U. D. Sindhu, M. Akhtar, M. N. Khan, M. M. Awais, A. Iqbal, Q. U. Ain, An account of botanical anthelmintics and methods used for their evaluation, Reviews in Veterinary and Animal Sciences, 2013, 1, 6-14.
93. [93] M. A. Dkhil, Anti-coccidial, anthelmintic and antioxidant activities of pomegranate (Punica granatum) peel extract, Parasitology Research, 2013, 112, 2639-2646, doi: 10.1007/s00436-013-3430-3
94. [94] R. Aggarwal, K. Kaur, M. Suri, U. Bagai, Anthelmintic potential of Calotropis procera, Azadirachta indica and Punica granatum against Gastrothylax indicus, Journal of Parasitic Diseases, 2016, 40, 1230-1238, doi: 10.1007/s00436-013-3430-3
95. [95] F. Castagna, D. Britti, M. Oliverio, A. Bosco, S. Bonacci, G. Iriti, M. Ragusa, V. Musolino, L. Rinaldi, E. Palma, V. Musella, In vitro anthelminthic efficacy of aqueous pomegranate (Punica granatum L.) extracts against gastrointestinal nematodes of sheep, Pathogens, 2020, 9, 1063, doi: 10.3390/pathogens9121063
96. [96] R. Kermanshai, B. E. McCarry, J. Rosenfeld, P. S. Summers, E. A. Weretilnyk, G. J. Sorger, Benzyl isothiocyanate is the chief or sole anthelmintic in papaya seed extracts, Phytochemistry, 2001, 57, 427-435, doi: 10.1016/S0031-9422(01)00077-2 .
97. [97] M. S. Hounzangbe-Adote, V. Paolini, I. Fouraste, K. Moutairou, H. Hoste, In vitro effects of four tropical plants on three life-cycle stages of the parasitic nematode, Haemonchus contortus, Research in Veterinary Science, 2005, 78, 155-160, doi: 10.1016/j.rvsc.2004.05.009
98. [98] P. J. Wabo, N. J. Ngankam, B. C. Bilong, M. Mpoame, A comparative study of the ovicidal and larvicidal activities of aqueous and ethanolic extracts of pawpaw seeds Carica papaya (Caricaceae) on Heligmosomoides bakeri, Asian Pacific Journal of Tropical Medicine, 2011, 4, 447-450, doi: 10.1016/S1995-7645(11)60123-5
99. [99] R. K. Bauri, M. N. Tigga, S. S. Kullu, A review on use of medicinal plants to control parasites, Indian Journal of Natural Products and Resources, 2015, 6, 268-277.
100. [100] T. O. Ajiboye, A. T. Kuvarega, D. C. Onwudiwe, Recent strategies for environmental remediation of organochlorine pesticides, Applied Sciences, 2020, 10, 6286, doi: 10.3390/app10186286
101. [101] O. A. Idris, O. A. Wintola, A. J. Afolayan, Helminthiases; prevalence, transmission, host-parasite interactions, resistance to common synthetic drugs and treatment, Heliyon, 2019, 5, e01161, doi: 10.1016/j.heliyon.2019.e01161
102. [102] N. Jeyathilakan, K. Murali, A. Anandaraj, S. Abdul Basith, In vitro evaluation of anthelmintic property of ethno-veterinary plant extracts against the liver fluke Fasciola gigantica, Journal of Parasitic Diseases, 2012, 36, 26–30, doi: 10.1007/s12639-011-0064-1
103. [103] O. A. Idris, O. A. Wintola, A. J. Afolayan, Helminthiases; prevalence, transmission, host-parasite interactions, resistance to common synthetic drugs and treatment, Heliyon, 2019, 5, e01161, doi: 10.1016/j.heliyon.2019.e01161
104. [104] J. Palacios-Landín, P. Mendoza-de Gives, D. O. Salinas-Sánchez, M. E. López-Arellano, E. Liébano-Hernández, V. M. Hernández-Velázquez, M. G. Valladares-Cisneros, In vitro and in vivo nematocidal activity of Allium sativum and Tagetes erecta extracts against Haemonchus contortus, Türkiye Parazitoloji Dergisi, 2015, 39, 260–265, doi: 10.5152/tpd.2015.4523
105. [105] R. V. Marques, Bryophytes, a source of inspiration for active ingredients discovery, 2021, https://orbit.dtu.dk/en/publications/bryophytes-a-source-of-inspiration-for-active-ingredients-discove/ Accessed 27 July 2026.
106. [106] D. Ramdani, E. Yuniarti, A. Jayanegara, A. S. Chaudhry, Roles of essential oils, polyphenols, and saponins of medicinal plants as natural additives and anthelmintics in ruminant diets: A systematic review, Animals, 2023, 13, 767, doi: 10.3390/ani13040767
107. [107] M. S. Akhtar, Z. Iqbal, M. N. Khan, M. Lateef, Anthelmintic activity of medicinal plants with particular reference to their use in animals in the Indo–Pakistan subcontinent, Small Ruminant Research, 2000, 38, 99–107, doi: 10.1016/S0921-4488(00)00163-2
108. [108] Z. Iqbal, Q. K. Nadeem, M. N. Khan, M. S. Akhtar, F. N. Waraich, In vitro anthelmintic activity of Allium sativum, Zingiber officinale, Curcurbita mexicana and Ficus religiosa, International Journal of Agriculture and Biology, 2001, 3, 454–457.
109. [109] C. A. Ukwubile, Anti-helminthic properties of some Nigerian medicinal plants on selected intestinal worms in children (age 5–13) in Ogurugu, South East Nigeria, Journal of Bacteriology & Parasitology, 2012, 3, 159, doi: 10.4172/2155-9597.1000159
110. [110] M. Tagoe, Y. D. Boakye, T. A. Agana, V. E. Boamah, C. Agyare, In vitro anthelmintic activity of ethanol stem bark extract of Albizia ferruginea (Guill. & Perr.) Benth., Journal of Parasitology Research, 2021, 6690869, doi: 10.1155/2021/6690869
111. [111] A. M. Mohamed, N. M. Metwally, S. S. Mahmoud, Sativa seeds against Schistosoma mansoni different stages, Memórias do Instituto Oswaldo Cruz, 2005, 100, 205–211, doi: 10.1590/S0074-02762005000200016
112. [112] L. Nie, H. Song, A. He, S. Yao, Recent research progress in natural bioactive constituents against lipid metabolic diseases, Current Topics in Medicinal Chemistry, 2016, 16, 2605–2624.
