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Introduction

What Do We Understand About Plant-Derived Exosome-Like Nanovesicles?

In recent years, Plant-derived exosome-like nanoparticles (PELNs) have attracted increasing attention as natural nanocarriers for biomedical applications. While mammalian-derived exosomes also demonstrate therapeutic potential, their clinical translation faces several challenges, including the risk of immune rejection, possible transmission of animal-borne pathogens, ethical concerns regarding the use of animal-derived materials, animal welfare considerations, low production yields, and the high cost of establishing large-scale culture systems. In contrast, PELNs are abundant and are typically derived from fruits, vegetables, and medicinal herbs, making them sustainable and readily available.¹ They carry unique bioactive cargos such as plant-specific proteins, lipids, nucleic acids, and microRNAs(miRNAs), offering them with distinct functional properties and broad translational promise.² Moreover, plant materials are easier to obtain, more economical to process, and simpler to store and transport, while avoiding the ethical concerns often associated with animal-derived exosomes in drug development.³

Currently, PELNs are primarily isolated using ultracentrifugation, ultrafiltration centrifugation, and density gradient centrifugation.⁴ However, variations in these protocols often result in substantial differences in yield, purity, and biological activity. Compared with size exclusion chromatography, ultracentrifugation allows the processing of larger sample volumes and achieves higher yields. In contrast to the more economical polymer precipitation method, it results in fewer co-precipitated impurities. Although density gradient centrifugation generally provides higher purity, both ultrafiltration centrifugation and ultracentrifugation are more practical for large-scale preparations, with ultracentrifugation often preferred for its balance of yield and feasibility (Table 1). ^2,4–6 Recent efforts have therefore focused on developing scalable isolation strategies that optimize yield and purity while preserving the functional integrity of PELNs.

Table 1 Comparison of Conventional Separation Techniques for PELNs

In the characterization of PELNs, biochemical profiling serves as a critical step in distinguishing them from other types of extracellular vesicles, particularly functional microvesicles.² Molecular characterization is commonly performed using Western blotting and flow cytometry. Potential markers for PELNs include surface proteins such as CD63,⁷ PEN1,⁸ TET8,^5,8 Exo70,⁵ TET3,⁵ Class I chitinase (PR-3) and Class I β-1,3-glucanase (PR-2).⁹ Internal proteins such as heat shock protein 70 (HSP70), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), and S-adenosylhomocysteine hydrolase (SAHH) are also frequently reported.^5,8 Nucleic acids, particularly small RNAs, can be profiled using next-generation sequencing or qPCR. Lipidomic analysis via mass spectrometry has revealed that PELNs possess lipid bilayers primarily composed of phosphatidylcholine and phosphatidylethanolamine. Notably, the lipid composition varies among PELNs derived from different plant sources.⁷ Phosphatidic acid, in particular, is a dominant lipid species that plays a key role in the uptake and absorption of PELNs by recipient cells⁸ (Figure 1).

Figure 1 Structural and molecular composition of PELNs. This figure illustrates a PELN, typically 50–200 nm in diameter, with a spherical or cup-shaped morphology and a lipid bilayer membrane. Characteristic surface markers include CD63, PEN1, TET8, TET3, Exo70, Class I chitinase (PR-3), and Class I β-1,3-glucanase (PR-2). Internal contents of PELNs consist of cytosolic proteins such as HSP70, GAPDH, nucleic acids (miRNA, RNA), lipids, and other biologically active constituents. The membrane composition mainly includes phosphatidylcholine (PC), phosphatidylethanolamine (PE), and phosphatidic acid (PA), which are critical for vesicle stability and cellular uptake. The figure also includes a symbolic legend indicating the molecular components. (By Figdraw).

Due to their low immunogenicity, excellent biocompatibility, inherent targeting capacity, and suitability for surface engineering, PELNs have been widely explored in drug delivery, diagnostics, and therapeutic intervention.^5,10 They have demonstrated promising effects across diverse disease models, including inflammation, oxidative stress, cancer, wound healing,⁸ immune regulation, neuroprotection, metabolic modulation, cardiovascular protection, gut homeostasis, osteoporosis, muscle atrophy, and premature ovarian failure. Notably, PELNs are compatible with multiple administration routes, including oral, intravenous, intratracheal, intranasal, and topical delivery.¹¹ This review therefore provides a comprehensive overview of therapeutic applications and signaling mechanisms associated with PELNs, offering insights to guide future translational research and clinical development.

