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Wound healing and signaling pathways

  • Zhe Liu und Yudong Fang EMAIL logo
Veröffentlicht/Copyright: 1. September 2025
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Open Life Sciences
Aus der Zeitschrift Open Life Sciences Band 20 Heft 1

Abstract

Wound healing is a precisely regulated dynamic process in which signaling pathways play a central role. This article provides a comprehensive review of the signaling pathways involved in wound healing, emphasizing their roles in inflammation, vascular regeneration, cell proliferation, and extracellular matrix remodeling. We further discuss the crosstalk between these pathways and their contributions to wound healing dysregulation. Finally, we explore emerging therapeutic strategies targeting these pathways, including small-molecule inhibitors, gene therapy, and biologics, summarizing their preclinical and clinical efficacy. By elucidating the molecular mechanisms underlying wound healing and potential interventions, this review aims to provide valuable insights for future research and translational applications in wound healing.

Keywords: wound healing; signaling pathway; crosstalk; therapeutic strategies

1 Introduction

Wound healing is a dynamic and multifaceted biological process that requires the precise coordination of inflammation, angiogenesis, cell proliferation, and extracellular matrix remodeling. While acute wounds progress through a well-defined sequence of hemostasis, inflammation, proliferation, and remodeling, chronic refractory wounds remain stalled in one or more of these phases, leading to prolonged tissue damage and increased susceptibility to infection. The condition of wound healing is usually also associated with diabetes, ischemia, venous insufficiency, and pressure ulcers. These factors can lead to difficulties in wound healing, significantly reduce the quality of life of patients, and impose a heavy burden on healthcare [1].

The persistent failure of wound healing is largely driven by dysregulated signaling networks that govern cellular responses to injury. Key pathways such as mitogen-activated protein kinase (MAPK), phosphatidylinositide 3-kinases (PI3K)/AKT, TGF-β, NF-κB, Wnt/β-catenin, Hippo-Yes-associated protein (YAP), and Notch are involved in regulating immune responses, vascular remodeling, fibroblast activation, and extracellular matrix turnover. Disruptions in these pathways contribute to excessive inflammation, inadequate angiogenesis, fibroblast dysfunction, and pathological fibrosis, ultimately leading to wound chronicity.

In recent years, increasing efforts have been directed toward identifying molecular targets within these pathways to develop novel therapeutic strategies. Emerging approaches, including small-molecule inhibitors, gene therapy, and biologics, hold promise for modulating key signaling pathways and restoring normal wound healing processes. However, challenges remain in translating these therapies into clinical practice due to issues related to delivery efficiency, off-target effects, and long-term safety.

This review aims to provide a comprehensive analysis of the key signaling pathways in wound healing, their pathological implications, and their interconnections. Furthermore, we discuss current and emerging therapeutic strategies targeting these pathways, summarizing their preclinical and clinical evidence. By elucidating these mechanisms, we hope to contribute to the development of more effective treatments for wound healing.

2 Overview of mechanisms involved in wound healing

2.1 Inflammatory response

Inflammation is an essential initial phase of wound healing, playing a pivotal role in eliminating pathogens and orchestrating tissue repair. Upon injury, damaged tissues release damage-associated molecular patterns and pro-inflammatory cytokines such as tumor necrosis factor-alpha (TNF-α), interleukin (IL)-1 beta, and IL-6, which recruit immune cells to the wound site. Neutrophils are the first responders, producing reactive oxygen species and antimicrobial peptides to clear invading pathogens. This is followed by monocyte infiltration and their differentiation into macrophages, which transition from an inflammatory (M1) to a reparative (M2) phenotype, facilitating tissue regeneration. However, in wound healing, persistent activation of the inflammatory response leads to excessive cytokine secretion, sustained neutrophil infiltration, and macrophage dysfunction, ultimately impairing wound healing and promoting tissue damage.

2.2 Angiogenesis

Angiogenesis, the formation of new blood vessels from pre-existing vasculature, is critical for delivering oxygen and nutrients to regenerating tissues. This process is tightly regulated by pro-angiogenic factors such as vascular endothelial growth factor (VEGF), fibroblast growth factors, and angiopoietins, while anti-angiogenic factors such as thrombospondins and endostatin counterbalance their effects. Under normal conditions, angiogenesis is activated in response to hypoxia and tissue injury, promoting capillary sprouting and endothelial cell (EC) migration. However, in wounds healing, dysregulated angiogenic signaling – often due to prolonged inflammation, endothelial dysfunction, or metabolic disturbances – leads to either insufficient or aberrant vascularization, resulting in tissue hypoxia and impaired healing.

2.3 Cell proliferation and migration

The proliferation and migration of keratinocytes are crucial for wound healing and repair. Epidermal stem cells, located in the basal layer of skin, are specific stem cells of skin tissue with infinite proliferation and differentiation potential. As the differentiation source of epidermal cells of all layers in the process of wound epithelialization, the proliferation and differentiation of epidermal stem cells contribute to the renewal of epidermal cells [2]. The interaction between cells and the extracellular matrix plays a crucial role in influencing stem cell differentiation through various cytokines that convey information.

2.4 Extracellular matrix remodeling

Extracellular matrix is a complex network composed of multiple macromolecules around cells, which can provide a suitable microenvironment for cells, mainly containing collagen, fibrin, elastin, laminin and other substances. As the main component of the extracellular matrix, collagen is distributed in various organs and tissues. Extracellular matrix remodeling is particularly important in the later stage of wound healing, and collagen synthesis and degradation are important factors affecting wound healing. Studies have shown that improving the content and stability of collagen can accelerate wound healing [3].

3 Overview of signaling pathways

3.1 MAPK signaling pathway

The MAPK pathway involves three-stage signaling processes: MAPK, MAPK kinase (MEK or MKK), and the kinase of MAPK kinase (MEKK or MKKK) (Figure 1). These three kinases can be activated sequentially and jointly regulate colossal amounts of important physiological/pathological effects such as cell growth and differentiation and stress and inflammatory responses. The MAPK pathway covers four main branching routes: ERK, JNK, P38/MAPK, and ERK5. The MAPK signaling pathway can induce cell migration and proliferation, so as to accelerate wound healing.

Figure 1 Schematic diagram showing the biological and pathological processes and signaling pathways underlying the wound healing.
Figure 1

Schematic diagram showing the biological and pathological processes and signaling pathways underlying the wound healing.

3.1.1 The MAPK signaling pathway can affect wound healing by affecting cell proliferation and repair

p38 is the most important member of the MAPK family in regulating inflammatory response, and the p38 signaling pathway is closely related to inflammatory response and apoptosis. Studies have shown that the p38/MAPK2 signaling pathway can regulate the expression of inflammatory factors and participate in wound healing, therefore plays a key role in the process of wound repair [4,5,6,7]. JNK is a stress-activated protein kinase in the cytoplasm. As a key molecule in the signal transduction process of various stressors, JNK plays an important role in osmotic pressure changes and inflammatory responses of cells [8]. The JNK signaling pathway is an important member of the MAPK inflammatory pathway. Studies have shown that JNK inhibitors can reduce the expression of related inflammatory factors in infected tissues, thereby alleviating the inflammatory response [9]. ERK signaling pathway is a classic MAPK signal transduction pathway, which not only participates in the regulation of cell proliferation and differentiation but also participates in the regulation of cell morphology and the redistribution of skeleton through phosphorylation of cytoskeletal components in the cytoplasm.

3.1.2 The MAPK pathways can affect wound healing by affecting vascular regeneration

Hou et al. [10] found that the microvascular density of wound was significantly increased in rats with open skin wounds treated with vaccinin, and the lymph node proteins and receptors related to the angiogenesis signaling pathway were activated (increased expression of phosphorylated ERK), confirming that the process was mediated by the MAPK/ERK signaling pathway. Based on the analysis of human wound cell culture medium, Geng et al. [11] found that activating the MAPK/ERK signaling pathway could promote the increase of VEGF level, suggesting that the activation of the MAPK/ERK signaling pathway has a positive effect on wound healing.

3.1.3 The MAPK pathways can affect wound healing by affecting extracellular matrix remodeling

Wang et al. [12] found that activating the ERK/MAPK signaling pathway can increase the expression of matrix metalloproteinase in dermal fibroblasts and regulate extracellular matrix remodeling, thereby promoting wound repair. Ji et al. [13] demonstrated that RUNX1 transcriptionally activates osteopontin (OPN), which suppresses keratinocyte migration and ECM degradation by inhibiting MAPK pathway activity. Silencing OPN or downregulating RUNX1 releases this inhibition, thereby activating ERK/p38 MAPK signaling to enhance HaCaT cell proliferation, migration, and MMP-mediated ECM remodeling, ultimately accelerating burn wound healing.

3.1.4 The MAPK signaling pathway can affect wound healing by affecting cell proliferation and repair

Konstantinou et al. [14] studied the mechanism of microcurrent in treating chronic wounds and promoting their healing. The in vitro experiment suggested that microcurrent stimulated the phosphorylation of ERK1/2 and p38 kinases related to the MAPK signaling pathway in fibroblasts, induced cell migration and proliferation, and thus accelerated wound healing. Zhang et al. [15] confirmed through in vivo and in vitro experiments that the P38/MAPK signaling pathway in damaged epidermal keratinocytes induced microtubule (MT)-associated protein 4 (MAP4) phosphorylation under hypoxia, which has the ability to regulate cell migration and proliferation, and at the same time, it induced MT decomposition and accelerated keratinocyte proliferation and migration, which promoted wound healing.