113. [113] K. G. Koorse, S. S. Sujith Samraj, P. J. Preethy John, P. M. Narayanan, D. S. T. S. Thampy, U. P. T. Ayyappan, G. V. Lonappan, Anthelmintic activity of fruit extract and fractions of Piper longum L. in vitro., Pharmacognosy Journal, 2018,10, 333-340, doi: 10.5530/pj.2018.2.57
114. [114] P. Pattanayak, P. Behera, D. Das, S. K. Panda, Ocimum sanctum Linn. A reservoir plant for therapeutic applications: An overview, Pharmacognosy Reviews, 2010, 4, 95–105, doi: 10.4103/0973-7847.65323
115. [115] N. Bano, A. Ahmed, M. Tanveer, G. M. Khan, M. T. Ansari, Pharmacological evaluation of Ocimum sanctum, Journal of Bioequivalence & Bioavailability, 2017, 9, 387–392, doi: 10.4172/jbb.1000330
116. [116] M. Mahardika, Anthelmintic activity of Ocimum sanctum Linn. leaves ethanol extract against Fasciola gigantica in vitro, In The Veterinary Medicine International Conference, KnE Life Sciences, 2017, 266-277, doi: 10.18502/kls.v3i6.1135
117. [117] U. J. Njoga, I. F. Jaja, O. S. Onwuka, S. U. Ilo, I. G. Eke, K. O. Abah, I. S. Ochiogu, Reproductive effects of medicinal plant (Azadirachta indica) used as forage and for ethnoveterinary practices: New insights from animal models, Challenges, 2022, 13, 40, doi: 10.3390/challe13020040
118. [118] G. Khuwaja, A. A. Chaudhary, A. M. Mashlawi, A. A. Alamri, F. Alfifi, K. Anjum, M. S. Alam, M. I. Alam, S. K. Ali, N. Raza, M. A. M. Ali, M. Imran, Biosynthesis of zinc oxide nanostructures using leaf extract of Azadirachta indica: Characterizations and in silico and nematicidal potentials, Catalysts, 2025, 15, 693, doi: 10.3390/catal15070693
119. [119] A. Azra, M. Kaleemullah, B. Khattak, N. Asma, A. U. R. Safi, J. Qaiser, M. Afzal, U. Tahir, Z. U. D. Sindhu, Y. Farhan, Comparative efficacy of domestic garlic (Allium sativum) and neem (Azadirachta indica) against Haemonchus contortus in small ruminants, Applied Ecology and Environmental Research, 2019, 17, 10389–10397, doi: 10.15666/aeer/1705_1038910397
120. [120] P. Pramu, R. W. Nurcahyo, J. Prastowo, I. Widiyono, Potential of secondary metabolite compounds of sweet potato (Ipomoea batatas) leaves as anthelmintic against Haemonchus contortus, Open Veterinary Journal, 2025, 15, 5538–5548, doi: 10.5455/OVJ.2025.v15.i11.10
121. [121] J. F. Islas, E. Acosta, Z. G. Buentello, J. L. Delgado-Gallegos, M. G. Moreno-Treviño, B. Escalante, J. E. Moreno-Cuevas, An overview of neem (Azadirachta indica) and its potential impact on health, Journal of Functional Foods, 2020, 74, 104171, doi: 10.1016/j.jff.2020.104171
122. [122] C. G. Stevens, F. Ugese, G. Otitoju, K. Baiyeri, Proximate and anti-nutritional composition of leaves and seeds of Moringa oleifera in Nigeria: A comparative study, Agro-Science, 2016, 14, 9–14, doi: 10.4314/as.v14i2.2
123. [123] A. M. Ibrahim, A. M. Abdalla, Impact of Moringa oleifera seed aqueous extract on some biological, biochemical, and histological aspects of Biomphalaria alexandrina snails, Environmental Science and Pollution Research, 2017, 24, 28072–28078, doi: 10.1007/s11356-017-0397-0
124. [124] D. E. Cabardo, H. P. Portugaliza, Anthelmintic activity of Moringa oleifera seed aqueous and ethanolic extracts against Haemonchus contortus eggs and third stage larvae, International Journal of Veterinary Science and Medicine, 2017, 5, 30–34, doi: 10.1016/j.ijvsm.2017.01.001
125. [125] J. Pedraza-Hernández, M. M. M. Y. Elghandour, A. Khusro, M. Z. M. Salem, L. M. Camacho-Diaz, A. Barbabosa-Pliego, A. Z. M. Salem, Assessment on bioactive role of Moringa oleifera leaves as anthelmintic agent and improved growth performance in goats, Tropical Animal Health and Production, 2021, 53, 318, doi: 10.1007/s11250-021-02745-9
126. [126] L. Drăgan, A. Györke, J. F. Ferreira, I. A. Pop, I. Dunca, M. Drăgan, V. Cozma, Effects of Artemisia annua and Foeniculum vulgare on chickens highly infected with Eimeria tenella (Phylum Apicomplexa), Acta Veterinaria Scandinavica, 2014, 56, 22, doi: 10.1186/1751-0147-56-22
127. [127] A. H. Khan, K. H. Bulbul, R. A. Shahardar, Z. A. Wani, I. M. Allaie, Prospects of botanical dewormers with especial reference to Kashmir, International Journal of Veterinary Science and Animal Husbandry, 2021, 6, 58–64, doi: 10.22271/veterinary.2021.v6.i4a.372
128. [128] S. Sharma, V. Goel, P. Kaur, K. Gadhave, N. Garg, L. D. Singla, D. Choudhury, Targeted drug delivery using beeswax-derived albendazole-loaded solid lipid nanoparticles in Haemonchus contortus, an albendazole-tolerant nematode, Experimental Parasitology, 2023, 253, 108593, doi: 10.1016/j.exppara.2023.108593
129. [129] P. K. Pandey, F. Jamal, Identification and evaluation of trypsin inhibitor in the seed extracts of Butea monosperma (Flame of Forest), International Journal of Biotechnology and Biochemistry, 2010, 6, 513–520.
130. [130] H. A. Bachaya, Z. Iqbal, M. N. Khan, A. Jabbar, A. H. Gilani, I. U. Din, In vitro and in vivo anthelmintic activity of Terminalia arjuna bark, International Journal of Agriculture & Biology, 2009, 11, 273–278.
