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Current Progress in Understanding the Molecular Pathogenesis of Burn Scar Contracture Jianglin Tan1 and Jun Wu2*

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Current Progress in Understanding the Molecular Pathogenesis of Burn Scar Contracture Jianglin Tan1 and Jun Wu2* Tan and Wu Burns & Trauma (2017) 5:14 DOI 10.1186/s41038-017-0080-1 REVIEW Open Access Current progress in understanding the molecular pathogenesis of burn scar contracture Jianglin Tan1 and Jun Wu2* Abstract Abnormal wound healing is likely to induce scar formation, leading to dysfunction, deformity, and psychological trauma in burn patients. Despite the advancement of medical care treatment, scar contracture in burn patients remains a challenge. Myofibroblasts play a key role in scar contracture. It has been demonstrated that myofibroblasts, as well as inflammatory cells, fibroblasts, endothelial cells, and epithelial cells, secrete transforming growth factor-β1(TGF-β1) and other cytokines, which can promote persistent myofibroblast activation via a positive regulation loop. In addition to the cellular contribution, the microenvironments, including the mechanical tension and integrin family, are also involved in scar contracture. Most recently, eukaryotic initiation factor 6 (eIF6), an upstream regulator of TGF-β1, has been demonstrated to be involved in myofibroblast differentiation and contraction in both in vitro fibroblast-populated collagen lattice (FPCL) and in vivo external mechanical stretch models. Moreover, the data showed that P311 could induce the transdifferentiation of epidermal stem cells to myofibroblasts by upregulating TGF-β1 expression, which mediated myofibroblast contraction. In this review, we briefly described the most current progress on the biological function of myofibroblasts in scar contracture and subsequently summarized the molecular events that initiated contracture. This would help us better understand the molecular basis of scar contracture as well as to find a comprehensive strategy for preventing/managing scar contracture. Keywords: Scar, Contracture, Burn, Molecular pathogenesis Background which meant 33% of patients had dysfunction in their It is commonly accepted that scars are a pathologic joints after burn injuries [5]. wound healing response to burn injuries, traumatic in- Wound healing proceeds through three overlapping juries, and surgeries. Hypertrophic scars and keloids, stages. The inflammatory stage is triggered by injury, which only occur in humans, present with exuberant wherein platelets, neutrophils, and macrophages release in- scar formation [1]. Although these disorders do not pose flammatory mediators and cytokines that participate in the a health risk, the scar contracture resulting in dys- recruitment of inflammatory cells, fibroblasts, endothelial function and deformity remains a challenge in the cells, and epithelial cells. The proliferative stage involves clinic [2, 3]. The management of the scar contracture, fibroblast activation, myofibroblast differentiation, and such as surgical intervention, drugs, silicone materials, extracellular matrix (ECM) deposition. In this phase, the pressure therapy, splinting, lasers, and radiation, is myofibroblasts have acquired contractile properties that used to control scar formation and contracture, but it can contract the wound and promote re-epithelialization. is still far from achieving our expected outcomes [4]. The third healing stage is matrix remodeling, including scar Schneider found that 620 of the 1865 analyzed adult tissue remodeling. During this stage, the persistent activa- burn patients developed at least one joint contracture, tion of myofibroblasts, imbalance of deposition and degrad- ation of ECM, and poor arrangement of newly formed * Correspondence: [email protected] fibers can lead to scar formation. 2Department of Burns, First Affiliated Hospital of Sun Yat-sen University, Myofibroblasts, a type of cell differentiated from quies- Guangzhou 510080, China cent fibroblasts and other cells, have been demonstrated to Full list of author information is available at the end of the article © The Author(s). 