{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T14:46:55Z","timestamp":1791557215331,"version":"4.3.4"},"reference-count":63,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2025,6,4]],"date-time":"2025-06-04T00:00:00Z","timestamp":1748995200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32072954"],"award-info":[{"award-number":["32072954"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32270555"],"award-info":[{"award-number":["32270555"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Hedgehog (HH) signaling plays important roles in the development of the nervous system (Sonic hedgehog), bone, cartilage (Indian Hedgehog) and testis (Desert Hedgehog). Research on HH and testes has mostly been conducted in HH-knockout mice and rats, etc. The relationship between HH and cellular junctions has mostly been found in the nervous system and intestine. However, few research studies concerning the link between HH signaling and cell junctions in testis function have been reported. We identified the members of HH signaling that are involved in Eriocheir sinensis testes: HH, Smoothen, Patched, Kif27 and Ci. HH has only one homolog in E. sinensis and is expressed in several types of germ cells in the testes. We found that Kif27 colocalized with Ci in the testes. The knockdown of HH induced enlarged interstitial spaces of the seminiferous tubules. A biotin\u2013streptavidin immunofluorescence experiment indicated that the hemolymph\u2013testis barrier (HTB) was disrupted. Western blot results showed that pinin, HH signaling and cell proliferation- and apoptosis-related protein levels were downregulated. Further immunofluorescent results showed the dislocation of several junction proteins, the abnormality of F-actin and the slowdown of germ cell proliferation and apoptosis. While \u03b2-catenin entered the spermatocyte nucleus, it did not activate Wnt-\u03b2-catenin signaling, which indicated that the disturbance of the cell cycle in germ cells was not caused by Wnt-\u03b2-catenin signaling. In summary, HH signaling plays some roles beyond our understanding in the regulation of the HTB and the germ cell cycle in E. sinensis testes.<\/jats:p>","DOI":"10.3390\/ijms26115378","type":"journal-article","created":{"date-parts":[[2025,6,4]],"date-time":"2025-06-04T06:00:49Z","timestamp":1749016849000},"page":"5378","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Hedgehog Signaling Functions in Spermatogenesis and Keeping Hemolymph\u2013Testis Barrier Stability in Eriocheir sinensis"],"prefix":"10.3390","volume":"26","author":[{"given":"Jun-Jie","family":"Yu","sequence":"first","affiliation":[{"name":"The Sperm Laboratory, College of Life Sciences, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hong-Yu","family":"Qi","sequence":"additional","affiliation":[{"name":"The Sperm Laboratory, College of Life Sciences, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhan","family":"Zhao","sequence":"additional","affiliation":[{"name":"The Sperm Laboratory, College of Life Sciences, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu","family":"Yang","sequence":"additional","affiliation":[{"name":"The Sperm Laboratory, College of Life Sciences, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuang-Yi","family":"Zhang","sequence":"additional","affiliation":[{"name":"The Sperm Laboratory, College of Life Sciences, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fu-Qing","family":"Tan","sequence":"additional","affiliation":[{"name":"School of Medicine, Zhejiang University, Hangzhou 310003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5695-0150","authenticated-orcid":false,"given":"Wan-Xi","family":"Yang","sequence":"additional","affiliation":[{"name":"The Sperm Laboratory, College of Life Sciences, Zhejiang University, Hangzhou 310058, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,6,4]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1275","DOI":"10.1038\/s41467-017-01364-z","article-title":"Hedgehog mediated degradation of Ihog adhesion proteins modulates cell segregation in Drosophila wing imaginal discs","volume":"8","author":"Hsia","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/S0960-9822(99)80018-9","article-title":"Sonic hedgehog signaling by the patched-smoothened receptor complex","volume":"9","author":"Murone","year":"1999","journal-title":"Curr. Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1998","DOI":"10.1016\/j.cub.2003.10.004","article-title":"Identification of a Functional Interaction between the Transmembrane Protein Smoothened and the Kinesin-Related Protein Costal2","volume":"13","author":"Ogden","year":"2003","journal-title":"Curr. Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/j.devcel.2005.01.001","article-title":"Hedgehog-Regulated Costal2-Kinase Complexes Control Phosphorylation and Proteolytic Processing of Cubitus Interruptus","volume":"8","author":"Zhang","year":"2005","journal-title":"Dev. Cell"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"a117","DOI":"10.1126\/scisignal.2005608","article-title":"The PPFIA1-PP2A protein complex promotes trafficking of Kif7 to the ciliary tip and Hedgehog signaling","volume":"7","author":"Liu","year":"2014","journal-title":"Sci. Signal."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1875","DOI":"10.1002\/ajmg.a.32909","article-title":"Hedgehog signaling update","volume":"152A","author":"Cohen","year":"2010","journal-title":"Am. J. Med. Genet. A"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1038\/ncb2988","article-title":"The kinesin-4 protein Kif7 regulates mammalian Hedgehog signalling by organizing the cilium tip compartment","volume":"16","author":"He","year":"2014","journal-title":"Nat. Cell Biol."},{"key":"ref_8","first-page":"1875","article-title":"KIF27 is one of orthologs for Drosophila Costal-2","volume":"25","author":"Katoh","year":"2004","journal-title":"Int. J. Oncol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1159\/000518308","article-title":"The Desert Hedgehog Signalling Pathway in Human Gonadal Development and Differences of Sex Development","volume":"16","author":"Pachernegg","year":"2022","journal-title":"Sex. Dev."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1007\/s11692-021-09542-0","article-title":"Enhanced Negative Regulation of the DHH Signaling Pathway as a Potential Mechanism of Ascrotal Testes in Laurasiatherians","volume":"48","author":"Chai","year":"2021","journal-title":"Evol. Biol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1530\/REP-21-0468","article-title":"Desert hedgehog mediates the proliferation of medaka spermatogonia through Smoothened signaling","volume":"163","author":"Zhao","year":"2022","journal-title":"Reproduction"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1002\/2211-5463.13600","article-title":"Single-cell transcriptome analysis of male chicken germ cells reveals changes in signaling pathway-related gene expression profiles during mitotic arrest","volume":"13","author":"Choi","year":"2023","journal-title":"FEBS Open Bio."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"119303","DOI":"10.1016\/j.gep.2022.119303","article-title":"Denovo RNA-Seq analysis of ovary and testis reveals potential differentially expressed transcripts associated with gonadal unsynchronization development in Onychostoma macrolepis","volume":"47","author":"Cao","year":"2023","journal-title":"Gene Expr. Patterns"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1038\/s41419-022-04658-3","article-title":"Leptin secreted from testicular microenvironment modulates hedgehog signaling to augment the endogenous function of Leydig cells","volume":"13","author":"Arora","year":"2022","journal-title":"Cell Death Dis."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e23739","DOI":"10.1002\/mrd.23739","article-title":"Fetal Leydig cells: What we know and what we don\u2019t","volume":"91","author":"Jiang","year":"2024","journal-title":"Mol. Reprod. Dev."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Nygaard, M.B., Almstrup, K., Lindb\u00e6k, L., Christensen, S.T., and Svingen, T. (2015). Cell context-specific expression of primary cilia in the human testis and ciliary coordination of Hedgehog signalling in mouse Leydig cells. Sci. Rep., 5.","DOI":"10.1038\/srep10364"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1006\/dbio.2002.0663","article-title":"Disruption of Testis Cords by Cyclopamine or Forskolin Reveals Independent Cellular Pathways in Testis Organogenesis","volume":"246","author":"Yao","year":"2002","journal-title":"Dev. Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.anireprosci.2017.02.002","article-title":"Hedgehog-Gli1 signaling regelates differentiation of chicken (Gallus gallus) embryonic stem cells to male germ cells","volume":"182","author":"Li","year":"2017","journal-title":"Anim. Reprod. Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1095\/biolreprod.108.067926","article-title":"Identification of Hedgehog Signaling Outcomes in Mouse Testis Development Using a Hanging Drop-Culture System1","volume":"80","author":"Szczepny","year":"2009","journal-title":"Biol. Reprod."