131. [131] B. Kundu, N. E. Kurland, S. Bano, C. Patra, F. B. Engel, V. K. Yadavalli, S. C. Kundu, Silk proteins for biomedical applications: Bioengineering perspectives, Progress in Polymer Science, 2014, 39, 251–267, doi: 10.1016/j.progpolymsci.2013.06.002
132. [132] A. Amalraj, A. Pius, S. Gopi, S. Gopi, Biological activities of curcuminoids, other biomolecules from turmeric and their derivatives – A review, Journal of Traditional and Complementary Medicine, 2017, 7, 205–233, doi: 10.1016/j.jtcme.2016.05.005
133. [133] S. Bano, A. Intisar, M. Rauf, A. Ghaffar, F. Yasmeen, W. U. Zaman, A. Aamir, Comparative analysis of oil composition and antibacterial activity of aerial parts of Terminalia arjuna (Roxb.), Natural Product Research, 2020, 34, 1311–1314, doi: 10.1080/14786419.2018.1557656
134. [134] S. A. Nirmal, A. S. Girme, R. D. Bhalke, Major constituents and anthelmintic activity of volatile oils from leaves and flowers of Cymbopogon martini Roxb, Natural Product Research, 2007, 21, 1217–1220, doi: 10.1080/14786410701552152
135. [135] L. M. Katiki, A. D. S. Chagas, H. R. Bizzo, J. F. S. Ferreira, A. F. T. D. T. Amarante, Anthelmintic activity of Cymbopogon martinii, Cymbopogon schoenanthus and Mentha piperita essential oils evaluated in four different in vitro tests, Veterinary Parasitology, 2011, 183, 103–108, doi: 10.1016/j.vetpar.2011.06.001
136. [136] I. M. Villaseñor, A. C. Sanchez, Menthalactone, a new analgesic from Mentha cordifolia Opiz. leaves, Zeitschrift für Naturforschung C, 2009, 64, 809–812, doi: 10.1515/znc-2009-11-1209
137. [137] Z. K. Hamdan, M. F. Namik, N. A. Mahmood, M. I. Soliman, M. A. Abdel-Rasol, A. H. Nigm, Mechanistic insights into Calotropis procera extract and green-silver nanoparticles as therapeutic agents in murine schistosomiasis: Targeting BAX/Bcl-2 and oxidative stress, Journal of Advanced Veterinary and Animal Research, 2025, 12, 1314–1332, doi: 10.5455/javar.2025.1992
138. [138] G. S. Cavalcante, S. M. D. Morais, W. P. P. André, J. V. D. Araújo-Filho, C. R. Muniz, L. O. D. Rocha, W. L. C. Ribeiro, A. L. M. Rodrigues, L. M. B. de Oliveira, C. M. L. Bevilaqua, M. V. Ramos, Chemical constituents of Calotropis procera latex and ultrastructural effects on Haemonchus contortus, Revista Brasileira de Parasitologia Veterinária, 2020, 29, e001320, doi: 10.1590/S1984-29612020045
139. [139] A. O. Aremu, J. F. Finnie, J. van Staden, Potential of South African medicinal plants used as anthelmintics – Their efficacy, safety concerns and reappraisal of current screening methods, South African Journal of Botany, 2012, 82, 134–150, doi: 10.1016/j.sajb.2012.05.006
140. [140] M. Mwale, P. J. Masika, In vivo anthelmintic efficacy of Aloe ferox, Agave sisalana, and Gunnera perpensa in village chickens naturally infected with Heterakis gallinarum, Tropical Animal Health and Production, 2015, 47, 131–138, doi: 10.1007/s11250-014-0696-0
141. [141] A. Darwish, A. J. Mohamed, S. H. Faraj, A. El-Sayed, M. A. Alghamdi, A. M. Sallam, A. Eissa, B. F. Farag, Y. Kamel, E. M. Embaby, A. Ateya, Analysis of potential genes, oxidative, metabolic, and hormonal markers associated with postpartum disorder susceptibility in Barki sheep (Ovis aries), Veterinary Sciences, 2025, 12, 219, doi: 10.3390/vetsci12030219
142. [142] M. Liu, S. K. Panda, W. Luyten, Plant-based natural products for the discovery and development of novel anthelmintics against nematodes, Biomolecules, 2020, 10, 426, doi: 10.3390/biom10030426
143. [143] G. X. Wang, Z. Zhou, D. X. Jiang, J. Han, J. F. Wang, L. W. Zhao, J. Li, In vivo anthelmintic activity of five alkaloids from Macleaya microcarpa (Maxim) Fedde against Dactylogyrus intermedius in Carassius auratus, Veterinary Parasitology, 2010, 171, 305–313, doi: 10.1016/j.vetpar.2010.03.018
144. [144] H. Huang, J. Yao, K. Liu, W. Yang, G. Wang, C. Shi, G. Yang, Sanguinarine has anthelmintic activity against the enteral and parenteral phases of Trichinella infection in experimentally infected mice, Acta Tropica, 2020, 201, 105226, doi: 10.1016/j.actatropica.2019.105226
145. [145] A. Novobilský, E. Stringano, C. H. Carbonero, L. M. J. Smith, H. L. Enemark, I. Mueller-Harvey, S. M. Thamsborg, In vitro effects of extracts and purified tannins of sainfoin (Onobrychis viciifolia) against two cattle nematodes, Veterinary Parasitology, 2013, 196, 532–537, doi: 10.1016/j.vetpar.2013.03.022
146. [146] O. Desrues, I. Mueller-Harvey, W. F. Pellikaan, H. L. Enemark, S. M. Thamsborg, Condensed tannins in the gastrointestinal tract of cattle after sainfoin (Onobrychis viciifolia) intake and their possible relationship with anthelmintic effects, Journal of Agricultural and Food Chemistry, 2017, 65, 1420–1427, doi: 10.1021/acs.jafc.6b05130
147. [147] M. Liu, S. K. Panda, W. Luyten, Plant-based natural products for the discovery and development of novel anthelmintics against nematodes, Biomolecules, 2020, 10, 426, doi: 10.3390/biom10030426
148. [148] P. S. Shukla, P. S. Shukla, B. Gopalakrishna, Investigation of in-vitro anthelmintic activity of cissampelos pareira linn against Pheretima posthuma, International Journal of Pharmaceutical Sciences and Research, 2012, 3, 265–267, doi: 10.13040/IJPSR.0975-8232.3(1).265-67
149. [149] K. Lalchhandama, B. Roy, B. K. Dutta, In vitro anthelmintic activity of Acacia oxyphylla: Changes in the levels of trace elements and activities of the tegumental enzymes of the cestode, Raillietina echinobothrida, Pharmacologyonline, 2007, 2, 307–317.