3.2 PI3K/AKT signaling pathway

PI3K is a kind of lipid kinase in the cytoplasm, which is involved in the signal transduction, transport, and metabolism of the cell membrane. PI3K is a dimer composed of regulatory subunit P85 and catalytic subunit P110. When it binds to growth factor receptors (such as EGFR), it can change the protein structure of AKT and activate it and, at the same time, activate or inhibit a series of downstream substrates such as apoptosis-related proteins Bad and Caspase9 through phosphorylation, thus regulating cell proliferation, differentiation, apoptosis, and migration. What is more, protein kinase B (AKT) can also activate IKK and produce crosstalk with the NF-κB pathway. The downstream target of PI3K/AKT is mammalian target of rapamycin (mTOR), and the downstream transcription factors of mTOR include HIF1α, c-MYC, FoxO, and other star molecules. The PI3K/AKT signaling pathway is closely related to wound healing [16].

3.2.1 The PI3K/AKT pathways can affect wound healing by affecting vascular regeneration

PI3K can induce the phosphorylation of AKT by catalyzing the phosphorylation of phosphatidylinositol diphosphate, thereby regulating the transcriptional level of downstream endothelial nitric oxide synthase (eNOS) and stimulating the synthesis and release of nitric oxide [17]. As a potent angiogenic medium, nitric oxide plays a role in regulating angiogenic factors, not only stimulating neoangiogenesis but also participating in regulating the proliferation, invasion, apoptosis, and lumen formation of ECs, which can significantly delay wound healing [18]. Yuan et al. [19] adopted different concentrations of cinnamaldehyde to interface with human umbilical vein colorectal cells (HUVECs) and verified that cinnamaldehyde could induce the phosphorylation of AKT and eNOS, activating and up-regulating PI3K, so as to accelerate the angiogenesis and wound healing. Dong et al. [20] proved through an in vitro experiment that paeoniflorin could enhance the angiogenesis ability of HUVECs and facilitate wound healing by means of the PI3K/AKT signaling pathway.

3.2.2 The PI3K/AKT signaling pathway can affect wound healing by influencing inflammatory responses

Yu et al. [21] studied the mechanism of insulin on macrophages phenotype conversion by establishing a burn wound in diabetic rats model and human monocyte THP-1 model and found that local injection of low-dose insulin around the wound could promote macrophages to change from proinflammatory (M1) phenotype to anti-inflammatory (M2) phenotype through the PI3K/AKT signaling pathway, as to accelerate wound healing.

3.2.3 The PI3K/AKT pathways can affect wound healing by affecting extracellular matrix remodeling

Wang et al. [22] studied the effect of hypoxic adipose stem cells’ exosomes (HypadSCs-ExO) on wound healing by establishing a human fibroblast model and full-thickness skin resection in a diabetic nude rat model. The results suggested that relying on PI3K/AKT signaling pathway, HypADSCs-exo may regulate the proliferation and migration of fibroblasts and the secretion of inflammation, vascular-associated growth factors and extracellular matrix, and accelerate the high-quality wound healing in diabetic targets. Zhang et al. [23] studied the wounds of mice with whole-layer skin incision and found that activating the PI3K/AKT signaling pathway could promote the proliferation and migration of fibroblasts, and collagen deposition, thereby accelerating wound repair.

3.2.4 The PI3K/AKT signaling pathway can affect wound healing by affecting cell proliferation and repair

Li et al. probed into the impact and mechanism of Human amniotic mesenchymal stem cells (hAMSCs) on burn wound healing through in vivo and in vitro experiments, which manifested that hAMSCs inhibited cell apoptosis by activating the PI3K/AKT signaling pathway, and this signaling pathway could activate glycogen synthase kinase-3 beta (GSK-3β)/β-catenin signaling pathway to promote cell proliferation and in turn achieve wound injury treatment [24,25].

3.3 TGF-β signaling pathway

TGF-β family is a kind of secretory polypeptide signaling molecule, which is involved in the regulation of cell proliferation, differentiation, apoptosis, and wound healing. Smad is usually the downstream effector transcription factor of the TGF-β signaling pathway, and TGF-β1/2/3 and BMP (bone morphogenetic protein) are the signal sources of this pathway. Studies have shown that inhibition of TGF-β signal transduction could accelerate the closure of scarless wound [26]. Among them, TGF-β1 is one of the main factors affecting wound healing and scar formation. TGF-β1 can stimulate fibroblast proliferation and promote the transformation of stromal cells into fibroblasts. TGF-β1 can also promote the transformation of fibroblasts into myofibroblasts, which is beneficial to wound contraction and early wound closure. Besides, TGF-β1 can stimulate the proliferation of vascular ECs and promote early vascular remodeling, which is conducive to granulation formation [27,28,29]. Smad protein is an intermediary molecule that conducts the extracellular TGF-β signal through the cytoplasm to the nucleus. The activation of the TGF-β1/Smad3 signaling pathway can stimulate the synthesis of other fibrous proteins, promote the proliferation and phenotypic transformation of fibroblasts, inhibit fibroblast apoptosis, and promote the synthesis of various extracellular matrices, which are of great significance for wound healing.

3.3.1 TGF-β pathways can affect wound healing by affecting extracellular matrix remodeling

Shi et al. constructed an in vitro cell model and an in vivo animal model of ischemic ear wound to study the effects of clinical-grade platelets exosome products (PEP) developed and injectable surgical fibrin sealant (TISSEEL) on chronic ischemic wounds. The results showed that TISSEEL–PEP regulated the TGF-β pathway, including SMAD2, RAS, and other pathways, and promoted epithelialization enhancement, fibroblast activation, and collagen production, thus promoting ischemic wound healing. Tang et al. [30] also studied the effectiveness of Ginsenoside Rg3 in the treatment of scars through in vitro experiments, and the results showed that Rg3 could inhibit fibroblasts proliferation, angiogenesis, and collagen synthesis through TGF-β/Smad and ERK signaling pathways, so as to promote the perfect wound healing and effectively inhibit scar growth. Pan et al. [31] studied the relationship between angiogenin and scars by cultivating scar fibroblast model, and the results suggested that the expression of angiopoietin was negatively correlated with the severity of burn scars, that is, the increase of angiopoietin inhibited the expression level of TGF-β1/Smad2 and vice versa, which further confirmed that the excessive scar hyperplasia may be lightened and wound healing may be headlined via this mechanism [32,33].

3.3.2 The TGF-β pathways can affect wound healing by affecting vascular regeneration

Miscianinov et al. [34] explored the relationship between the TGF-β pathway and ROtung-to-mesenchymal transition (EndMT) by establishing an EC model, and the results showed that high expression level of Mir-148b enhanced EC proliferation and migration and in vitro vascular formation by regulating TGFB2 and SMAD2, which are targeted genes in the TGF-β signaling pathway, and at the same time, it weakened the EndMT process and accelerated wound closure.

3.4 Wnt signaling pathway

Wnt signaling pathway is closely related to wound repair, involving the proliferation and migration of fibroblasts and keratinocytes, extracellular matrix and collagen contraction, angiogenesis, etc. [35,36,37,38,39]. Generally, the Wnt pathway mainly refers to the classical signaling pathway mediated by β-Catenin, which is an important biomarker for detecting Wnt activation. The content of β-catenin in the cytoplasm directly affects the proliferation and differentiation of epidermal stem cells. The higher the content of β-catenin, the stronger the differentiation and proliferation abilities of epidermal cells. β-Catenin can promote the development of skin and its appendages, wound angiogenesis, and epithelial remodeling, thereby accelerating wound healing [40]. Studies have reported that in the process of diabetic wound healing, the differentiation and proliferation abilities of epidermal cells are enhanced with the increase in Wnt and β-catenin expression, which accelerates wound healing [41].

3.4.1 The Wnt pathways can affect wound healing by affecting extracellular matrix remodeling

Gay et al. [42] investigated the relevance between macrophages and fibrotic scars by establishing the wound-induced hair neogenesis (WIHN) in rat model, and the results showed that chronic Wnt activity in the WIHN model was correlated with fibrotic WIHN-scar formation, and fibrotic skin healing was achieved by the way that late macrophage devoured SFRP4, an inhibitor of Wnt, to drive chronic Wnt activity. Hu et al. [43] inquired into the effects of ganoderma lucidum polysaccharides (GL-PS) on wound healing by establishing a human skin fibroblast model and a mouse full-thickness dermoplasty model, and the results indicated that GL-PS up-regulated the expression of Wnt/β-catenin and TGF-β and boosted the viability and migration ability of fibroblasts, which promoted the wound healing rate and shortened the healing time.

3.4.2 The Wnt signaling pathway can affect wound healing by affecting cell proliferation and repair

He et al. [44] studied the mechanism of MalAT1-containing exosomes derived from adipose-derived stem cells (ADSC-ExOS) amid wound healing through in vitro experiments. The results manifested that ADSC-ExOS containing MALAT1 could significantly promote cell proliferation and migration and inhibit cell apoptosis, and in turn induce wound healing by activating target Mir-124 in the Wnt/β-catenin pathway.

3.5 NF-κB signaling pathway

As an important pathway that causes inflammatory response in the body, the NF-κB signaling pathway is widely involved in various inflammatory processes and plays an important role in the occurrence and treatment of various diseases. After transferring to the nucleus, the activated NF-κB binds to the promoter or enhancer region of target gene, to induce the production of adhesion molecules in ECs and enhance the inflammatory response of leukocytes and the proliferation of fibroblasts, thus exerting pro-inflammatory effects [45,46,47].