2017 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Tan and Wu Burns & Trauma (2017) 5:14 Page 2 of 6 play an essential role in the induction and maintenance of Smad complex translocates into the nucleus to enhance scar contracture. In normal acute wound healing, the myo- gene transcription via cooperation with DNA transcription fibroblasts are temporally limited and cleared by apoptosis factors. Additionally, it induces overproduction of α-SMA, in the third healing phase when the tissues are repaired. collagen I, collagen III, fibronectin (FN), and other However, in hypertrophic scars and keloids, these myofi- cytokines [16]. TGF-β1 can also reduce matrix metallopro- broblasts persist at a high number for long periods and pro- teinases (MMPs) activity via decreasing proteases, such as mote the synthesis of α-smooth muscle actin (α-SMA), tissue inhibitor of metalloproteinases (TIMPs) I and II [17]. transforming growth factor-β1(TGF-β1), and other growth The TGF-β/Smads signaling pathway is a positive autocrine factors,andtheyhavesustained contractile ability via the loop in both hypertrophic scar and keloid formation, which TGF-β1 positive loop [6]. then increases the stress fiber stabilization and stiffness of the microenvironment. Importantly, TGF-β1 is an inducer of myofibroblast dif- Review ferentiation, which is considered a potential therapeutic The origins and characteristics of myofibroblasts target for hypertrophic scars and keloids. It has been re- In the inflammatory stage, fibrocytes and fibroblasts are ported that peroxisome proliferator-activated receptor-γ believed to be activated in response to inflammatory fac- (PPARγ) ligands, 15d-PGJ2 and GW7845, could inhibit tors; they then migrate to the location of injury based on a the expression and phosphorylation of TGF-β1/Smads chemoattractant gradient and differentiate into myofibro- [18]. Either disruption or neutralization of TGF-β/Smads blasts. In general, most myofibroblasts are derived from signaling by botulinum toxin type A, tetrandrine, baica- fibroblast differentiation around the local wound area [7]. lein, loureirin B, or the Uighur medicine ASMq can de- In addition, other myofibroblasts may originate from peri- crease the myofibroblast properties [19]. In addition, cytes [8], smooth muscle cells from the vasculature [9], TGF-β1 could also promote myofibroblast differentiation fibrocytes from bone marrow-derived peripheral blood independent of Smads signaling and instead act via the [10], epithelial cells through the epithelial-mesenchymal wnt, p38, and PI3K/PKA2 signaling pathways [20]. P311, transition (EMT) [11], epidermal stem cells [12, 13], local identified by suppressive subtraction hybridization as mesenchymal stem cells, and bone marrow-derived mes- potentially involved in smooth muscle (SM) myogenesis, enchymal stem cells [14]. was highly expressed in hypertrophic scars and could The myofibroblast cell is a phenotypically intermediate induce a TGF-β1-independent, nonfibrogenic myofibro- cell type between fibroblasts and smooth muscle cells. blast phenotype [21, 22]. Furthermore, in the renal fibrosis The myofibroblasts exhibit the characteristics of smooth model, it was found that overexpression of P311 was muscle cells, containing high-contractile stress fibers. concurrent with the expression of α-SMA and TGF-β1via The stress fibers consist of α-SMA protein, which can be the TGF-β1/Smad signaling pathway [23]. Eukaryotic initi- used to differentiate between myofibroblasts and fibro- ation factor 6 (eIF6), acting as a key binding protein of blasts in tissues. However, it remains difficult and P311 [24], has recently been demonstrated by our team as complicated to discriminate between myofibroblasts and a novel upstream regulator to inhibit myofibroblast differ- other contractile cells, such as smooth muscle cells, peri- entiation at the TGF-β1 transcription level via H2A.Z cytes, and myoepithelial cells, even if the smooth muscle occupancy and Sp1 recruitment. Additionally, there is cells express smooth muscle myosin heavy chain, h-cal- downregulation of α-SMA and collagen I expression desmon (H-CAD), smoothelin, and the muscle inter- [25]. In addition, our next study demonstrated that mediate filament protein, desmin, which are absent from eIF6-mediated TGF-β1 can also be regulated by exter- myofibroblasts [15]. There are a lot of cytokines and nal mechanical stretch [26]. mechanical microenvironment factors that contribute to scar contracture (Fig. 1). Positive growth factors and cytokines In addition, many other growth factors show positive roles The cytokine contribution to scar contracture in myofibroblast differentiation, such as connective tissue Transforming growth factor-β family growth factor (CTGF), platelet-derived growth factor It has been demonstrated that TGF-β1isoneofthemost (PDGF), insulin growth factor (IGF), and vascular endothe- important factors controlling myofibroblast differentiation lial growth factor (VEGF) [27, 28]. CTGF could synergize and function. TGF-β is found in all tissues, and it consists the action of TGF-β, promoting ECM production and scar of three isoforms,
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