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The Roles of Fibroblast Growth Factors in the Testicular Development and Tumor

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The Roles of Fibroblast Growth Factors in the Testicular Development and Tumor Hindawi Publishing Corporation Journal of Diabetes Research Volume 2013, Article ID 489095, 8 pages http://dx.doi.org/10.1155/2013/489095 Review Article The Roles of Fibroblast Growth Factors in the Testicular Development and Tumor Xin Jiang,1,2 Melissa Skibba,2 Chi Zhang,2,3 Yi Tan,2,3 Ying Xin,2,4 and Yaqin Qu1 1 The First Hospital of Jilin University, Changchun 130021, China 2 KCHRI at the Department of Pediatrics, The University of Louisville, Louisville 40202, USA 3 The Chinese-American Research Institute for Diabetic Complications, Wenzhou 325200, China 4 Key Laboratory of Pathobiology, Ministry of Education, Jilin University, 126 Xinmin Street, Changchun 130021, China Correspondence should be addressed to Ying Xin; [email protected] and Yaqin Qu; [email protected] Received 4 June 2013; Accepted 19 August 2013 Academic Editor: Subrata Chakrabarti Copyright © 2013 Xin Jiang et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Fibroblast growth factors (FGFs) are classically known as hormonal factors and recent studies have revealed that FGFs have a key role in regulating growth and development of several reproductive organs, including the testis. The testis is mainly consisted of germ cells, Sertoli cells and Leydig cells to develop and maintain the male phenotype and reproduction. This review summarizes the structure and fuctions of testis, the roles of FGFs on testicular development and potential involvement in testicular tumor and its regulatory mechanism. Among 23 members of FGFs, the FGF-1, FGF-2, FGF-4, FGF-8, FGF-9, and FGF-21 were involved and describe in details. Understanding the roles and mechanism of FGFs is the foundation to modeling testicular development and treatments in testicular disease. Therefore, in the last part, the potential therapy with FGFs for the testis of cancer and diabetes was also discussed. 1. Introduction Beside being regulated by endocrine factors, function of the testis are also mediated by paracrine pathway, including The adult mammalian testis is the important organ within the hormones, growth factors, and cytokines [5–7]. Growth fac- male reproductive system and has two major physiological tors, such as fibroblast growth factors (FGFs), are important functions: the production of spermatozoa via spermatogen- hormone-related substances that promote cell proliferation, esis and testosterone (TE) via steroidogenesis [1]. Develop- regulate tissue differentiation, and modulate organogenesis ment and maintenance of the male function of fertility are [8]. The FGF family consists of 23 members and have been all dependent on the activity of the germ cells, Sertoli cells, localizedinsomekindofcellsofthemalereproductive andLeydigcellsofthetestis.Germcellsareoneofthetwo tract and have shown to be important in regulating the types of cells in the body and the other is somatic cells. During the male development, germ cells originate from the gut of growth and the development of several reproductive organs, theembryoandmigratetothedevelopinggonads.Thenthey including the testis [9, 10]. The regulation of testicular cell can undergo mitosis or meiosis and differentiate into sperm growth is required for the maintenance of spermatogenesis [2, 3], which fuses with oocytes during fertilization. Sertoli in the adult testis. On the other hand, testicular apoptosis, cells, known as the nursery cell, affect and support germ cells as programmed cell death, also plays an important role by anchoring junctions in seminiferous epithelium. Leydig in controlling the number of male germ cells and elimi- cells secrete testosterone to stimulate Sertoli cell activity and nating defective germ cells during testicular development germ cell proliferation (spermatogenesis) [4]. Germ cells are and spermatogenesis as well as preventing testicular tumor. also affected by follicle-stimulating hormone (FSH) that is Testicular apoptosis is controlled by hormones and FGFs produced from the pituitary gland and is a key regulator in [11–13].ThereviewarticlewilloutlinetheroleofFGFson the production and function of testosterone. testicular development, proliferation and apoptosis in normal 2 Journal of Diabetes Research and pathological conditions, and the regulation of testicular stages of development: spermatogonia, primary sperma- tumor. tocyte, secondary spermatocyte, pachytene spermatocytes, round spermatids, elongating spermatids, and spermato- 2. FGFs Signaling zoon. Between these tubules is the interstitium, containing the androgen producing Leydig cells, red blood cells, and FGFs are hormonal factors that provide a range of