3.5.1 The NF-κB signaling pathway can affect wound healing by influencing inflammatory responses

Kong et al., through in vivo and in vitro experiments, discovered that the inhibition of NF-κB p65 nuclear translocation could suppress the inflammatory phenotype conversion of vascular smooth muscle cells (VSMCs), lighten inflammation responses, and promote angiogenesis. Sangiovanni et al. [48] explored the mechanism of Cannabis sativa L. ethanolic extract (CSE) in skin inflammation and wound injury through in vitro experiments. The results showed that CSE could inhibit TNF-α-induced NF-κB-driven transcription, and IL-8 and MMP-9 release, and exert anti-inflammatory activity, so as to promote wound healing. Chen et al. [49] studied the therapeutic effect and mechanism of thalidomide on rosacea-like mouse skin model induced by LL37. The results manifested that the overexpression of NF-κB in rosaceas led to distinct inflammatory reaction, and at the same time, thalidomide improved skin inflammation and promoted skin healing by inhibiting NF-κB expression. Romana-Souza et al. [50] studied the restorative effect of caffeic acid phenethyl ester (CAPE) on pressure ulcers through in vivo experiments, and the results showed that NF-κB was involved in the inflammatory reaction of pressure ulcers, leading to chronic inflammation and delaying wound closure, and CAPE might reduce the activation of NF-κB P65 and promote wound healing [51,52,53,54,55].

3.5.2 The NF-κB pathways can affect wound healing by affecting vascular regeneration

Studies have shown that VEGF transcription is regulated by the activation of the NF-κB signaling pathway; therefore, inhibiting the activation of the NF-κB signaling pathway can significantly inhibit VEGF expression and capillary formation [56].

3.6 Notch signaling pathway

Notch pathway does not transfer signals by gradual activation of kinase phosphorylation, but it releases the Notch protein fragments (NICD or ICN) with transcriptional regulatory activity through three-step protease hydrolysis and then binds them to the transcription factor CSL to regulate downstream gene expression. Notch signaling pathway is closely related to wound healing [57,58,59,60].

3.6.1 The Notch pathways can affect wound healing by affecting vascular regeneration

Li et al. [61] constructed a transgenic zebrafish model to study the impact of tetraspanin 18 (Tspan18) on angiogenesis, and the results demonstrated that arterial venous specification was mainly regulated by the Notch pathway, Tspan18 was expressed in blood vessels, and in addition, Notch and VEGF were regulated to promote angiogenesis and wound healing.

3.6.2 The Notch signaling pathway can affect wound healing by influencing inflammatory responses

He et al. used the RBP-J gene down-regulated by the Notch signaling to establish a wound inflammation model. In the experimental process, it was found that the blockade of the Notch signaling pathway could reduce the expression of inflammatory cytokines such as TGF-β1, CCL2, and TNF-α in macrophages and the infiltration of inflammatory cells, thus inhibiting the formation of pathological scars and promoting wound healing [62,63,64].

3.6.3 The Notch pathways can affect wound healing by affecting extracellular matrix remodeling

Ebrahim et al. verified that the joint regulation of platelet-rich plasma therapy and adipose-derived mesenchymal stem cell therapy effectively promoted wound re-epithelialization and granulation tissue formation in diabetic rats by regulating the Notch pathway, and the collagen area percentage, epidermal thickness, and angiogenesis were also significantly increased thereby [65,66].

3.7 Hippo signaling pathway

The Hippo pathway consists of a group of conserved kinases. After sensing the signals from the extracellular environment, the upstream membrane protein receptors undergo a series of kinase phosphorylation reactions and finally act on the downstream effectors YAP and TAZ. YAP/TAZ has the function of transcriptional regulation, which can fine-regulate the cell phenotype. Moreover, the Hippo pathway regulates cell proliferation, differentiation, and apoptosis by acting on the downstream effectors YAP and TAZ [67].

3.7.1 The Hippo pathways can affect wound healing by affecting extracellular matrix remodeling

Mascharak et al. [68] probed into the connection between ENF and scar formation by establishing cell transplantation and transgenic mouse models, and the results indicated that mechanical tension drove Engrailed-1 (EN-1) activation through typical mechanical transduction signals, and inhibition of YAP might block En1 activation in vivo to reduce scar production and promote wound regeneration and healing through ENF. Brewer et al. [69] found that the scarless regenerative wound healing characteristic of spiny mice was related to the target protein YAP in the Hippo signaling pathway and activation of YAP in vitro could prevent fibrosis and provide a new idea for promoting scarless regenerative wound healing in vivo.

3.7.2 The Hippo signaling pathway can affect wound healing by affecting cell proliferation and repair

Yuan et al. [70] demonstrated through in vivo and in vitro experiments that YAP1/TaZ-Tead inhibition would destroy skin homeostasis and lead to skin ulceration in mice. Shome et al. [71] found that the upregulation of Hippo transcription factor YAP and its downstream effectors CTGF and Cyr61 drove paracrine signals in the mechanism of cold atmospheric pressure plasma treatment for wound healing, which facilitated to cell migration and accelerated wound healing.

4 Crosstalk between signaling pathways

4.1 PI3K/AKT and MAPK synergy in angiogenesis

The interplay between the PI3K/AKT and MAPK signaling pathways is crucial for regulating angiogenesis in wound healing (Figure 2). PI3K/AKT signaling is primarily activated by receptor tyrosine kinases such as VEGFR2 and FGFR, leading to the phosphorylation of AKT, which stabilizes HIF-1α and enhances VEGF expression. Meanwhile, the MAPK/ERK pathway promotes EC proliferation and migration by activating downstream effectors such as matrix metalloproteinases and endothelial nitric oxide synthase.

Figure 2 Schematic diagram showing cross-talk between signaling pathways implicated in chronic wound healing.
Figure 2

Schematic diagram showing cross-talk between signaling pathways implicated in chronic wound healing.

Recent studies highlight that PI3K/AKT and MAPK pathways exhibit bidirectional regulation at multiple levels. PI3K/AKT can inhibit Raf phosphorylation, thereby downregulating MAPK signaling under certain conditions [72]. Conversely, ERK activation can phosphorylate and suppress TSC2, relieving mTOR inhibition and boosting AKT activity. This reciprocal regulation ensures a controlled balance between endothelial proliferation and vessel maturation [73].

In chronic wounds, prolonged hyperglycemia and oxidative stress impair PI3K/AKT activation, reducing VEGF-mediated angiogenesis. Simultaneously, sustained activation of MAPK, particularly the JNK and p38 branches, leads to cell apoptosis and excessive inflammatory signaling, further hindering capillary formation [74]. Moreover, the dysregulation of upstream regulators such as integrins and focal adhesion kinase disrupts cross-signaling between PI3K/AKT and MAPK, compounding angiogenic failure. Targeting these pathways simultaneously, for example, by using PI3K agonists alongside controlled MAPK inhibition, may help restore vascular homeostasis and improve wound healing [10].

4.2 NF-κB and TGF-β interplay in inflammation and extracellular matrix remodeling

The NF-κB and TGF-β signaling pathways exhibit intricate crosstalk in the regulation of inflammation and extracellular matrix remodeling. NF-κB is primarily activated by inflammatory stimuli via toll-like receptors, leading to increased transcription of cytokines such as TNF-α and IL-1β. Meanwhile, TGF-β is involved in the later stages of wound healing, facilitating fibroblast activation and extracellular matrix synthesis through Smad2 and Smad3-dependent transcriptional regulation.

Recent insights suggest that NF-κB and TGF-β pathways regulate each other through multiple feedback loops. NF-κB suppresses Smad3 activity, reducing TGF-β-mediated fibroblast differentiation and extracellular matrix production. In contrast, TGF-β can inhibit NF-κB signaling via Smad7, which interferes with IκB kinase activation, thereby limiting inflammatory responses [75]. However, in chronic wounds, persistent NF-κB activation prevents the transition to the proliferative phase, leading to prolonged inflammation and excessive matrix metalloproteinase expression, which degrades extracellular matrix components [76]. Additionally, TGF-β’s role in fibrosis is exacerbated in chronic wounds due to an imbalance in Smad-independent pathways, particularly the p38 MAPK and PI3K/AKT branches. Excessive TGF-β signaling leads to myofibroblast overactivation and excessive extracellular matrix deposition, contributing to fibrosis [77]. Furthermore, recent findings highlight that microRNAs, such as miR-146a and miR-200b, play a role in modulating the NF-κB-TGF-β interaction by regulating key signaling intermediates such as TRAF6 and Smad4 [72]. Therapeutic interventions targeting microRNAs or modulating the NF-κB–TGF-β balance may provide novel approaches to controlling inflammation and fibrosis in chronic wound healing.

4.3 Wnt/β-catenin and Hippo signaling crosstalk in cell proliferation

The interplay between Wnt/β-catenin and Hippo signaling is crucial for regulating keratinocyte and fibroblast proliferation during wound healing. Wnt/β-catenin signaling promotes cell proliferation by stabilizing β-catenin, allowing it to translocate into the nucleus and activate target genes involved in cell cycle progression. In contrast, Hippo signaling functions as a growth suppressor by phosphorylating and inactivating YAP and transcriptional coactivator with PDZ-binding motif (TAZ), thereby preventing excessive cell proliferation.