},{"key":"ref_20","first-page":"1655","article-title":"SUMO regulates somatic cyst stem cells maintenance and directly targets hedgehog pathway in adult Drosophila testis","volume":"143","author":"Lv","year":"2016","journal-title":"Development"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1530\/REP-11-0110","article-title":"Hedgehog signalling promotes germ cell survival in the rat testis","volume":"142","author":"Saario","year":"2011","journal-title":"Reproduction"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Kothandapani, A., Lewis, S.R., Noel, J.L., Zacharski, A., Krellwitz, K., Baines, A., Winske, S., Vezina, C.M., Kaftanovskaya, E.M., and Agoulnik, A.I. (2020). GLI3 resides at the intersection of hedgehog and androgen action to promote male sex differentiation. PLoS Genet., 16.","DOI":"10.1371\/journal.pgen.1008810"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1016\/j.crtox.2020.10.001","article-title":"Hedgehog signal disruption, gonadal dysgenesis and reproductive disorders: Is there a link to endocrine disrupting chemicals?","volume":"1","author":"Johansson","year":"2020","journal-title":"Curr. Res. Toxicol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1825","DOI":"10.1095\/biolreprod63.6.1825","article-title":"Desert hedgehog (Dhh) gene is required in the mouse testis for formation of adult-type Leydig cells and normal development of peritubular cells and seminiferous tubules","volume":"63","author":"Clark","year":"2000","journal-title":"Biol. Reprod."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1530\/REP-10-0006","article-title":"A missense mutation of the Dhh gene is associated with male pseudohermaphroditic rats showing impaired Leydig cell development","volume":"141","author":"Kawai","year":"2011","journal-title":"Reproduction"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Wei, J., Wu, J., Ru, W., Chen, G., Gao, L., and Tang, D. (2022). Novel compound heterozygous mutations in the desert hedgehog (DHH) gene in cases of siblings with 46,XY disorders of sexual development. BMC Med. Genom., 15.","DOI":"10.1186\/s12920-022-01334-5"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Baz-Red\u00f3n, N., Soler-Colomer, L., Fern\u00e1ndez-Cancio, M., Benito-Sanz, S., Garrido, M., Molin\u00e9, T., Clemente, M., Camats-Tarruella, N., and Yeste, D. (2022). Novel variant in HHAT as a cause of different sex development with partial gonadal dysgenesis associated with microcephaly, eye defects, and distal phalangeal hypoplasia of both thumbs: Case report. Front. Endocrinol., 13.","DOI":"10.3389\/fendo.2022.957969"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1871","DOI":"10.1007\/s10815-021-02140-1","article-title":"Mutations in the desert hedgehog (DHH) gene in the disorders of sexual differentiation and male infertility","volume":"38","author":"Mehta","year":"2021","journal-title":"J. Assist. Reprod. Gen."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"3704","DOI":"10.1210\/en.2006-1731","article-title":"Distinct Roles for Steroidogenic factor 1 and Desert hedgehog Pathways in Fetal and Adult Leydig Cell Development","volume":"148","author":"Park","year":"2007","journal-title":"Endocrinology"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2657","DOI":"10.1096\/fj.12-225060","article-title":"Dynamic changes in fetal Leydig cell populations influence adult Leydig cell populations in mice","volume":"27","author":"Barsoum","year":"2013","journal-title":"FASEB J."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1663","DOI":"10.1095\/biolreprod65.6.1663","article-title":"GLI1 Localization in the Germinal Epithelial Cells Alternates Between Cytoplasm and Nucleus: Upregulation in Transgenic Mice Blocks Spermatogenesis in Pachytene1","volume":"65","author":"Kroft","year":"2001","journal-title":"Biol. Reprod."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"772","DOI":"10.1093\/biolre\/ioad090","article-title":"Smoothened mediates medaka spermatogonia proliferation via Gli1\u2013Rgcc\u2013Cdk1 axis","volume":"109","author":"Zhao","year":"2023","journal-title":"Biol. Reprod."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"108981","DOI":"10.1016\/j.fsi.2023.108981","article-title":"Hedgehog signaling is essential in the regulation of limb regeneration in the Chinese mitten crab, Eriocheir sinensis","volume":"140","author":"Li","year":"2023","journal-title":"Fish Shellfish Immun."