The Key Therapeutic Applications of Plant-Derived Exosome-Like Nanoparticles

Among the various biomolecules encapsulated in PELNs, miRNAs are key post-transcriptional regulators of gene expression. Their therapeutic potential lies in their ability to mediate cross-kingdom communication, thereby increasing the diversity of miRNAs in mammalian cells and exerting multi-target effects.^12,13 PELNs can protect miRNAs from degradation in the gastrointestinal tract while maintaining specific concentrations. Studies suggest that PELNs contain hundreds of miRNAs, and a single miRNA can target hundreds of mRNAs. Thus, when PELNs levels reach a certain baseline, they may produce significant regulatory effects.¹⁴

For example, ginseng-derived plant exosomes carry mtr-miR159 and deliver it into bone marrow mesenchymal stem cells. This upregulates Tmem100 and activates the PI3K/Akt signaling pathway, promoting neural differentiation and enhancing peripheral nerve regeneration in a rat model of peripheral nerve injury.¹⁵ In another study, ginseng-derived exosome-like nanoparticles delivered their endogenous vvi-miR396b and ptc-miR396f into glioma cells, silencing the oncogenes c-MYC and BCL2, effectively inhibiting tumor growth and achieving efficient blood-brain barrier penetration.¹⁶

Similar to miRNA, small interfering RNA (siRNA) is a double-stranded RNA molecule approximately 20–25 nucleotides in length. It can bind perfectly to the target mRNA and induce its degradation, leading to specific gene silencing. siRNA has important potential for precision therapy, especially in cancer treatment. A typical example is ginger-derived exosome-like nanoparticles (GELNs), which deliver Bcl2 siRNA into tumor cells. By silencing the anti-apoptotic gene Bcl2, they activate the apoptosis pathway in cancer cells and significantly suppress tumor growth in a mouse breast cancer model.¹⁷

PELNs have recently been shown to influence cell fate, inflammatory responses, oxidative stress, tissue repair, metabolic regulation, and tumor immunity through diverse molecular mechanisms.¹⁸ The following studies reveal their considerable potential in disease treatment.

PELNs can regulate cell proliferation, apoptosis, differentiation, and stemness maintenance through multiple signaling pathways and key molecules, thereby contributing to tissue repair, disease therapy, and regenerative medicine. In terms of cell proliferation, PELNs from Momordica charantia are a typical example. They activate the PI3K/Akt and ERK signaling pathways, upregulate PCNA, Cyclin D1, Cyclin B1, and Ki-67, promote cell cycle progression, and enhance cell proliferation, which improves the repair capacity of cardiomyocytes after radiation injury.¹⁹ Regarding apoptosis, PELNs from Brucea javanica carry natural miRNAs (such as the let-7 family) that inhibit the PI3K/Akt/mTOR pathway while activating ROS/caspase-dependent apoptosis, leading to caspase-3 and PARP cleavage. This suppresses tumor cell survival and induces programmed cell death, highlighting the potential of plant exosomes in antitumor therapy.²⁰ In differentiation, PELNs from Panax ginseng activate the PI3K/Akt pathway through miRNAs, significantly upregulate Nestin, β3-tubulin, NGF, BDNF, and bFGF, and drive bone marrow mesenchymal stem cells to differentiate into neurons, providing new insights into neural repair and the treatment of neurodegenerative diseases.¹⁵ In stemness maintenance, PELNs from Grape inhibit GSK-3β activity, stabilize β-catenin nuclear translocation, and activate the Wnt/β-catenin pathway. This induces transcription factors such as c-Myc, Lgr5⁺, SOX2, Nanog, OCT4, and KLF4, which enhance the self-renewal and regeneration of intestinal stem cells.²¹ These studies indicate that PELNs regulate cell cycle factors, apoptosis pathways, differentiation markers, and stemness-related transcription factors, thus playing important roles in cell fate and opening new directions for tissue repair, antitumor therapy, and regenerative medicine.