Crosstalk between these pathways occurs at multiple regulatory points. Hippo signaling components, particularly large tumor suppressor kinases (LATS1/2), can phosphorylate YAP/TAZ, promoting their cytoplasmic retention and degradation. This suppresses β-catenin activity, as YAP/TAZ are known to interact with β-catenin to enhance its nuclear translocation and transcriptional activation of Wnt target genes [78]. Conversely, Wnt signaling inhibits Hippo pathway activation by suppressing GSK-3β, which not only stabilizes β-catenin but also promotes YAP/TAZ activity. Additionally, YAP has been shown to bind to disheveled (DVL), a key mediator of Wnt signaling, further enhancing β-catenin-driven transcription [79]. In the context of wound healing, disruption of this crosstalk contributes to impaired tissue regeneration. Persistent inflammation and oxidative stress often suppress Wnt signaling, leading to reduced β-catenin activity and delayed keratinocyte migration. Simultaneously, aberrant Hippo pathway suppression can result in uncontrolled YAP/TAZ activation, driving excessive fibroblast proliferation and ECM deposition, ultimately leading to fibrosis and scar formation. This imbalance hinders proper wound closure and functional tissue restoration [72].

Targeting the Wnt/β-catenin-Hippo signaling crosstalk presents a promising therapeutic strategy. Enhancing Wnt signaling while fine-tuning Hippo pathway activity could promote controlled cell proliferation and re-epithelialization without exacerbating fibrosis. Small-molecule activators of Wnt or inhibitors of LATS1/2 could help restore β-catenin and YAP/TAZ activity in a regulated manner, fostering an optimal wound healing environment.

5 Therapeutic strategies targeting key signaling pathways in wound healing

Effective wound healing requires precise regulation of signaling pathways involved in inflammation resolution, angiogenesis, cell proliferation, and extracellular matrix remodeling. Dysregulation of these pathways in wound healing contributes to delayed healing and pathological tissue remodeling. Recent advances in targeted therapies, including small-molecule inhibitors, gene therapy, and biologics, have shown promise in modulating these pathways to restore normal wound healing dynamics. This section summarizes current therapeutic approaches targeting PI3K/AKT, MAPK, NF-κB, TGF-β, Wnt/β-catenin, and Hippo signaling.

5.1 Small-molecule inhibitors

Small-molecule inhibitors have been extensively investigated for their ability to modulate key signaling pathways in chronic wound healing. The PI3K/AKT pathway plays a central role in angiogenesis and cell survival, making it a potential target for wound therapy. Small-molecule agonists, such as SC79, an AKT activator, have been shown to promote EC proliferation and enhance vascular regeneration in preclinical wound models [80]. Conversely, excessive MAPK activation, particularly through the JNK and p38 branches, can contribute to inflammation-induced apoptosis. p38 inhibitors such as SB203580 have demonstrated efficacy in reducing inflammation and improving wound closure by preventing excessive fibroblast senescence [81].

Persistent NF-κB activation in chronic wounds sustains an inflammatory microenvironment, while excessive TGF-β signaling promotes fibrosis. Small-molecule NF-κB inhibitors have been explored for their ability to downregulate pro-inflammatory cytokines [82]. In contrast, selective TGF-β modulators, including losartan and pirfenidone, have been investigated for their antifibrotic properties, promoting balanced extracellular matrix remodeling [83,84].

Wnt activators such as lithium chloride and CHIR99021 enhance cellular proliferation and migration by stabilizing β-catenin [85,86]. Meanwhile, Hippo pathway inhibitors, such as verteporfin, reduce excessive fibroblast proliferation by preventing YAP/TAZ nuclear translocation, thus limiting fibrosis [87].

5.2 Gene therapy

Gene therapy has emerged as a promising strategy for directly modulating key signaling pathways involved in chronic wound pathophysiology. Viral and non-viral gene delivery strategies have been employed to enhance AKT activity in chronic wounds. Gene therapy using VEGF-A-expressing collagen-mimetic peptide tethers has been shown to activate the PI3K/AKT pathway, promoting angiogenesis and accelerating wound closure [88]. In parallel, CRISPR/Cas9-mediated editing of TGF-β receptors has been investigated for selectively modulating fibrotic responses while preserving its role in extracellular matrix deposition [89]. Overexpression of Wnt ligands or β-catenin-stabilizing constructs has been shown to improve epithelial regeneration in chronic wounds. Conversely, knockdown of YAP using shRNA-based gene therapy has been explored to prevent fibroblast hyperproliferation and excessive scarring [90].

5.3 Biologic therapy

Recombinant VEGF and PDGF have been used to enhance angiogenesis by stimulating PI3K/AKT and MAPK pathways, though their efficacy has been variable due to rapid degradation in the wound microenvironment. Encapsulation strategies using nanoparticles have been developed to enhance the stability and delivery efficiency of these growth factors [91].

Monoclonal antibodies targeting pro-inflammatory cytokines, such as anti-TNF agents (infliximab) and IL-6 inhibitors (tocilizumab), have been investigated for modulating NF-κB-mediated inflammation. Meanwhile, TGF-β-neutralizing antibodies, such as fresolimumab, have been tested for preventing excessive fibrosis in chronic wounds [92].

Exosomes derived from stem cells have gained attention for their ability to modulate multiple signaling pathways simultaneously. Mesenchymal stem cell-derived exosomes have been shown to enhance diabetic wound healing and skin regeneration [93].

6 Conclusion and future directions

Wound healing arises from the complex dysregulation of multiple signaling pathways, including PI3K/AKT, MAPK, NF-κB, TGF-β, Wnt/β-catenin, and Hippo. While significant progress has been made in understanding their roles in wound healing, several challenges remain. The intricate crosstalk between these pathways is not yet fully elucidated, making it difficult to develop precise therapeutic strategies that effectively restore wound healing without unintended side effects. Existing treatments, such as growth factors and anti-inflammatory agents, often suffer from limited efficacy due to rapid degradation, poor bioavailability, and non-specific targeting. Although preclinical studies on small-molecule inhibitors, gene therapy, and biologics have demonstrated promising results, the translation of these findings into clinical practice remains challenging due to safety concerns, inconsistent outcomes, and a lack of large-scale clinical validation. Moreover, patient variability, including differences in wound etiology, metabolic conditions, and genetic predispositions, complicates the development of universally effective therapies.

Future research should focus on elucidating the dynamic interactions between these pathways using advanced multi-omics technologies, such as single-cell RNA sequencing and proteomics, to better understand how signaling networks contribute to impaired healing. Instead of targeting single pathways, combination therapies that fine-tune multiple signaling cascades may offer better outcomes by addressing both inflammation and tissue regeneration while preventing fibrosis. Advances in nanotechnology-based drug delivery systems, including hydrogels and biomaterials, hold great potential for improving the stability, bioavailability, and targeted release of therapeutic agents at the wound site. Additionally, personalized medicine approaches, guided by artificial intelligence and machine learning, could help stratify patients based on molecular profiles, enabling precision-targeted therapies tailored to individual wound characteristics. Finally, bridging the gap between preclinical research and clinical application will require well-designed clinical trials that assess the long-term efficacy and safety of emerging therapies, optimize dosing strategies, and minimize adverse effects. By addressing these challenges, future research can pave the way for more effective and personalized treatments for wound healing, ultimately improving patient outcomes and reducing healthcare burdens.

# These authors contributed equally to this work.

  1. Funding information: The work was supported by the Science and Technology Young Talent” Program of Shanghai Integrated Traditional Chinese and Western Medicine Hospital (2024KJCY002), Traditional Chinese Medicine Scientific Research Project of Health Commission of Hongkou District, Shanghai (HKQGYQY-ZYY-2023-02), Scientific and Technological Development Project of Shanghai University of Traditional Chinese Medicine (24KFL085), and National Natural Science Foundation of China (82174382).

  2. Author contributions: Zhe Liu participated in the search and collection of data. Zhe Liu drafted the manuscript, while Yudong Fang provided guidance on article conceptualization and revision suggestions. All authors read and approved the final manuscript.

  3. Conflict of interest: Authors state no conflict of interest.

  4. Data availability statement: Data sharing is not applicable to this article as no datasets were generated or analyzed during the current study.

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Received: 2024-11-12
Revised: 2025-07-14
Accepted: 2025-07-26
Published Online: 2025-09-01

© 2025 the author(s), published by De Gruyter

This work is licensed under the Creative Commons Attribution 4.0 International License.