150. [150] K. Lalchhandama, B. Roy, B. K. Dutta, Anthelmintic activity of Acacia oxyphylla stem bark against Ascaridia galli, Pharmaceutical Biology, 2009, 47, 578–583, doi: 10.1080/13880200902902463
151. [151] S. Dasgupta, B. Roy, Antiparasitic activity of methanolic extract of Acacia oxyphylla (Leguminosae) against Raillietina echinobothrida, Journal of Parasitic Diseases, 2010 34,14-19, doi: 10.1007/s12639-010-0001-8
152. [152] H. Ahmed, S. G. Kilinc, F. Celik, H. K. Kesik, S. Simsek, K. S. Ahmad, M. S. Afzal, S. Farrakh, W. Safdar, F. Pervaiz, S. Liaqat, J. Zhang, J. Cao, An inventory of anthelmintic plants across the globe, Pathogens, 2023 12, 131, doi: 10.3390/pathogens12010131
153. [153] A. Swargiary, A. Daimari, M. Daimari, N. Basumatary, E. Narzary, Phytochemicals, antioxidant, and anthelmintic activity of selected traditional wild edible plants of lower Assam, Indian journal of pharmacology, 2016, 48, 418, doi: 10.4103/0253-7613.186212
154. [154] H. O. Kim, S. W. Park, H. D. Park, Inactivation of Escherichia coli O157: H7 by cinnamic aldehyde purified from Cinnamomum cassia shoot, Food Microbiology, 2004, 21, 105-110, doi: 10.1016/S0740-0020(03)00010-8
155. [155] A. R. Williams, A. Ramsay, T. V. Hansen, H. M. Ropiak, H. Mejer, P. Nejsum, I. Mueller-Harvey, S. M. Thamsborg, Anthelmintic activity of trans-cinnamaldehyde and A-and B-type proanthocyanidins derived from cinnamon (Cinnamomum verum), Scientific Reports, 2015, 5, 14791, doi: 10.1038/srep14791
156. [156] M. A. Chama, H. A. Onyame, C. Fleischer, D. Osei-Safo, R. Waibel, J. Otchere, I. Addae-Mensah, M. Wilson, In vitro activities of crude extracts and triterpenoid constituents of Dichapetalum crassifolium Chodat against clinical isolates of Schistosoma haematobium, Heliyon, 2020, 6, e04460, doi: 10.1016/j.heliyon.2020.e04460
157. [157] P. E. Pensel, M. A. Maggiore, L. B. Gende, M. J. Eguaras, M. G. Denegri, M. C. Elissondo, Efficacy of essential oils of Thymus vulgaris and Origanum vulgare on Echinococcus granulosus, Interdisciplinary perspectives on infectious diseases, 2014, 2014, 693289, doi: 10.1155/2014/693289
158. [158] L. E. Ferreira, B. I. Benincasa, A. L. Fachin, S. C. Franca, S. S. Contini, A. C. Chagas, R. O. Beleboni, Thymus vulgaris L. essential oil and its main component thymol: Anthelmintic effects against Haemonchus contortus from sheep, Veterinary parasitology, 2016, 228, 70-76, doi: 10.1016/j.vetpar.2016.08.011
159. [159] C. Gómez-Rincón, E. Langa, P. Murillo, M. S. Valero, C. Berzosa, V. López, Activity of tea tree (Melaleuca alternifolia) essential oil against L3 larvae of Anisakis simplex, BioMed research international, 2014, 2014, 549510, doi: 10.1155/2014/54951010.1155/2014/549510
160. [160] N. S. Lam, X. Long, X. Z. Su, F. Lu, Melaleuca alternifolia (tea tree) oil and its monoterpene constituents in treating protozoan and helminthic infections, Biomedicine & Pharmacotherapy, 2020, 130, 110624, doi: 10.1016/j.biopha.2020.110624
161. [161] P. Wangchuk, M. S. Pearson, P. R. Giacomin, L. Becker, J. Sotillo, D. Pickering, M. J. Smout, A. Loukas, Compounds derived from the Bhutanese daisy, Ajania nubigena, demonstrate dual anthelmintic activity against Schistosoma mansoni and Trichuris muris, PLoS Neglected Tropical Diseases, 2016, 10, e0004908, doi: 10.1371/journal.pntd.0004908
162. [162] S. Preston, P. K. Korhonen, L. Mouchiroud, M. Cornaglia, S. L. McGee, N. D. Young, R. A. Davis, S. Crawford, C. Nowell, B. R. Ansell, and G. M. Fisher, Deguelin exerts potent nematocidal activity via the mitochondrial respiratory chain, The FASEB Journal, 2017, 31, 4515–4532, doi: 10.1096/fj.201700288R
163. [163] H. Akkari, O. Ezzine, S. Dhahri, F. B’chir, M. Rekik, S. Hajaji, M. Gharbi, Chemical composition, insecticidal and in vitro anthelmintic activities of Ruta chalepensis (Rutaceae) essential oil, Industrial Crops and Products, 2015, 74, 745–751, doi: 10.1016/j.indcrop.2015.06.008
164. [164] L. Nahar, H. R. El-Seedi, S. A. Khalifa, M. Mohammadhosseini, S. D. Sarker, Ruta essential oils: Composition and bioactivities, Molecules, 2021, 26, 4766, doi: 10.3390/molecules26164766
165. [165] E. von Son-de Fernex, M. Á. Alonso-Díaz, B. Valles-de la Mora, P. Mendoza-de Gives, M. González-Cortazar, A. Zamilpa, Anthelmintic effect of 2H-chromen-2-one isolated from Gliricidia sepium against Cooperia punctata, Experimental Parasitology, 2017, 178, 1–6, doi: 10.1016/j.exppara.2017.04.013
166. [166] M. Doligalska, K. Jóźwicka, K. Donskow-Łysoniewska, M. Kalinowska, The antiparasitic activity of avenacosides against intestinal nematodes, Veterinary Parasitology, 2017, 241, 5–13, doi: 10.1016/j.vetpar.2017.05.003
167. [167] G. J. Díaz, G. T. Hernández, A. Zamilpa, C. M. B. Pérez, J. E. R. Bribiesca, O. H. Mendo, P. M. de Gives, In vitro assessment of Argemone mexicana, Taraxacum officinale, Ruta chalepensis and Tagetes filifolia against Haemonchus contortus nematode eggs and infective (L3) larvae, Microbial Pathogenesis, 2017, 109, 162–168, doi: 10.1016/j.micpath.2017.05.048
168. [168] G. F. Castillo-Mitre, A. Olmedo-Juárez, R. Rojo-Rubio, M. González-Cortázar, P. Mendoza-de Gives, E. E. Hernández-Beteta, A. Zamilpa, Caffeoyl and coumaroyl derivatives from Acacia cochliacantha exhibit ovicidal activity against Haemonchus contortus, Journal of Ethnopharmacology, 2017, 204, 125–131, doi: 10.1016/j.jep.2017.04.010
169. [169] S. S. Santa Rosa, F. O. Santos, H. G. Lima, I. M. A. Reis, D. S. A. Cassiano, I. J. C. Vieira, M. J. M. Batatinha, in vitro anthelmintic and cytotoxic activities of extracts of Persea willdenovii Kosterm (Lauraceae), Journal of Helminthology, 2018, 92, 674–680, doi: 10.1017/S0022149X17000979