PELNs also exhibit strong anti-inflammatory and immunomodulatory activities. PELNs from Panax notoginseng inhibit M1 macrophage polarization and promote M2 polarization, reducing TNF-α and IL-6 while increasing IL-10, thereby alleviating inflammation.²² PELNs from Garlic suppress the TLR4/NF-κB pathway, downregulate inflammatory cytokines such as IL-6 and TNF-α, and enhance the tight junction protein ZO-1 to maintain intestinal barrier integrity.²³ PELNs from Broccoli mainly act through the AMPK pathway, promoting tolerogenic dendritic cells and Tregs to restore immune homeostasis.²⁴ These findings suggest that PELNs regulate immune cell phenotypes and cytokine levels through multiple pathways and hold therapeutic potential for inflammatory diseases.

In addition, PELNs demonstrate antioxidative potential. For example, PELNs from Mung bean sprouts activate the PI3K/Akt-Nrf2 pathway to upregulate HO-1 and SOD, and reduce oxidative stress.²⁵ PELNs from Carrot enhance HO-1 and NQO1 via the Nrf2/ARE pathway and decrease ROS production, improving cellular antioxidant defense.²⁶ PELNs from Ginger, which contain 6-Shogaol, also activate Nrf2 and further enhance the ability to scavenge free radicals.²⁷ These mechanisms show that PELNs can effectively strengthen antioxidant capacity, providing new approaches for preventing tissue injury and delaying aging.

In metabolic regulation, PELNs display broad effects. PELNs from Garlic upregulate GLP-1 and IRS1/2, enhance insulin signaling, and improve glucose utilization.²⁸ PELNs from Mung bean sprouts increase GLUT4 expression and decrease GSK-3β activity, promoting glucose uptake and glycogen synthesis and improving insulin resistance.²⁵ PELNs from Citrus limon suppress lipid metabolism genes such as ACACA, DDHD1, and DHCR24, reduce lipid synthesis, and induce tumor cell apoptosis.²⁹ These results indicate that PELNs can improve glucose metabolism, enhance insulin sensitivity, and regulate lipid metabolism, showing promise in the treatment of metabolic diseases.

In immune regulation and antitumor responses, PELNs demonstrate unique mechanisms. Exosomes from Artemisia annua contain plant mitochondrial DNA, which activates the cGAS-STING pathway, enhances IFN-I production, and promotes CD8⁺ T cell activation, thereby improving antitumor immunity.³⁰ PELNs from Catharanthus roseus act through the TNF-α/NF-κB/PU.1 axis to strengthen immune cell function and relieve chemotherapy-induced immunosuppression.³¹ PELNs from Panax ginseng activate TLR4 signaling, drive tumor-associated macrophages toward the M1 phenotype, and enhance local immune activity.³² These studies indicate that PELNs can regulate both innate and adaptive immunity, enhance antitumor responses, and provide new strategies for cancer immunotherapy.

PELNs are currently under clinical investigation for a variety of human diseases. Ongoing clinical trials are evaluating their therapeutic efficacy in the treatment of colorectal cancer (NCT01294072), head and neck cancer (NCT01668849), and IBD treatment (NCT04879810). Table 2 and Table 3 summarize the classification of PELN-based therapies according to disease types and the therapeutic drugs delivered by PELNs, respectively.³³

Table 2 Classification of PELNs Therapies by Disease Types

Table 3 Therapeutic Drugs Delivered by PELNs

Effects of Plant-Derived Exosome-Like Nanoparticles on Disease-Associated Signaling Pathways

PELNs exert therapeutic effects by modulating multiple critical signaling pathways, including PI3K/Akt, NF-κB, Wnt, AMPK, MAPK, the NLRP3 inflammasome, cGAS/STING, and Nrf2/ARE. Through these regulatory axes, PELNs influence key biological processes such as metabolic homeostasis, anti-inflammation, antioxidation, wound healing, neuroprotection, and tumor suppression, thereby offering therapeutic promise in diverse pathological conditions, including diabetes, neurodegenerative diseases, cardiovascular disorders, inflammatory diseases, and cancer.