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  9. Efficacy of alginate-coated gold nanoparticles against antibiotics-resistant Staphylococcus and Streptococcus pathogens of acne origins
  10. Battling COVID-19 leveraging nanobiotechnology: Gold and silver nanoparticle–B-escin conjugates as SARS-CoV-2 inhibitors
  11. Neurodegenerative diseases and neuroinflammation-induced apoptosis
  12. Impact of fracture fixation surgery on cognitive function and the gut microbiota in mice with a history of stroke
  13. COLEC10: A potential tumor suppressor and prognostic biomarker in hepatocellular carcinoma through modulation of EMT and PI3K-AKT pathways
  14. High-temperature requirement serine protease A2 inhibitor UCF-101 ameliorates damaged neurons in traumatic brain-injured rats by the AMPK/NF-κB pathway
  15. SIK1 inhibits IL-1β-stimulated cartilage apoptosis and inflammation in vitro through the CRTC2/CREB1 signaling
  16. Rutin–chitooligosaccharide complex: Comprehensive evaluation of its anti-inflammatory and analgesic properties in vitro and in vivo
  17. Knockdown of Aurora kinase B alleviates high glucose-triggered trophoblast cells damage and inflammation during gestational diabetes
  18. Calcium-sensing receptors promoted Homer1 expression and osteogenic differentiation in bone marrow mesenchymal stem cells
  19. ABI3BP can inhibit the proliferation, invasion, and epithelial–mesenchymal transition of non-small-cell lung cancer cells
  20. Changes in blood glucose and metabolism in hyperuricemia mice
  21. Rapid detection of the GJB2 c.235delC mutation based on CRISPR-Cas13a combined with lateral flow dipstick
  22. IL-11 promotes Ang II-induced autophagy inhibition and mitochondrial dysfunction in atrial fibroblasts
  23. Short-chain fatty acid attenuates intestinal inflammation by regulation of gut microbial composition in antibiotic-associated diarrhea
  24. Application of metagenomic next-generation sequencing in the diagnosis of pathogens in patients with diabetes complicated by community-acquired pneumonia
  25. NAT10 promotes radiotherapy resistance in non-small cell lung cancer by regulating KPNB1-mediated PD-L1 nuclear translocation
  26. Phytol-mixed micelles alleviate dexamethasone-induced osteoporosis in zebrafish: Activation of the MMP3–OPN–MAPK pathway-mediating bone remodeling
  27. Association between TGF-β1 and β-catenin expression in the vaginal wall of patients with pelvic organ prolapse
  28. Primary pleomorphic liposarcoma involving bilateral ovaries: Case report and literature review
  29. Effects of de novo donor-specific Class I and II antibodies on graft outcomes after liver transplantation: A pilot cohort study
  30. Sleep architecture in Alzheimer’s disease continuum: The deep sleep question
  31. Ephedra fragilis plant extract: A groundbreaking corrosion inhibitor for mild steel in acidic environments – electrochemical, EDX, DFT, and Monte Carlo studies
  32. Langerhans cell histiocytosis in an adult patient with upper jaw and pulmonary involvement: A case report
  33. Inhibition of mast cell activation by Jaranol-targeted Pirin ameliorates allergic responses in mouse allergic rhinitis
  34. Aeromonas veronii-induced septic arthritis of the hip in a child with acute lymphoblastic leukemia
  35. Clusterin activates the heat shock response via the PI3K/Akt pathway to protect cardiomyocytes from high-temperature-induced apoptosis
  36. Research progress on fecal microbiota transplantation in tumor prevention and treatment
  37. Low-pressure exposure influences the development of HAPE
  38. Stigmasterol alleviates endplate chondrocyte degeneration through inducing mitophagy by enhancing PINK1 mRNA acetylation via the ESR1/NAT10 axis
  39. AKAP12, mediated by transcription factor 21, inhibits cell proliferation, metastasis, and glycolysis in lung squamous cell carcinoma
  40. Association between PAX9 or MSX1 gene polymorphism and tooth agenesis risk: A meta-analysis
  41. A case of bloodstream infection caused by Neisseria gonorrhoeae
  42. Case of nasopharyngeal tuberculosis complicated with cervical lymph node and pulmonary tuberculosis
  43. p-Cymene inhibits pro-fibrotic and inflammatory mediators to prevent hepatic dysfunction
  44. GFPT2 promotes paclitaxel resistance in epithelial ovarian cancer cells via activating NF-κB signaling pathway
  45. Transfer RNA-derived fragment tRF-36 modulates varicose vein progression via human vascular smooth muscle cell Notch signaling
  46. RTA-408 attenuates the hepatic ischemia reperfusion injury in mice possibly by activating the Nrf2/HO-1 signaling pathway
  47. Decreased serum TIMP4 levels in patients with rheumatoid arthritis
  48. Sirt1 protects lupus nephritis by inhibiting the NLRP3 signaling pathway in human glomerular mesangial cells
  49. Sodium butyrate aids brain injury repair in neonatal rats
  50. Interaction of MTHFR polymorphism with PAX1 methylation in cervical cancer
  51. Convallatoxin inhibits proliferation and angiogenesis of glioma cells via regulating JAK/STAT3 pathway
  52. The effect of the PKR inhibitor, 2-aminopurine, on the replication of influenza A virus, and segment 8 mRNA splicing
  53. Effects of Ire1 gene on virulence and pathogenicity of Candida albicans
  54. Small cell lung cancer with small intestinal metastasis: Case report and literature review
  55. GRB14: A prognostic biomarker driving tumor progression in gastric cancer through the PI3K/AKT signaling pathway by interacting with COBLL1
  56. 15-Lipoxygenase-2 deficiency induces foam cell formation that can be restored by salidroside through the inhibition of arachidonic acid effects
  57. FTO alleviated the diabetic nephropathy progression by regulating the N6-methyladenosine levels of DACT1
  58. Clinical relevance of inflammatory markers in the evaluation of severity of ulcerative colitis: A retrospective study
  59. Zinc valproic acid complex promotes osteoblast differentiation and exhibits anti-osteoporotic potential
  60. Primary pulmonary synovial sarcoma in the bronchial cavity: A case report
  61. Metagenomic next-generation sequencing of alveolar lavage fluid improves the detection of pulmonary infection
  62. Uterine tumor resembling ovarian sex cord tumor with extensive rhabdoid differentiation: A case report
  63. Genomic analysis of a novel ST11(PR34365) Clostridioides difficile strain isolated from the human fecal of a CDI patient in Guizhou, China
  64. Effects of tiered cardiac rehabilitation on CRP, TNF-α, and physical endurance in older adults with coronary heart disease
  65. Changes in T-lymphocyte subpopulations in patients with colorectal cancer before and after acupoint catgut embedding acupuncture observation
  66. Modulating the tumor microenvironment: The role of traditional Chinese medicine in improving lung cancer treatment
  67. Alterations of metabolites related to microbiota–gut–brain axis in plasma of colon cancer, esophageal cancer, stomach cancer, and lung cancer patients
  68. Research on individualized drug sensitivity detection technology based on bio-3D printing technology for precision treatment of gastrointestinal stromal tumors
  69. CEBPB promotes ulcerative colitis-associated colorectal cancer by stimulating tumor growth and activating the NF-κB/STAT3 signaling pathway
  70. Oncolytic bacteria: A revolutionary approach to cancer therapy
  71. A de novo meningioma with rapid growth: A possible malignancy imposter?
  72. Diagnosis of secondary tuberculosis infection in an asymptomatic elderly with cancer using next-generation sequencing: Case report
  73. Hesperidin and its zinc(ii) complex enhance osteoblast differentiation and bone formation: In vitro and in vivo evaluations
  74. Research progress on the regulation of autophagy in cardiovascular diseases by chemokines
  75. Anti-arthritic, immunomodulatory, and inflammatory regulation by the benzimidazole derivative BMZ-AD: Insights from an FCA-induced rat model
  76. Immunoassay for pyruvate kinase M1/2 as an Alzheimer’s biomarker in CSF
  77. The role of HDAC11 in age-related hearing loss: Mechanisms and therapeutic implications
  78. Evaluation and application analysis of animal models of PIPNP based on data mining
  79. Therapeutic approaches for liver fibrosis/cirrhosis by targeting pyroptosis
  80. Fabrication of zinc oxide nanoparticles using Ruellia tuberosa leaf extract induces apoptosis through P53 and STAT3 signalling pathways in prostate cancer cells
  81. Haplo-hematopoietic stem cell transplantation and immunoradiotherapy for severe aplastic anemia complicated with nasopharyngeal carcinoma: A case report
  82. Modulation of the KEAP1-NRF2 pathway by Erianin: A novel approach to reduce psoriasiform inflammation and inflammatory signaling
  83. The expression of epidermal growth factor receptor 2 and its relationship with tumor-infiltrating lymphocytes and clinical pathological features in breast cancer patients
  84. Innovations in MALDI-TOF Mass Spectrometry: Bridging modern diagnostics and historical insights
  85. BAP1 complexes with YY1 and RBBP7 and its downstream targets in ccRCC cells
  86. Hypereosinophilic syndrome with elevated IgG4 and T-cell clonality: A report of two cases
  87. Electroacupuncture alleviates sciatic nerve injury in sciatica rats by regulating BDNF and NGF levels, myelin sheath degradation, and autophagy
  88. Polydatin prevents cholesterol gallstone formation by regulating cholesterol metabolism via PPAR-γ signaling
  89. RNF144A and RNF144B: Important molecules for health
  90. Analysis of the detection rate and related factors of thyroid nodules in the healthy population
  91. Artesunate inhibits hepatocellular carcinoma cell migration and invasion through OGA-mediated O-GlcNAcylation of ZEB1