170. [170] L. F. Wanderley, A. M. D. S. Soares, C. R. E. Silva, I. M. D. Figueiredo, A. T. D. S. Ferreira, J. Perales, H. R. D. O. Mota, J. T. A. Oliveira, L. M. Costa Junior, A cysteine protease from the latex of Ficus benjamina has in vitro anthelmintic activity against Haemonchus contortus, Revista Brasileira de Parasitologia Veterinária, 2018, 27(4), 473–480, doi: 10.1590/S1984-296120180070
171. [171] J. A. Cortes-Morales, A. Olmedo-Juárez, G. Trejo-Tapia, M. González-Cortazar, B. E. Domínguez-Mendoza, P. Mendoza-de Gives, and A. Zamilpa, In vitro ovicidal activity of Baccharis conferta Kunth against Haemonchus contortus, Experimental Parasitology, 2019, 197, 20–28, doi: 10.1016/j.exppara.2019.01.003
172. [172] G. S. Castañeda-Ramírez, J. F. de Jesús Torres-Acosta, C. A. Sandoval-Castro, R. Borges-Argáez, M. Cáceres-Farfán, G. Mancilla-Montelongo, C. Mathieu, Bio-guided fractionation to identify Senegalia gaumeri leaf extract compounds with anthelmintic activity against Haemonchus contortus eggs and larvae, Veterinary Parasitology, 2019, 270, 13–19, doi: 10.1016/j.vetpar.2019.05.001
173. [173] S. E. Payne, G. R. Flematti, A. Reeder, A. C. Kotze, Z. Durmic, P. E. Vercoe, Procyanidin A2 in the Australian plant Alectryon oleifolius has anthelmintic activity against equine cyathostomins in vitro, Veterinary Parasitology, 2018, 249, 63–69, doi: 10.1016/j.vetpar.2017.11.008
174. [174] F. A. Sinott, Â. Sena-Lopes, K. S. Leal, M. T. de Oliveira Silva, M. C. de Freitas, M. Q. de Moura, M. E. A. Berne, S. Borsuk, Essential oil from Brazilian Red Propolis exhibits anthelmintic activity against larvae of Toxocara cati, Experimental Parasitology, 2019, 200, 37–41, doi: 10.1016/j.exppara.2019.03.014
175. [175] M. P. Silva, T. M. Silva, A. C. Mengarda, M. C. Salvadori, F. S. Teixeira, S. M. Alencar, G. C. Luz Filho, B. Bueno-Silva, J. de Moraes, Brazilian red propolis exhibits antiparasitic properties in vitro and reduces worm burden and egg production in an mouse model harboring either early or chronic Schistosoma mansoni infection, Journal of Ethnopharmacology, 2021, 264, 113387, doi: 10.1016/j.jep.2020.113387
176. [176] M. Patel, A. Ganeshpurkar, D. Bansal, N. Dubey, Experimental evaluation of anthelmintic effect of Gallic Acid, Pharmacognosy Communications, 2015, 5, 145–147, doi: 10.5530/pc.2015.2.6
177. [177] C. García-Hernández, R. Rojo-Rubio, A. Olmedo-Juárez, A. Zamilpa, P. M. de Gives, I. A. Antonio-Romo, L. Aguilar-Marcelino, J. Arece-García, D. Tapia-Maruri, M. González-Cortazar, Galloyl derivatives from Caesalpinia coriaria exhibit in vitro ovicidal activity against cattle gastrointestinal parasitic nematodes, Experimental Parasitology, 2019, 200, 16–23, doi: 10.1016/j.exppara.2019.03.012
178. [178] C. Kamaraj, A. A. Rahuman, A. Bagavan, G. Elango, G. Rajakumar, A. A. Zahir, S. Marimuthu, T. Santhoshkumar, C. Jayaseelan, Evaluation of medicinal plant extracts against blood-sucking parasites, Parasitology Research, 2010, 106, 1403–1412, doi: 10.1007/s00436-010-1816-z
179. [179] T. Banerjee, A. Singh, S. Kumar, T. Dhanani, N. A. Gajbhiye, T. K. Koley, A. Maurya, J. Filgona, Ovicidal and larvicidal effects of extracts from leaves of Andrographis paniculata (Burm. f.) Wall. ex Nees against field isolates of human hookworm (Ancylostoma duodenale), Journal of Ethnopharmacology, 2019, 235, 489–500, doi: 10.1016/j.jep.2019.02.021
180. [180] Q. AlGabbani, Nanotechnology: A promising strategy for the control of parasitic infections, Experimental Parasitology, 2023, 250, 108548, doi: 10.1016/j.exppara.2023.108548
181. [181] O. Dagdag, R. Haldhar, T. W. Quadri, W. Daoudi, E. Berdimurodov, H. Kim, Nanomaterials and their properties, In Nano-Hybrid Smart Coatings: Advancements in Industrial Efficiency and Corrosion Resistance, American Chemical Society, 2024, 17-40, doi: 10.1021/bk-2024-1469.ch002
182. [182] S. Sharma, V. Goel, P. Kaur, K. Gadhave, N. Garg, L. D. Singla, D. Choudhury, Targeted drug delivery using beeswax-derived albendazole-loaded solid lipid nanoparticles in Haemonchus contortus, an albendazole-tolerant nematode, Experimental Parasitology, 2023, 253, 108593, doi: 10.1016/j.exppara.2023.108593
183. [183] C. M. Ferraz, L. C. Comério, V. B. S. Segantine, J. P. B. de Assis, L. P. Costa Silva, L. D. N. R. Bezerra, J. V. de Araújo, V. L. R. Vilela, F. E. de Freitas Soares, G. A. M. Rossi, F. L. Tobias, H. Langoni, F. R. Braga, Silver nanoparticles from Duddingtonia flagrans: Evaluation of potential ovicidal activity on Toxocara canis eggs, Pathogens, 2024, 13, 1043, doi: 10.3390/pathogens13121043
184. [184] C. A. Pimentel-Acosta, F. N. Morales-Serna, M. C. Chávez-Sánchez, H. H. Lara, A. Pestryakov, N. Bogdanchikova, E. J. Fajer-Ávila, Efficacy of silver nanoparticles against the adults and eggs of monogenean parasites of fish, Parasitology Research, 2019, 118, 1741–1749, doi: 10.1007/s00436-019-06315-9
185. [185] S. Hande, V. Sonkar, P. Bhoj, N. Togre, K. Goswami, D. Dash, The role of oxidative and nitrosative stress of silver nanoparticles in human parasitic helminth Brugia malayi: A mechanistic insight, Acta Parasitologica, 2021, 66, 1212–1221, doi: 10.1007/s11686-021-00394-4
186. [186] A. Rehman, R. Ullah, I. Uddin, I. Zia, L. Rehman, S. M. A. Abidi, In vitro anthelmintic effect of biologically synthesized silver nanoparticles on liver amphistome, Gigantocotyle explanatum, Experimental Parasitology, 2019, 198, 95–104, doi: 10.1016/j.exppara.2019.02.005