PELNs activate PI3K/Akt pathway to promote cell survival, proliferation, and metabolic regulation. In metabolic diseases such as diabetes, PELNs enhance Glucose Transporter Type 4 (GLUT4) expression via the PI3K/Akt pathway, thereby improving insulin resistance.²⁵ In neuroprotection (eg, neurodegeneration, ischemic stroke, and ischemia-reperfusion injury), they inhibit apoptosis and maintain blood-brain barrier (BBB) integrity.^22,77,78 This pathway also facilitates wound healing by promoting skin cell proliferation, migration, extracellular matrix secretion, and angiogenesis.⁶⁷ In tumors, PELNs modulate cancer cell survival, proliferation, and metabolism while inhibiting invasion and metastasis.^20,58,59

NF-κB pathway is primarily involved in regulating inflammation, immune responses, and cell survival. In inflammatory diseases (eg, colitis), PELNs downregulate pro-inflammatory cytokines such as TNF-α and IL-6, alleviating inflammation.²³ In bone metabolism disorders (eg, osteoporosis), they inhibit osteoclast activation to reduce bone loss.⁸³ They also enhance immune function (eg, post-chemotherapy immunomodulation) by activating lymphocytes and macrophages,³¹ and contribute to anti-aging effects in skin by promoting collagen expression.⁷²

Wnt signaling pathway regulates cell proliferation, differentiation, and tissue homeostasis. PELNs promote intestinal stem cell proliferation and differentiation via Wnt/TCF4 activation, thus supporting intestinal repair.¹⁰⁴ In inflammatory conditions like colitis, they modulate neural stem cell differentiation in the intestine to enhance regenerative capacity.²¹

AMPK pathway is a master regulator of cellular energy metabolism and homeostasis. In muscle atrophy, PELNs upregulate myogenesis-related factors, enhance metabolic activity, and improve mitochondrial function.⁸⁵ In inflammation (eg, colitis), they attenuate inflammation by suppressing pro-inflammatory cytokines and promoting anti-inflammatory mediators.²⁴

MAPK pathway governs cell proliferation, differentiation, stress responses, apoptosis, and inflammation. In inflammatory liver injury (eg, acetaminophen (APAP)-induced hepatotoxicity), PELNs inhibit phosphorylation of key proteins, reducing hepatocyte apoptosis and inflammation.⁹⁶ In neuroprotection and cardioprotection, they improve cell survival and prevent radiation-induced apoptosis.^19,77 Additionally, they promote tissue regeneration (eg, wound healing), bone remodeling (eg, osteoporosis),^67,82–84 and exert antitumor effects by suppressing proliferation, inducing apoptosis, and reducing tumor cell invasiveness.^29,58

NLRP3 inflammasome regulates innate immunity and the release of pro-inflammatory cytokines. In conditions such as hepatic injury, sepsis-induced acute lung injury, and ulcerative colitis, PELNs inhibit NLRP3 inflammasome assembly, reduce cytokine release, and mitigate inflammatory damage.^{35,42,44,50,105}

cGAS/STING pathway plays a crucial role in innate immunity, antiviral defense, and antitumor immunity. In metabolic disorders such as insulin resistance and type 2 diabetes, PELNs improve metabolic function by reducing inflammation and promoting insulin receptor substrate expression.^28,76 In tumor immunoregulation, they reshape macrophage phenotypes and enhance antitumor immunity, thereby suppressing tumor growth.³⁰

Nrf2/ARE pathway alleviates oxidative stress by upregulating antioxidant defenses. In neurodegenerative diseases (eg, Parkinson’s disease), PELNs reduce oxidative damage and enhance neuronal survival.²⁶ In cardiovascular diseases (eg, myocardial infarction) and inflammatory liver injury (eg, alcoholic fatty liver), they suppress ROS production and induce antioxidant enzymes such as HO-1 and NQO1, thereby reducing tissue injury.^{26,27,79,81,106}

Collectively, PELNs modulate diverse signaling pathways to regulate metabolism, inflammation, oxidative stress, tissue regeneration, and tumor progression, highlighting their broad clinical potential across multiple disease spectrums. (Figure 2).