  92. Endovascular management of post-pancreatectomy hemorrhage caused by a hepatic artery pseudoaneurysm: Case report and review of the literature
  93. Efficacy and safety of anti-PD-1/PD-L1 antibodies in patients with relapsed refractory diffuse large B-cell lymphoma: A meta-analysis
  94. SATB2 promotes humeral fracture healing in rats by activating the PI3K/AKT pathway
  95. Overexpression of the ferroptosis-related gene, NFS1, corresponds to gastric cancer growth and tumor immune infiltration
  96. Understanding risk factors and prognosis in diabetic foot ulcers
  97. Atractylenolide I alleviates the experimental allergic response in mice by suppressing TLR4/NF-kB/NLRP3 signalling
  98. FBXO31 inhibits the stemness characteristics of CD147 (+) melanoma stem cells
  99. Immune molecule diagnostics in colorectal cancer: CCL2 and CXCL11
  100. Inhibiting CXCR6 promotes senescence of activated hepatic stellate cells with limited proinflammatory SASP to attenuate hepatic fibrosis
  101. Cadmium toxicity, health risk and its remediation using low-cost biochar adsorbents
  102. Pulmonary cryptococcosis with headache as the first presentation: A case report
  103. Solitary pulmonary metastasis with cystic airspaces in colon cancer: A rare case report
  104. RUNX1 promotes denervation-induced muscle atrophy by activating the JUNB/NF-κB pathway and driving M1 macrophage polarization
  105. Morphometric analysis and immunobiological investigation of Indigofera oblongifolia on the infected lung with Plasmodium chabaudi
  106. The NuA4/TIP60 histone-modifying complex and Hr78 modulate the Lobe2 mutant eye phenotype
  107. Experimental study on salmon demineralized bone matrix loaded with recombinant human bone morphogenetic protein-2: In vitro and in vivo study
  108. A case of IgA nephropathy treated with a combination of telitacicept and half-dose glucocorticoids
  109. Analgesic and toxicological evaluation of cannabidiol-rich Moroccan Cannabis sativa L. (Khardala variety) extract: Evidence from an in vivo and in silico study
  110. Wound healing and signaling pathways
  111. Combination of immunotherapy and whole-brain radiotherapy on prognosis of patients with multiple brain metastases: A retrospective cohort study
  112. To explore the relationship between endometrial hyperemia and polycystic ovary syndrome
  113. Research progress on the impact of curcumin on immune responses in breast cancer
  114. Biogenic Cu/Ni nanotherapeutics from Descurainia sophia (L.) Webb ex Prantl seeds for the treatment of lung cancer
  115. Dapagliflozin attenuates atrial fibrosis via the HMGB1/RAGE pathway in atrial fibrillation rats
  116. Glycitein alleviates inflammation and apoptosis in keratinocytes via ROS-associated PI3K–Akt signalling pathway
  117. ADH5 inhibits proliferation but promotes EMT in non-small cell lung cancer cell through activating Smad2/Smad3
  118. Apoptotic efficacies of AgNPs formulated by Syzygium aromaticum leaf extract on 32D-FLT3-ITD human leukemia cell line with PI3K/AKT/mTOR signaling pathway
  119. Novel cuproptosis-related genes C1QBP and PFKP identified as prognostic and therapeutic targets in lung adenocarcinoma
  120. Bee venom promotes exosome secretion and alters miRNA cargo in T cells
  121. Treatment of pure red cell aplasia in a chronic kidney disease patient with roxadustat: A case report
  122. Comparative bioinformatics analysis of the Wnt pathway in breast cancer: Selection of novel biomarker panels associated with ER status
  123. Kynurenine facilitates renal cell carcinoma progression by suppressing M2 macrophage pyroptosis through inhibition of CASP1 cleavage
  124. RFX5 promotes the growth, motility, and inhibits apoptosis of gastric adenocarcinoma cells through the SIRT1/AMPK axis
  125. ALKBH5 exacerbates early cardiac damage after radiotherapy for breast cancer via m6A demethylation of TLR4
  126. Phytochemicals of Roman chamomile: Antioxidant, anti-aging, and whitening activities of distillation residues
  127. Circadian gene Cry1 inhibits the tumorigenicity of hepatocellular carcinoma by the BAX/BCL2-mediated apoptosis pathway
  128. The TNFR-RIPK1/RIPK3 signalling pathway mediates the effect of lanthanum on necroptosis of nerve cells
  129. Ecology and Environmental Science
  130. Optimization and comparative study of Bacillus consortia for cellulolytic potential and cellulase enzyme activity
  131. The complete mitochondrial genome analysis of Haemaphysalis hystricis Supino, 1897 (Ixodida: Ixodidae) and its phylogenetic implications
  132. Epidemiological characteristics and risk factors analysis of multidrug-resistant tuberculosis among tuberculosis population in Huzhou City, Eastern China
  133. Indices of human impacts on landscapes: How do they reflect the proportions of natural habitats?
  134. Genetic analysis of the Siberian flying squirrel population in the northern Changbai Mountains, Northeast China: Insights into population status and conservation
  135. Diversity and environmental drivers of Suillus communities in Pinus sylvestris var. mongolica forests of Inner Mongolia
  136. Global assessment of the fate of nitrogen deposition in forest ecosystems: Insights from 15N tracer studies
  137. Fungal and bacterial pathogenic co-infections mainly lead to the assembly of microbial community in tobacco stems
  138. Influencing of coal industry related airborne particulate matter on ocular surface tear film injury and inflammatory factor expression in Sprague-Dawley rats
  139. Agriculture
  140. Integrated analysis of transcriptome, sRNAome, and degradome involved in the drought-response of maize Zhengdan958
  141. Variation in flower frost tolerance among seven apple cultivars and transcriptome response patterns in two contrastingly frost-tolerant selected cultivars
  142. Heritability of durable resistance to stripe rust in bread wheat (Triticum aestivum L.)
  143. Animal Science
  144. Effect of sex ratio on the life history traits of an important invasive species, Spodoptera frugiperda
  145. Plant Sciences
  146. Hairpin in a haystack: In silico identification and characterization of plant-conserved microRNA in Rafflesiaceae
  147. Widely targeted metabolomics of different tissues in Rubus corchorifolius
  148. The complete chloroplast genome of Gerbera piloselloides (L.) Cass., 1820 (Carduoideae, Asteraceae) and its phylogenetic analysis
  149. Field trial to correlate mineral solubilization activity of Pseudomonas aeruginosa and biochemical content of groundnut plants
  150. Correlation analysis between semen routine parameters and sperm DNA fragmentation index in patients with semen non-liquefaction: A retrospective study
  151. Plasticity of the anatomical traits of Rhododendron L. (Ericaceae) leaves and its implications in adaptation to the plateau environment
  152. Effects of Piriformospora indica and arbuscular mycorrhizal fungus on growth and physiology of Moringa oleifera under low-temperature stress
  153. Effects of different sources of potassium fertiliser on yield, fruit quality and nutrient absorption in “Harward” kiwifruit (Actinidia deliciosa)
  154. Comparative efficiency and residue levels of spraying programs against powdery mildew in grape varieties
  155. The DREB7 transcription factor enhances salt tolerance in soybean plants under salt stress
  156. Food Science
  157. Phytochemical analysis of Stachys iva: Discovering the optimal extract conditions and its bioactive compounds
  158. Review on role of honey in disease prevention and treatment through modulation of biological activities
  159. Computational analysis of polymorphic residues in maltose and maltotriose transporters of a wild Saccharomyces cerevisiae strain
  160. Optimization of phenolic compound extraction from Tunisian squash by-products: A sustainable approach for antioxidant and antibacterial applications
  161. Liupao tea aqueous extract alleviates dextran sulfate sodium-induced ulcerative colitis in rats by modulating the gut microbiota
  162. Toxicological qualities and detoxification trends of fruit by-products for valorization: A review
  163. Polyphenolic spectrum of cornelian cherry fruits and their health-promoting effect
  164. Optimizing the encapsulation of the refined extract of squash peels for functional food applications: A sustainable approach to reduce food waste
  165. Advancements in curcuminoid formulations: An update on bioavailability enhancement strategies curcuminoid bioavailability and formulations
  166. Impact of saline sprouting on antioxidant properties and bioactive compounds in chia seeds
  167. The dilemma of food genetics and improvement
  168. Bioengineering and Biotechnology
  169. Impact of hyaluronic acid-modified hafnium metalorganic frameworks containing rhynchophylline on Alzheimer’s disease
  170. Emerging patterns in nanoparticle-based therapeutic approaches for rheumatoid arthritis: A comprehensive bibliometric and visual analysis spanning two decades
  171. Application of CRISPR/Cas gene editing for infectious disease control in poultry
  172. Preparation of hafnium nitride-coated titanium implants by magnetron sputtering technology and evaluation of their antibacterial properties and biocompatibility
  173. Preparation and characterization of lemongrass oil nanoemulsion: Antimicrobial, antibiofilm, antioxidant, and anticancer activities
  174. Corrigendum
  175. Corrigendum to “Utilization of convolutional neural networks to analyze microscopic images for high-throughput screening of mesenchymal stem cells”
  176. Corrigendum to “Effects of Ire1 gene on virulence and pathogenicity of Candida albicans”
https://doi.org/10.1515/biol-2025-1166
Schlagwörter für diesen Artikel
wound healing; signaling pathway; crosstalk; therapeutic strategies
Creative Commons
BY 4.0