187. [187] M. T. Rahimi, E. Ahmadpour, B. R. Esboei, A. Spotin, M. H. K. Koshki, A. Alizadeh, S. Honary, H. Barabadi, M. A. Mohammadi, Scolicidal activity of biosynthesized silver nanoparticles against Echinococcus granulosus protoscolices, International Journal of Surgery, 2015, 19, 128–133, doi: 10.1016/j.ijsu.2015.05.043
188. [188] Y. Cheng, X. Chen, W. Song, Z. Kong, P. Li, Y. Liu, Contribution of silver ions to the inhibition of infectivity of Schistosoma japonicum cercariae caused by silver nanoparticles, Parasitology, 2013, 140, 617–625, doi: 10.1017/S0031182012002211
189. [189] C. S. Nwagwu, A. A. Ugwu, C. P. Agbo, A. L. Onugwu, S. O. Onugwu, S. W. Uzondu, O. B. Onokala, S. C. Emencheta, A. A. Attama, P. O. Nnamani, Surface-modified nanocarriers in the design of targeted drug delivery systems for parasitic diseases: a comprehensive review, BMC Chemistry, 2026, 20, 95, doi: 10.1186/s13065-026-01772-7
190. [190] B. Y. S. Kim, J. T. Rutka, W. C. W. Chan, Nanomedicine, New England Journal of Medicine, 2010, 363, 2434–2443, doi: 10.1056/NEJMra0912273
191. [191] M. M. O. Rashid, J. Ferdous, S. Banik, M. R. Islam, A. H. M. M. Uddin, F. N. Robel, Anthelmintic activity of silver-extract nanoparticles synthesized from the combination of silver nanoparticles and M. charantia fruit extract, BMC Complementary and Alternative Medicine, 2016, 16, 242, doi: 10.1186/s12906-016-1219-5
192. [192] A. Shelar, J. Sangshetti, S. Chakraborti, A. V. Singh, R. Patil, S. Gosavi, Helminthicidal and larvicidal potentials of biogenic silver nanoparticles synthesized from medicinal plant Momordica charantia, Medicinal Chemistry, 2019, 15, 781–789, doi: 10.2174/1573406415666190430142637
193. [193] V. Goel, P. Kaur, L. D. Singla, D. Choudhury, Biomedical evaluation of Lansium parasiticum extract-protected silver nanoparticles against Haemonchus contortus, a parasitic worm, Frontiers in Molecular Biosciences, 2020, 7, 595646, doi: 10.3389/fmolb.2020.595646
194. [194] C. M. Ferraz, L. C. Comério, V. B. S. Segantine, J. P. B. de Assis, L. P. Costa Silva, L. D. N. R. Bezerra, J. V. de Araújo, V. L. R. Vilela, F. E. de Freitas Soares, G. A. M. Rossi, F. L. Tobias, H. Langoni, F. R. Braga, Silver nanoparticles from Duddingtonia flagrans: Evaluation of potential ovicidal activity on Toxocara canis eggs, Pathogens, 2024, 13, 1043, doi: 10.3390/pathogens13121043
195. [195] C. A. Pimentel-Acosta, F. N. Morales-Serna, M. C. Chávez-Sánchez, H. H. Lara, A. Pestryakov, N. Bogdanchikova, E. J. Fajer-Ávila, Efficacy of silver nanoparticles against the adults and eggs of monogenean parasites of fish, Parasitology Research, 2019, 118, 1741–1749, doi: 10.1007/s00436-019-06315-9
196. [196] S. Hande, V. Sonkar, P. Bhoj, N. Togre, K. Goswami, D. Dash, The role of oxidative and nitrosative stress of silver nanoparticles in human parasitic helminth Brugia malayi: a mechanistic insight, Acta Parasitologica, 2021, 66, 1212–1221, doi: 10.1007/s11686-021-00394-4
197. [197] A. Rehman, R. Ullah, I. Uddin, I. Zia, L. Rehman, S. M. A. Abidi, In vitro anthelmintic effect of biologically synthesized silver nanoparticles on liver amphistome, Gigantocotyle explanatum, Experimental Parasitology, 2019, 198, 95–104, doi: 10.1016/j.exppara.2019.02.005
198. [198] M. T. Rahimi, E. Ahmadpour, B. R. Esboei, A. Spotin, M. H. K. Koshki, A. Alizadeh, S. Honary, H. Barabadi, M. A. Mohammadi, Scolicidal activity of biosynthesized silver nanoparticles against Echinococcus granulosus protoscolices, International Journal of Surgery, 2015, 19, 128–133, doi: 10.1016/j.ijsu.2015.05.043
199. [199] Y. Cheng, X. Chen, W. Song, Z. Kong, P. Li, Y. Liu, Contribution of silver ions to the inhibition of infectivity of Schistosoma japonicum cercariae caused by silver nanoparticles, Parasitology, 2013, 140, 617–625, doi: 10.1017/S0031182012002211
200. [200] K. Morsy, S. Fahmy, A. Mohamed, S. Ali, M. El-Garhy, M. Shazly, Optimizing and evaluating the antihelminthic activity of the biocompatible zinc oxide nanoparticles against the ascaridid nematode, Parascaris equorum in vitro, Acta Parasitologica, 2019, 64, 873–886, doi: 10.2478/s11686-019-00111-2
201. [201] Y. A. Khan, B. R. Singh, R. Ullah, M. Shoeb, A. H. Naqvi, S. M. A. Abidi, Anthelmintic effect of biocompatible zinc oxide nanoparticles (ZnO NPs) on Gigantocotyle explanatum, a neglected parasite of Indian water buffalo, PLOS ONE, 2015, 10, e0133086, doi: 10.1371/journal.pone.0133086
202. [202] Z. Baghbani, B. Esmaeilnejad, S. Asri-Rezaei, Assessment of oxidative/nitrosative stress biomarkers and DNA damage in Teladorsagia circumcincta following exposure to zinc oxide nanoparticles, Journal of Helminthology, 2020, 94, e115, doi: 10.1017/S0022149X19001068
203. [203] M. Shakir, M. Faraz, M. S. Khan, S. I. Al-Resayes, The photocatalytic, in vitro anthelmintic activity of biomolecule-inspired CDS nanoparticles, Comptes Rendus. Chimie, 2015, 18, 966–978, doi: 10.1016/j.crci.2015.07.009
204. [204] P. K. Kar, S. Murmu, S. Saha, V. Tandon, K. Acharya, Anthelmintic efficacy of gold nanoparticles derived from a phytopathogenic fungus, Nigrospora oryzae, PLOS ONE, 2014, 9, e84693, doi: 10.1371/journal.pone.0084693
205. [205] G. Benelli, Gold nanoparticles – against parasites and insect vectors, Acta Tropica, 2018, 178, 73–80, doi: 10.1016/j.actatropica.2017.10.021
206. [206] M. Sonane, N. Moin, A. Satish, The role of antioxidants in attenuation of Caenorhabditis elegans lethality on exposure to TiO₂ and ZnO nanoparticles, Chemosphere, 2017, 187, 240–247, doi: 10.1016/j.chemosphere.2017.08.080