determin- abloodvessel[1, 31]. The mature spermatozoa detaches ing and regulating functions in some tissues or organs [14, 15]. from the epithelium in the center of seminiferous tubule Although FGFs were first discovered as a mitogen on 3T3 in the process of sperm release, known as spermiation. fibroblasts, only some members of the FGF family promote Development and maintenance of the male phenotype and growth and strictly act on fibroblasts2 [ ]. Among the 23 establishment of fertility are all dependent upon the activity members of FGF family, seven subfamilies are further divided of the Sertoli cells and Leydig cells of the testis. Sertoli cell, based on phylogeny, common structural characteristics, and also known as the nursery cell, supports as many as 50 germ sequence identity rather than on functional similarity [16]. cells in the epithelium and plays a crucial role in germ cell FGF plays an important role in many processes including maturation. Sertoli cells are associated with germ cells by development, morphogenesis, angiogenesis, hematopoiesis, different anchoring junctions with the basement membrane, cell proliferation, differentiation, survival, and migration17 [ – such as cell to cell adherent junctions during different stages 19]. of their development [1]. In essence, Sertoli and germ cells, The biological significance of FGF signaling system for particularly spermatogonia, are resting on the basement human health and development is illustrated in recent obser- membrane at different stages of the seminiferous epithelial vations [20]. The correct maintenance and regulation of FGF cycle, relying on its structural and hormonal support [32]. signaling is evident from human and mouse genetic studies, Leydig cells are dependent on LH and FSH, and Sertoli cell which showed a variety of developmental disorders including can also affect Leydig cell function. Testosterone secreted by dominant skeletal diseases, infertility, and cancer if some the Leydig cells acts with FSH to stimulate Sertoli cell activity signaling mutations lead to the disruption of FGFs [10, 21, 22]. and spermatogenesis [33]. Accumulating evidence indicates the role of FGFs as a critical regulator for long-term energy balance and metab- 4. FGFs and Testicular Development olism. In particular, the endocrine-acting FGF-19 subfamily including FGF-19, FGF-21, and FGF-23 was shown to be FGFs are classically considered to be paracrine factors and are involved in glucose, lipid, bile acid, phosphate, and vitamin known for their roles in tissue patterning and organogenesis D metabolism; however, the mechanisms underlying their during embryogenesis. FGFs have been localized in many functions as metabolic regulators still need to be defined cells of the male reproductive tract and have been shown to be [13, 23, 24]. important in the regulation for the growth and development Numerous studies have focused on the expression and the of several reproductive organs including the testis [8, 34, 35]. role of FGFs, including FGF-1 (aFGF), FGF-2 (bFGF), FGF- StudieshaveshownthatFGF-9signalingcanstimulatemes- 4, FGF-5, FGF-8, FGF-9, FGF-13, FGF-18, and FGF-21 in the enchymal cell proliferation and migration of mesonephric testis [8, 10, 13, 25–27]. Moreover, the role of FGFs is different cells into the testis during development, contributing to the anditwillbeinvolvedindevelopment,cellproliferation, formation of the interstitial compartment of the testis [36]. differentiation, and apoptosis in the testis. In mammals, there is a poleward expansion of testiculo- genic programs along the anteroposterior axis of developing 3. Testicular Structure XY gonads. Sry (sex determining region of Chr Y) and its target gene, Sox9, are essential in the supporting cells The adult mammalian testis is a vital organ within the for initiating Sertoli cell differentiation and testis formation scrotum of the male reproductive system and has two major [37, 38]. Sry and Sox9 alone cannot drive center-to-pole physiological functions as mentioned above [1]. The surface expansion of testiculogenic programs at the initial phase of of testis has a transparent layer of dense connective tissue, testis differentiation. However, FGF-9 signaling is crucial for namedtesticulartunicaalbuginea,providingaprotectivebar- the maintenance of Sry and Sox9 expression, and FGF-9 can rier on the testis. The testicular tunica albuginea thicken and act as a diffusible conductor for the poleward expansion of extend into the testis which will be separated into many small tubulogenic programs and testis cord formation in develop- testis lobules, filled with testicular essence28 [ , 29]. Testicular ing XY gonads [39]. essence is mainly composed of seminiferous tubules, which Also, the role of FGF-9 on testis development is relative
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