Artikel in diesem Heft

  1. Biomedical Sciences
  2. Mechanism of triptolide regulating proliferation and apoptosis of hepatoma cells by inhibiting JAK/STAT pathway
  3. Maslinic acid improves mitochondrial function and inhibits oxidative stress and autophagy in human gastric smooth muscle cells
  4. Comparative analysis of inflammatory biomarkers for the diagnosis of neonatal sepsis: IL-6, IL-8, SAA, CRP, and PCT
  5. Post-pandemic insights on COVID-19 and premature ovarian insufficiency
  6. Proteome differences of dental stem cells between permanent and deciduous teeth by data-independent acquisition proteomics
  7. Optimizing a modified cetyltrimethylammonium bromide protocol for fungal DNA extraction: Insights from multilocus gene amplification
  8. Preliminary analysis of the role of small hepatitis B surface proteins mutations in the pathogenesis of occult hepatitis B infection via the endoplasmic reticulum stress-induced UPR-ERAD pathway
  9. Efficacy of alginate-coated gold nanoparticles against antibiotics-resistant Staphylococcus and Streptococcus pathogens of acne origins
  10. Battling COVID-19 leveraging nanobiotechnology: Gold and silver nanoparticle–B-escin conjugates as SARS-CoV-2 inhibitors
  11. Neurodegenerative diseases and neuroinflammation-induced apoptosis
  12. Impact of fracture fixation surgery on cognitive function and the gut microbiota in mice with a history of stroke
  13. COLEC10: A potential tumor suppressor and prognostic biomarker in hepatocellular carcinoma through modulation of EMT and PI3K-AKT pathways
  14. High-temperature requirement serine protease A2 inhibitor UCF-101 ameliorates damaged neurons in traumatic brain-injured rats by the AMPK/NF-κB pathway
  15. SIK1 inhibits IL-1β-stimulated cartilage apoptosis and inflammation in vitro through the CRTC2/CREB1 signaling
  16. Rutin–chitooligosaccharide complex: Comprehensive evaluation of its anti-inflammatory and analgesic properties in vitro and in vivo
  17. Knockdown of Aurora kinase B alleviates high glucose-triggered trophoblast cells damage and inflammation during gestational diabetes
  18. Calcium-sensing receptors promoted Homer1 expression and osteogenic differentiation in bone marrow mesenchymal stem cells
  19. ABI3BP can inhibit the proliferation, invasion, and epithelial–mesenchymal transition of non-small-cell lung cancer cells
  20. Changes in blood glucose and metabolism in hyperuricemia mice
  21. Rapid detection of the GJB2 c.235delC mutation based on CRISPR-Cas13a combined with lateral flow dipstick
  22. IL-11 promotes Ang II-induced autophagy inhibition and mitochondrial dysfunction in atrial fibroblasts
  23. Short-chain fatty acid attenuates intestinal inflammation by regulation of gut microbial composition in antibiotic-associated diarrhea
  24. Application of metagenomic next-generation sequencing in the diagnosis of pathogens in patients with diabetes complicated by community-acquired pneumonia
  25. NAT10 promotes radiotherapy resistance in non-small cell lung cancer by regulating KPNB1-mediated PD-L1 nuclear translocation
  26. Phytol-mixed micelles alleviate dexamethasone-induced osteoporosis in zebrafish: Activation of the MMP3–OPN–MAPK pathway-mediating bone remodeling
  27. Association between TGF-β1 and β-catenin expression in the vaginal wall of patients with pelvic organ prolapse
  28. Primary pleomorphic liposarcoma involving bilateral ovaries: Case report and literature review
  29. Effects of de novo donor-specific Class I and II antibodies on graft outcomes after liver transplantation: A pilot cohort study
  30. Sleep architecture in Alzheimer’s disease continuum: The deep sleep question
  31. Ephedra fragilis plant extract: A groundbreaking corrosion inhibitor for mild steel in acidic environments – electrochemical, EDX, DFT, and Monte Carlo studies
  32. Langerhans cell histiocytosis in an adult patient with upper jaw and pulmonary involvement: A case report
  33. Inhibition of mast cell activation by Jaranol-targeted Pirin ameliorates allergic responses in mouse allergic rhinitis
  34. Aeromonas veronii-induced septic arthritis of the hip in a child with acute lymphoblastic leukemia
  35. Clusterin activates the heat shock response via the PI3K/Akt pathway to protect cardiomyocytes from high-temperature-induced apoptosis
  36. Research progress on fecal microbiota transplantation in tumor prevention and treatment
  37. Low-pressure exposure influences the development of HAPE
  38. Stigmasterol alleviates endplate chondrocyte degeneration through inducing mitophagy by enhancing PINK1 mRNA acetylation via the ESR1/NAT10 axis
  39. AKAP12, mediated by transcription factor 21, inhibits cell proliferation, metastasis, and glycolysis in lung squamous cell carcinoma
  40. Association between PAX9 or MSX1 gene polymorphism and tooth agenesis risk: A meta-analysis
  41. A case of bloodstream infection caused by Neisseria gonorrhoeae
  42. Case of nasopharyngeal tuberculosis complicated with cervical lymph node and pulmonary tuberculosis
  43. p-Cymene inhibits pro-fibrotic and inflammatory mediators to prevent hepatic dysfunction
  44. GFPT2 promotes paclitaxel resistance in epithelial ovarian cancer cells via activating NF-κB signaling pathway
  45. Transfer RNA-derived fragment tRF-36 modulates varicose vein progression via human vascular smooth muscle cell Notch signaling
  46. RTA-408 attenuates the hepatic ischemia reperfusion injury in mice possibly by activating the Nrf2/HO-1 signaling pathway
  47. Decreased serum TIMP4 levels in patients with rheumatoid arthritis
  48. Sirt1 protects lupus nephritis by inhibiting the NLRP3 signaling pathway in human glomerular mesangial cells
  49. Sodium butyrate aids brain injury repair in neonatal rats
  50. Interaction of MTHFR polymorphism with PAX1 methylation in cervical cancer
  51. Convallatoxin inhibits proliferation and angiogenesis of glioma cells via regulating JAK/STAT3 pathway
  52. The effect of the PKR inhibitor, 2-aminopurine, on the replication of influenza A virus, and segment 8 mRNA splicing
  53. Effects of Ire1 gene on virulence and pathogenicity of Candida albicans
  54. Small cell lung cancer with small intestinal metastasis: Case report and literature review
  55. GRB14: A prognostic biomarker driving tumor progression in gastric cancer through the PI3K/AKT signaling pathway by interacting with COBLL1
  56. 15-Lipoxygenase-2 deficiency induces foam cell formation that can be restored by salidroside through the inhibition of arachidonic acid effects
  57. FTO alleviated the diabetic nephropathy progression by regulating the N6-methyladenosine levels of DACT1
  58. Clinical relevance of inflammatory markers in the evaluation of severity of ulcerative colitis: A retrospective study
  59. Zinc valproic acid complex promotes osteoblast differentiation and exhibits anti-osteoporotic potential
  60. Primary pulmonary synovial sarcoma in the bronchial cavity: A case report
  61. Metagenomic next-generation sequencing of alveolar lavage fluid improves the detection of pulmonary infection
  62. Uterine tumor resembling ovarian sex cord tumor with extensive rhabdoid differentiation: A case report
  63. Genomic analysis of a novel ST11(PR34365) Clostridioides difficile strain isolated from the human fecal of a CDI patient in Guizhou, China
  64. Effects of tiered cardiac rehabilitation on CRP, TNF-α, and physical endurance in older adults with coronary heart disease
  65. Changes in T-lymphocyte subpopulations in patients with colorectal cancer before and after acupoint catgut embedding acupuncture observation
  66. Modulating the tumor microenvironment: The role of traditional Chinese medicine in improving lung cancer treatment
  67. Alterations of metabolites related to microbiota–gut–brain axis in plasma of colon cancer, esophageal cancer, stomach cancer, and lung cancer patients
  68. Research on individualized drug sensitivity detection technology based on bio-3D printing technology for precision treatment of gastrointestinal stromal tumors
  69. CEBPB promotes ulcerative colitis-associated colorectal cancer by stimulating tumor growth and activating the NF-κB/STAT3 signaling pathway
  70. Oncolytic bacteria: A revolutionary approach to cancer therapy
  71. A de novo meningioma with rapid growth: A possible malignancy imposter?
  72. Diagnosis of secondary tuberculosis infection in an asymptomatic elderly with cancer using next-generation sequencing: Case report
  73. Hesperidin and its zinc(ii) complex enhance osteoblast differentiation and bone formation: In vitro and in vivo evaluations
  74. Research progress on the regulation of autophagy in cardiovascular diseases by chemokines
  75. Anti-arthritic, immunomodulatory, and inflammatory regulation by the benzimidazole derivative BMZ-AD: Insights from an FCA-induced rat model
  76. Immunoassay for pyruvate kinase M1/2 as an Alzheimer’s biomarker in CSF
  77. The role of HDAC11 in age-related hearing loss: Mechanisms and therapeutic implications
  78. Evaluation and application analysis of animal models of PIPNP based on data mining
  79. Therapeutic approaches for liver fibrosis/cirrhosis by targeting pyroptosis
  80. Fabrication of zinc oxide nanoparticles using Ruellia tuberosa leaf extract induces apoptosis through P53 and STAT3 signalling pathways in prostate cancer cells
  81. Haplo-hematopoietic stem cell transplantation and immunoradiotherapy for severe aplastic anemia complicated with nasopharyngeal carcinoma: A case report
  82. Modulation of the KEAP1-NRF2 pathway by Erianin: A novel approach to reduce psoriasiform inflammation and inflammatory signaling
  83. The expression of epidermal growth factor receptor 2 and its relationship with tumor-infiltrating lymphocytes and clinical pathological features in breast cancer patients
  84. Innovations in MALDI-TOF Mass Spectrometry: Bridging modern diagnostics and historical insights
  85. BAP1 complexes with YY1 and RBBP7 and its downstream targets in ccRCC cells
  86. Hypereosinophilic syndrome with elevated IgG4 and T-cell clonality: A report of two cases
  87. Electroacupuncture alleviates sciatic nerve injury in sciatica rats by regulating BDNF and NGF levels, myelin sheath degradation, and autophagy