207. [207] M. F. Charão, C. Souto, N. Brucker, A. Barth, D. S. Jornada, D. Fagundez, D. S. Ávila, V. L. Eifler-Lima, S. S. Guterres, A. R. Pohlmann, S. C. Garcia, Caenorhabditis elegans as an alternative in vivo model to determine oral uptake, nanotoxicity, and efficacy of melatonin-loaded lipid-core nanocapsules on paraquat damage, International Journal of Nanomedicine, 2015, 10, 5093–5106, doi: 10.2147/IJN.S84909
208. [208] A. A. Bauomy, Zinc oxide nanoparticles and L-carnitine effects on neuro-schistosomiasis mansoni induced in mice, Environmental Science and Pollution Research, 2020, 27, 18699–18707, doi: 10.1007/s11356-020-08356-5
209. [209] A. Tauseef, Hisamuddin, A. Khalilullah, I. Uddin, Role of MgO nanoparticles in the suppression of Meloidogyne incognita, infecting cowpea and improvement in plant growth and physiology, Experimental Parasitology, 2021, 220, 108045, doi: 10.1016/j.exppara.2020.108045
210. [210] N. Torabi, F. Dobakhti, A. Haniloo, Albendazole and praziquantel chitosan nanoparticles: preparation, characterization, and in vitro release study, Iranian Journal of Science and Technology, Transactions A: Science, 2018, 42, 1269–1275, doi: 10.1007/s40995-017-0150-z
211. [211] M. A. Moustafa, H. S. Mossalem, R. M. Sarhan, A. A. Abdel-Rahman, E. M. Hassan, The potential effects of silver and gold nanoparticles as molluscicides and cercaricides on Schistosoma mansoni, Parasitology Research, 2018, 117, 3867–3880, doi: 10.1007/s00436-018-6093-2
212. [212] A. Piechulek, L. C. Berwanger, A. von Mikecz, Silica nanoparticles disrupt OPT-2/PEP-2-dependent trafficking of nutrient peptides in the intestinal epithelium, Nanotoxicology, 2019, 13, 1133–1148, doi: 10.1080/17435390.2019.1643048
213. [213] R. V. Pena, R. Silva Brito, O. A. Araújo, L. Damacena-Silva, C. A. Y. Harayashiki, T. L. Rocha, Hazardous effects of nickel ferrite nanoparticles and nickel chloride in early life stages of the freshwater snail Biomphalaria glabrata (Say, 1818), Environmental Science and Pollution Research, 2024, 31, 58324–58334, doi: 10.1007/s11356-024-35011-0
214. [214] A. Kazan, Ö. Yeşil-Çeliktas, Y. S. Zhang, Fabrication of thymoquinone-loaded albumin nanoparticles by microfluidic particle synthesis and their effect on planarian regeneration, Macromolecular Bioscience, 2019, 19, e1900182, doi: 10.1002/mabi.201900182
215. [215] S. Sathya, B. Shanmuganathan, B. Balasubramaniam, K. Balamurugan, K. P. Devi, Phytol loaded PLGA nanoparticles regulate the expression of Alzheimer's related genes and neuronal apoptosis against amyloid-β induced toxicity in Neuro-2a cells and transgenic Caenorhabditis elegans, Food and Chemical Toxicology, 2020, 136, 110962, doi: 10.1016/j.fct.2019.110962
216. [216] L. L. Li, Y. H. Cui, L. Y. Lu, Y. L. Liu, C. J. Zhu, L. J. Tian, W. W. Li, X. Zhang, H. Cheng, J. Y. Ma, J. Chu, Z. H. Tong, H. Q. Yu, Selenium stimulates cadmium detoxification in Caenorhabditis elegans through thiols-mediated nanoparticles formation and secretion, Environmental Science & Technology, 2019, 53, 2344–2352, doi: 10.1021/acs.est.8b04200
217. [217] M. J. Mashock, T. Zanon, A. D. Kappell, L. N. Petrella, E. C. Andersen, K. R. Hristova, Copper oxide nanoparticles impact several toxicological endpoints and cause neurodegeneration in Caenorhabditis elegans, PLOS ONE, 2016, 11, e0167613, doi: 10.1371/journal.pone.0167613
218. [218] R. Dorostkar, M. Ghalavand, A. Nazarizadeh, M. Tat, M. S. Hashemzadeh, Anthelmintic effects of zinc oxide and iron oxide nanoparticles against Toxocara vitulorum, International Nano Letters, 2017, 7, 157–164, doi: 10.1007/s40089-016-0198-3
219. [219] S. Ramya, T. Shanmugasundaram, R. Balagurunathan, Actinobacterial enzyme mediated synthesis of selenium nanoparticles for antibacterial, mosquito larvicidal and anthelminthic applications, Particulate Science and Technology, 2020, 38, 63–72, doi: 10.1080/02726351.2018.1508098
220. [220] L. Gonzalez-Moragas, S. M. Yu, N. Benseny-Cases, S. Stürzenbaum, A. Roig, A. Laromaine, Toxicogenomics of iron oxide nanoparticles in the nematode C. elegans, Nanotoxicology, 2017, 11, 647–657, doi: 10.1080/17435390.2017.1342011
221. [221] S. Sharma, V. Goel, P. Kaur, K. Gadhave, N. Garg, L. D. Singla, D. Choudhury, Bioinspired dual-functional solid lipid nanoformulations for targeted drug delivery and sustained release for enhancement of potency of albendazole, an antihelminthic drug, bioRxiv, 2021, 2021-2107, doi: 10.1101/2021.07.24.453620
222. [222] F. K. Alves de Souza, M. L. F. Della Costa, M. A. Morales, A. G. Magdalena, Enhancement of curcumin bioavailability using a controlled drug delivery system based on chitosan and green iron oxide nanoparticles (Fe₃O₄), Química Nova, 2025, 48, e-20250202, doi: 10.21577/0100-4042.20250202
223. [223] C. Jayaseelan, A. A. Rahuman, G. Rajakumar, T. Santhoshkumar, A. V. Kirthi, S. Marimuthu, A. Bagavan, C. Kamaraj, A. A. Zahir, G. Elango, K. Velayutham, K. V. B. K. Rao, L. Karthik, S. Raveendran, Efficacy of plant-mediated synthesized silver nanoparticles against hematophagous parasites, Parasitology Research, 2012, 111, 921–933, doi: 10.1007/s00436-011-2473-6
224. [224] M. Rai, A. P. Ingle, R. Pandit, P. Paralikar, N. Anasane, C. A. D. Santos, Curcumin and curcumin-loaded nanoparticles: antipathogenic and antiparasitic activities, Expert Review of Anti-Infective Therapy, 2020, 18, 367–379, doi: 10.1080/14787210.2020.1730815
225. [225] A. E. Al-Snafi, Antiparasitic effects of medicinal plants (part 1)—a review, IOSR Journal of Pharmacy, 2016, 6, 51–66.