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1136\/svn-2022-001907","article-title":"Oestrogen ameliorates blood-brain barrier damage after experimental subarachnoid haemorrhage via the SHH pathway in male rats","volume":"8","author":"Zhang","year":"2023","journal-title":"Stroke Vasc. Neurol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.ydbio.2007.07.008","article-title":"Cross regulation of intercellular gap junction communication and paracrine signaling pathways during organogenesis in Drosophila","volume":"310","author":"Lechner","year":"2007","journal-title":"Dev. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"720","DOI":"10.1007\/s12975-019-00771-2","article-title":"Activation of the Hedgehog Pathway Promotes Recovery of Neurological Function After Traumatic Brain Injury by Protecting the Neurovascular Unit","volume":"11","author":"Wu","year":"2020","journal-title":"Transl. Stroke Res."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Xing, G., Zhao, T., Zhang, X., Li, H., Li, X., Cui, P., Li, M., Li, D., Zhang, N., and Jiang, W. (2020). Astrocytic Sonic Hedgehog Alleviates Intracerebral Hemorrhagic Brain Injury via Modulation of Blood-Brain Barrier Integrity. Front. Cell. Neurosci., 14.","DOI":"10.3389\/fncel.2020.575690"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1111\/jcmm.13311","article-title":"Ectodysplasin A regulates epithelial barrier function through sonic hedgehog signalling pathway","volume":"22","author":"Li","year":"2018","journal-title":"J. Cell. Mol. Med."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4703","DOI":"10.1242\/dev.126.21.4703","article-title":"Gap junction-mediated transfer of left-right patterning signals in the early chick blastoderm is upstream of Shh asymmetry in the node","volume":"126","author":"Levin","year":"1999","journal-title":"Development"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"107801","DOI":"10.1016\/j.celrep.2020.107801","article-title":"H1N1 Influenza Virus Cross-Activates Gli1 to Disrupt the Intercellular Junctions of Alveolar Epithelial Cells","volume":"31","author":"Ruan","year":"2020","journal-title":"Cell Rep."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"996","DOI":"10.1016\/j.bbrc.2005.11.106","article-title":"Hedgehog peptide promotes cell polarization and lumen formation in developing mouse submandibular gland","volume":"339","author":"Hashizume","year":"2006","journal-title":"Biochem. Bioph. Res. Co."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.1091\/mbc.e10-08-0675","article-title":"N-cadherin-mediated cell adhesion restricts cell proliferation in the dorsal neural tube","volume":"22","author":"Chalasani","year":"2011","journal-title":"Mol. Biol. Cell"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Liu, W., Ruan, T., Ji, X., Ran, D., Sun, J., Shi, H., Prinz, R.A., Sun, J., Pan, Z., and Jiao, X. (2020). The Gli1-Snail axis contributes to Salmonella Typhimurium-induced disruption of intercellular junctions of intestinal epithelial cells. Cell. Microbiol., 22.","DOI":"10.1111\/cmi.13211"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Singh, V.B., Singh, M.V., Gorantla, S., Poluektova, L.Y., and Maggirwar, S.B. (2016). Smoothened Agonist Reduces Human Immunodeficiency Virus Type-1-Induced Blood-Brain Barrier Breakdown in Humanized Mice. Sci. Rep., 6.","DOI":"10.1038\/srep26876"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1723","DOI":"10.1126\/science.1206936","article-title":"The Hedgehog pathway promotes blood-brain barrier integrity and CNS immune quiescence","volume":"334","author":"Alvarez","year":"2011","journal-title":"Science"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1609","DOI":"10.1126\/science.1121449","article-title":"alpha E-catenin controls cerebral cortical size by regulating the hedgehog signaling pathway","volume":"311","author":"Lien","year":"2006","journal-title":"Science"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1158\/0008-5472.CAN-10-4621","article-title":"GLI1 Inhibition Promotes Epithelial-to-Mesenchymal Transition in Pancreatic Cancer Cells","volume":"72","author":"Joost","year":"2012","journal-title":"Cancer Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"4489","DOI":"10.1038\/sj.onc.1210241","article-title":"Gli1 acts through Snail and E-cadherin to promote nuclear signaling by b-catenin","volume":"26","author":"Li","year":"2007","journal-title":"Oncogene"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1038\/onc.2010.459","article-title":"GLI1 facilitates the migration and invasion of pancreatic cancer cells through MUC5AC-mediated attenuation of E-cadherin","volume":"30","author":"Inaguma","year":"2011","journal-title":"Oncogene"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"3548","DOI":"10.1111\/bph.14423","article-title":"Modulation of sonic hedgehog-induced mouse embryonic stem cell behaviours through E-cadherin expression and integrin \u03b21-dependent F-actin formation","volume":"175","author":"Oh","year":"2018","journal-title":"Brit. J. Pharmacol."