Table 4 Summary of Signaling Pathways Affected by PELNs

Figure 2 Hierarchical representation of PELNs-regulated signaling pathways and associated diseases. This three-tier circular diagram illustrates the relationship between key signaling pathways modulated by PELNs, the major disease categories they influence, and specific pathological conditions. This three-tier circular figure illustrates the relationship between key signaling pathways modulated by plant-derived exosome-like nanoparticles (PELNs), the major disease categories they influence, and specific pathological conditions. Inner ring (core): Key signaling pathways involved in PELNs-mediated therapeutic effects, including PI3K/Akt, NF-κB, MAPK, AMPK, Wnt, NLRP3 inflammasome, cGAS/STING, Nrf2/ARE, among others. Middle ring: Broad disease categories influenced by the corresponding pathways, such as metabolic disorders, inflammatory diseases, neurodegenerative conditions, cardiovascular diseases, musculoskeletal disorders, cancers, and tissue regeneration. Outer ring: Representative diseases within each category (eg, diabetes, ulcerative colitis, ischemic stroke, myocardial infarction, osteoporosis, TNBC, etc.). The figure highlights the multifunctional regulatory roles of PELNs across diverse pathological contexts. (By Figdraw).

In addition to the major signaling pathways discussed above, several less common but biologically relevant pathways modulated by PELNs have also been identified and are comprehensively summarized in Table 4. With the continued elucidation of the molecular mechanisms by which PELNs regulate intracellular signaling, their application in precision drug delivery is expected to expand significantly. Owing to their low cytotoxicity, high biocompatibility, and minimal intrinsic immunogenicity, PELNs offer a unique therapeutic modality that integrates drug delivery, signaling modulation, and dynamic response to pathological stimuli. This multifaceted functionality enables a synergistic “delivery–regulation–therapy” strategy to enhance therapeutic efficacy. Furthermore, the inherent targeting capacity of PELNs, coupled with their surface modifiability, holds great promise for the development of intelligent drug delivery systems. Such systems would possess disease-site recognition, stimulus responsiveness, and controlled release capabilities, offering a more efficient, safe, and personalized treatment approach for chronic disorders, cancer, and inflammatory diseases.

Beyond signaling pathways, several bioactive molecules have been identified as mediators of PELN function. For example, Houttuynia cordata PELNs contain flavonoids such as luteolin,³⁵ ginger-derived PELNs carry 6-shogaol,²⁷ broccoli-derived PELNs deliver sulforaphane,²⁴ and Artemisia annua PELNs contain mitochondrial DNA,³⁰ engineered ginseng-derived ELNs are loaded with miR-182-5p,⁵⁰ tomato-derived PELNs carry miR164a/b-5p,⁸¹ ginseng-derived nanoparticles contain various miRNAs,¹⁵ Momordica charantia PELNs include miR-5266 and miR-5813,⁷⁸ and Brucea javanica PELNs contain functional miRNAs such as let-7.²⁰ These findings show more clearly which bioactive components of PELNs are responsible for their therapeutic effects.

Future Prospective

With the progress of research on PELNs in drug delivery and disease treatment, their potential as novel therapeutic tools is becoming increasingly evident. Current preclinical studies have demonstrated that PELNs possess low immunogenicity, favorable biocompatibility, and can be given through multiple administration routes. They have shown positive therapeutic effects in models of inflammation, cancer, cardiovascular disease, neurodegeneration, and metabolic disorders. Importantly, PELNs can not only serve as carriers for drugs and nucleic acids (eg, miRNA and siRNA), but also provide therapeutic benefits through their own bioactive components. This dual role makes them promising for complex diseases.

In the future, several challenges still need to be addressed before PELNs can be widely used in clinical practice. First, their therapeutic effects must be reproducible and stable At present, PELNs derived from different plants may show differences in composition and function, and these differences need systematic study. Second, although animal experiments have demonstrated that PELNs can suppress inflammation, reduce oxidative stress, and promote tissue repair, their safety, effective dosage, and long-term benefits in humans remain to be clarified. Furthermore, combining PELNs with existing therapies could further improve outcomes. For example, in cancer therapy, PELNs may serve as natural carriers for nucleic acids and be used in combination with chemotherapy or immune checkpoint inhibitors, which might increase efficacy and reduce side effects.

Looking ahead, PELNs show broad prospects in treatment. They may become not only the next generation of drug delivery platforms, but also independent therapeutic agents. With further progress in preparation methods, mechanistic studies, and clinical validation, PELNs may bring new breakthroughs for the treatment of currently intractable diseases.

Funding

National Natural Science Foundation of China (82405273), Natural Science Foundation of Hubei Province (2022CFD023 and 2024AFD299). Basic scientific research project of the Educational Department of Liaoning Province (JYTMS20230584). Natural Science Foundation of Liaoning Province (2023-MSLH-028).

Disclosure

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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