226. [226] T. P. Simanjuntak, M. Hatta, A. M. Tahir, R. H. Sirait, M. B. Karo, T. Tambaib, R. Dwiyanti, R. A. Noviyanthi, A. R. Junita, Analysis of anti-toxoplasma immunoglobulin G and immunoglobulin M antibody levels after intervention with Curcuma longa extract on early pregnant mice with acute toxoplasmosis, Journal of Global Infectious Diseases, 2019, 11, 25–29, doi: 10.4103/jgid.jgid_28_18
227. [227] J. R. do Carmo Neto, R. O. Guerra, J. R. Machado, A. C. A. Silva, M. V. da Silva, Antiprotozoal and anthelmintic activity of zinc oxide nanoparticles, Current Medicinal Chemistry, 2022, 29, 2127–2141, doi: 10.2174/0929867328666210709105850
228. [228] A. Arenal Cruz, M. B. Molento, Nanotechnology: meeting the future of Veterinary Parasitology Research, Pesquisa Veterinária Brasileira, 2015, 35, 842–843, doi: 10.1590/S0100-736X2015001000004
229. [229] H. K. Makadia, S. J. Siegel, Poly lactic-co-glycolic acid (PLGA) as biodegradable controlled drug delivery carrier, Polymers, 2011, 3, 1377–1397, doi: 10.3390/polym3031377
230. [230] S. Talegaonkar, A. Bhattacharyya, Potential of lipid nanoparticles (SLNs and NLCs) in enhancing oral bioavailability of drugs with poor intestinal permeability, AAPS PharmSciTech, 2019, 20, 121, doi: 10.1208/s12249-019-1337-8
231. [231] P. Liu, G. Chen, J. Zhang, A review of liposomes as a drug delivery system: current status of approved products, regulatory environments, and future perspectives, Molecules, 2022, 27, 1372, doi: 10.3390/molecules27041372
232. [232] M. Mir, S. Ishtiaq, S. Rabia, M. Khatoon, A. Zeb, G. M. Khan, A. ur Rehman, F. ud Din, Nanotechnology: from in vivo imaging system to controlled drug delivery, Nanoscale Research Letters, 2017, 12, 500, doi: 10.1186/s11671-017-2249-8
233. [233] L. Wang, L. Yan, J. Liu, C. Chen, Y. Zhao, Quantification of nanomaterial/nanomedicine trafficking in vivo, Analytical Chemistry, 2018, 90, 589–614, doi: 10.1021/acs.analchem.7b04765
234. [234] S. Soares, J. Sousa, A. Pais, C. Vitorino, Nanomedicine: principles, properties, and regulatory issues, Frontiers in Chemistry, 2018, 6, 360, doi: 10.3389/fchem.2018.00360
235. [235] F. Xiaoli, Q. Chen, W. Guo, Y. Zhang, H. Chen, J. Wu, L. Shao, Toxicology data of graphene-family nanomaterials: an update, Archives of Toxicology, 2020, 94, 1915–1939, doi: 10.1007/s00204-020-02717-2
236. [236] M. Auffan, J. Rose, M. R. Wiesner, J. Y. Bottero, Chemical stability of metallic nanoparticles: a parameter controlling their potential cellular toxicity in vitro, Environmental Pollution, 2009, 157, 1127–1133, doi: 10.1016/j.envpol.2008.10.002
237. [237] Y. Sun, D. Chen, Y. Pan, W. Qu, H. Hao, X. Wang, Z. Liu, S. Xie, Nanoparticles for antiparasitic drug delivery, Drug Delivery, 2019, 26, 1206–1221, doi: 10.1080/10717544.2019.1692968
238. [238] S. Saraf, A. Jain, A. Tiwari, A. Verma, P. K. Panda, S. K. Jain, Advances in liposomal drug delivery to cancer: an overview, Journal of Drug Delivery Science and Technology, 2020, 56, 101549, doi: 10.1016/j.jddst.2020.101549
239. [239] N. Filipczak, J. Pan, S. S. K. Yalamarty, V. P. Torchilin, Recent advancements in liposome technology, Advanced Drug Delivery Reviews, 2020, 156, 4–22, doi: 10.1016/j.addr.2020.06.022
240. [240] D. Guimarães, A. Cavaco-Paulo, E. Nogueira, Design of liposomes as drug delivery system for therapeutic applications, International Journal of Pharmaceutics, 2021, 601, 120571, doi: 10.1016/j.ijpharm.2021.120571
241. [241] V. Rajendran, S. Rohra, M. Raza, G. M. Hasan, S. Dutt, P. C. Ghosh, Stearylamine liposomal delivery of monensin in combination with free artemisinin eliminates blood stages of Plasmodium falciparum in culture and P. berghei infection in murine malaria, Antimicrobial Agents and Chemotherapy, 2016, 60, 1304–1318, doi: 10.1128/AAC.01796-15
242. [242] A. Ghosh, L. Li, L. Xu, R. P. Dash, N. Gupta, J. Lam, Q. Jin, V. Akshintala, G. Pahapale, W. Liu, A. Sarkar, R. Rais, D. H. Gracias, F. M. Selaru, Gastrointestinal-resident, shape-changing microdevices extend drug release in vivo, Science Advances, 2020, 6, eabb4133, doi: 10.1126/sciadv.abb4133

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