  88. Polydatin prevents cholesterol gallstone formation by regulating cholesterol metabolism via PPAR-γ signaling
  89. RNF144A and RNF144B: Important molecules for health
  90. Analysis of the detection rate and related factors of thyroid nodules in the healthy population
  91. Artesunate inhibits hepatocellular carcinoma cell migration and invasion through OGA-mediated O-GlcNAcylation of ZEB1
  92. Endovascular management of post-pancreatectomy hemorrhage caused by a hepatic artery pseudoaneurysm: Case report and review of the literature
  93. Efficacy and safety of anti-PD-1/PD-L1 antibodies in patients with relapsed refractory diffuse large B-cell lymphoma: A meta-analysis
  94. SATB2 promotes humeral fracture healing in rats by activating the PI3K/AKT pathway
  95. Overexpression of the ferroptosis-related gene, NFS1, corresponds to gastric cancer growth and tumor immune infiltration
  96. Understanding risk factors and prognosis in diabetic foot ulcers
  97. Atractylenolide I alleviates the experimental allergic response in mice by suppressing TLR4/NF-kB/NLRP3 signalling
  98. FBXO31 inhibits the stemness characteristics of CD147 (+) melanoma stem cells
  99. Immune molecule diagnostics in colorectal cancer: CCL2 and CXCL11
  100. Inhibiting CXCR6 promotes senescence of activated hepatic stellate cells with limited proinflammatory SASP to attenuate hepatic fibrosis
  101. Cadmium toxicity, health risk and its remediation using low-cost biochar adsorbents
  102. Pulmonary cryptococcosis with headache as the first presentation: A case report
  103. Solitary pulmonary metastasis with cystic airspaces in colon cancer: A rare case report
  104. RUNX1 promotes denervation-induced muscle atrophy by activating the JUNB/NF-κB pathway and driving M1 macrophage polarization
  105. Morphometric analysis and immunobiological investigation of Indigofera oblongifolia on the infected lung with Plasmodium chabaudi
  106. The NuA4/TIP60 histone-modifying complex and Hr78 modulate the Lobe2 mutant eye phenotype
  107. Experimental study on salmon demineralized bone matrix loaded with recombinant human bone morphogenetic protein-2: In vitro and in vivo study
  108. A case of IgA nephropathy treated with a combination of telitacicept and half-dose glucocorticoids
  109. Analgesic and toxicological evaluation of cannabidiol-rich Moroccan Cannabis sativa L. (Khardala variety) extract: Evidence from an in vivo and in silico study
  110. Wound healing and signaling pathways
  111. Combination of immunotherapy and whole-brain radiotherapy on prognosis of patients with multiple brain metastases: A retrospective cohort study
  112. To explore the relationship between endometrial hyperemia and polycystic ovary syndrome
  113. Research progress on the impact of curcumin on immune responses in breast cancer
  114. Biogenic Cu/Ni nanotherapeutics from Descurainia sophia (L.) Webb ex Prantl seeds for the treatment of lung cancer
  115. Dapagliflozin attenuates atrial fibrosis via the HMGB1/RAGE pathway in atrial fibrillation rats
  116. Glycitein alleviates inflammation and apoptosis in keratinocytes via ROS-associated PI3K–Akt signalling pathway
  117. ADH5 inhibits proliferation but promotes EMT in non-small cell lung cancer cell through activating Smad2/Smad3
  118. Apoptotic efficacies of AgNPs formulated by Syzygium aromaticum leaf extract on 32D-FLT3-ITD human leukemia cell line with PI3K/AKT/mTOR signaling pathway
  119. Novel cuproptosis-related genes C1QBP and PFKP identified as prognostic and therapeutic targets in lung adenocarcinoma
  120. Bee venom promotes exosome secretion and alters miRNA cargo in T cells
  121. Treatment of pure red cell aplasia in a chronic kidney disease patient with roxadustat: A case report
  122. Comparative bioinformatics analysis of the Wnt pathway in breast cancer: Selection of novel biomarker panels associated with ER status
  123. Kynurenine facilitates renal cell carcinoma progression by suppressing M2 macrophage pyroptosis through inhibition of CASP1 cleavage
  124. RFX5 promotes the growth, motility, and inhibits apoptosis of gastric adenocarcinoma cells through the SIRT1/AMPK axis
  125. ALKBH5 exacerbates early cardiac damage after radiotherapy for breast cancer via m6A demethylation of TLR4
  126. Phytochemicals of Roman chamomile: Antioxidant, anti-aging, and whitening activities of distillation residues
  127. Circadian gene Cry1 inhibits the tumorigenicity of hepatocellular carcinoma by the BAX/BCL2-mediated apoptosis pathway
  128. The TNFR-RIPK1/RIPK3 signalling pathway mediates the effect of lanthanum on necroptosis of nerve cells
  129. Ecology and Environmental Science
  130. Optimization and comparative study of Bacillus consortia for cellulolytic potential and cellulase enzyme activity
  131. The complete mitochondrial genome analysis of Haemaphysalis hystricis Supino, 1897 (Ixodida: Ixodidae) and its phylogenetic implications
  132. Epidemiological characteristics and risk factors analysis of multidrug-resistant tuberculosis among tuberculosis population in Huzhou City, Eastern China
  133. Indices of human impacts on landscapes: How do they reflect the proportions of natural habitats?
  134. Genetic analysis of the Siberian flying squirrel population in the northern Changbai Mountains, Northeast China: Insights into population status and conservation
  135. Diversity and environmental drivers of Suillus communities in Pinus sylvestris var. mongolica forests of Inner Mongolia
  136. Global assessment of the fate of nitrogen deposition in forest ecosystems: Insights from 15N tracer studies
  137. Fungal and bacterial pathogenic co-infections mainly lead to the assembly of microbial community in tobacco stems
  138. Influencing of coal industry related airborne particulate matter on ocular surface tear film injury and inflammatory factor expression in Sprague-Dawley rats
  139. Agriculture
  140. Integrated analysis of transcriptome, sRNAome, and degradome involved in the drought-response of maize Zhengdan958
  141. Variation in flower frost tolerance among seven apple cultivars and transcriptome response patterns in two contrastingly frost-tolerant selected cultivars
  142. Heritability of durable resistance to stripe rust in bread wheat (Triticum aestivum L.)
  143. Animal Science
  144. Effect of sex ratio on the life history traits of an important invasive species, Spodoptera frugiperda
  145. Plant Sciences
  146. Hairpin in a haystack: In silico identification and characterization of plant-conserved microRNA in Rafflesiaceae
  147. Widely targeted metabolomics of different tissues in Rubus corchorifolius
  148. The complete chloroplast genome of Gerbera piloselloides (L.) Cass., 1820 (Carduoideae, Asteraceae) and its phylogenetic analysis
  149. Field trial to correlate mineral solubilization activity of Pseudomonas aeruginosa and biochemical content of groundnut plants
  150. Correlation analysis between semen routine parameters and sperm DNA fragmentation index in patients with semen non-liquefaction: A retrospective study
  151. Plasticity of the anatomical traits of Rhododendron L. (Ericaceae) leaves and its implications in adaptation to the plateau environment
  152. Effects of Piriformospora indica and arbuscular mycorrhizal fungus on growth and physiology of Moringa oleifera under low-temperature stress
  153. Effects of different sources of potassium fertiliser on yield, fruit quality and nutrient absorption in “Harward” kiwifruit (Actinidia deliciosa)
  154. Comparative efficiency and residue levels of spraying programs against powdery mildew in grape varieties
  155. The DREB7 transcription factor enhances salt tolerance in soybean plants under salt stress
  156. Food Science
  157. Phytochemical analysis of Stachys iva: Discovering the optimal extract conditions and its bioactive compounds
  158. Review on role of honey in disease prevention and treatment through modulation of biological activities
  159. Computational analysis of polymorphic residues in maltose and maltotriose transporters of a wild Saccharomyces cerevisiae strain
  160. Optimization of phenolic compound extraction from Tunisian squash by-products: A sustainable approach for antioxidant and antibacterial applications
  161. Liupao tea aqueous extract alleviates dextran sulfate sodium-induced ulcerative colitis in rats by modulating the gut microbiota
  162. Toxicological qualities and detoxification trends of fruit by-products for valorization: A review
  163. Polyphenolic spectrum of cornelian cherry fruits and their health-promoting effect
  164. Optimizing the encapsulation of the refined extract of squash peels for functional food applications: A sustainable approach to reduce food waste
  165. Advancements in curcuminoid formulations: An update on bioavailability enhancement strategies curcuminoid bioavailability and formulations
  166. Impact of saline sprouting on antioxidant properties and bioactive compounds in chia seeds
  167. The dilemma of food genetics and improvement
  168. Bioengineering and Biotechnology
  169. Impact of hyaluronic acid-modified hafnium metalorganic frameworks containing rhynchophylline on Alzheimer’s disease
  170. Emerging patterns in nanoparticle-based therapeutic approaches for rheumatoid arthritis: A comprehensive bibliometric and visual analysis spanning two decades
  171. Application of CRISPR/Cas gene editing for infectious disease control in poultry
  172. Preparation of hafnium nitride-coated titanium implants by magnetron sputtering technology and evaluation of their antibacterial properties and biocompatibility
  173. Preparation and characterization of lemongrass oil nanoemulsion: Antimicrobial, antibiofilm, antioxidant, and anticancer activities
  174. Corrigendum
  175. Corrigendum to “Utilization of convolutional neural networks to analyze microscopic images for high-throughput screening of mesenchymal stem cells”
  176. Corrigendum to “Effects of Ire1 gene on virulence and pathogenicity of Candida albicans”
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Heruntergeladen am 19.10.2025 von https://www.degruyterbrill.com/document/doi/10.1515/biol-2025-1166/html?srsltid=AfmBOoptgVOA2IQNv1uMqer_m4JPukFKefQOKYmZJ7GKwW-aTdRo3j24
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