},{"key":"ref_51","first-page":"G1252","article-title":"Hedgehog signaling regulates E-cadherin expression for the maintenance of the actin cytoskeleton and tight junctions","volume":"299","author":"Xiao","year":"2010","journal-title":"Am. J. Physiol.-Gastr. L."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Min, M., Song, T., Sun, M., Wang, T., Tan, J., and Zhang, J. (2022). Dhh signaling pathway regulates reconstruction of seminiferous tubule-like structure. Reprod. Biol., 22.","DOI":"10.1016\/j.repbio.2022.100684"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1007\/BF00744995","article-title":"Laminin ultrastructural immunolocalization in rat testis during ontogenesis","volume":"95","author":"El","year":"1991","journal-title":"Histochemistry"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1124\/pr.110.002790","article-title":"The Blood-Testis Barrier and Its Implications for Male Contraception","volume":"64","author":"Cheng","year":"2011","journal-title":"Pharmacol. Rev."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1007\/s00441-022-03680-3","article-title":"mTORC1\/C2 regulate spermatogenesis in Eriocheir sinensis via alterations in the actin filament network and cell junctions","volume":"390","author":"Li","year":"2022","journal-title":"Cell Tissue Res."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Liu, D., Li, Z., Zhao, Y., Wang, L., Qi, H., Zhao, Z., Tan, F., and Yang, W. (2023). Es-\u03b2-CATENIN affects the hemolymph-testes barrier in Eriocheir sinensis by disrupting cell junctions and cytoskeleton. Int. J. Biol. Macromol., 242.","DOI":"10.1016\/j.ijbiomac.2023.124867"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Qi, H., Li, Z., Wang, L., Zhao, Z., Wang, J., Tan, F., and Yang, W. (2023). Myosin VI stabilizes intercellular junctions in the testis through the LHR and MAPK signalling pathway during spermatogenesis in Eriocheir sinensis. Int. J. Biol. Macromol., 248.","DOI":"10.1016\/j.ijbiomac.2023.125842"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Zhao, Z., Qi, H., Li, Z., Wang, L., Wang, J., Tan, F., and Yang, W. (2024). Vangl2 regulates intercellular junctions by remodeling actin-based cytoskeleton through the Rock signaling pathway during spermatogenesis in Eriocheir sinensis. Int. J. Biol. Macromol., 279.","DOI":"10.1016\/j.ijbiomac.2024.135264"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1007\/s00441-014-2092-5","article-title":"Morphological alterations of all stages of spermatogenesis and acrosome reaction in Chinese mitten crab Eriocheir sinensis","volume":"360","author":"Wang","year":"2015","journal-title":"Cell Tissue Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"121952","DOI":"10.1016\/j.envpol.2023.121952","article-title":"TiO2 nanoparticles affect spermatogenesis and adhesion junctions via the ROS-mediated mTOR signalling pathway in Eriocheir sinensis testes","volume":"331","author":"Wang","year":"2023","journal-title":"Environ. Pollut."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1848212","DOI":"10.1080\/21688370.2020.1848212","article-title":"Cell-cell junctions: Structure and regulation in physiology and pathology","volume":"9","author":"Adil","year":"2021","journal-title":"Tissue Barriers"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Leu, S. (2021). The role and regulation of Pnn in proliferative and non-dividing cells: Form embryogenesis to pathogenesis. Biochem. Pharmacol., 192.","DOI":"10.1016\/j.bcp.2021.114672"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1111\/j.2047-2927.2014.00187.x","article-title":"Dynamic Hedgehog signalling pathway activity in germline stem cells","volume":"2","author":"Sahin","year":"2014","journal-title":"Andrology"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/11\/5378\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:46:19Z","timestamp":1760031979000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/11\/5378"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,6,4]]},"references-count":63,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2025,6]]}},"alternative-id":["ijms26115378"],"URL":"https:\/\/doi.org\/10.3390\/ijms26115378","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,6,4]]}}}