B. Demouveaux, V. Gouyer, F. Gottrand, T. Narita, and J. Desseyn, Gel-forming mucin interactome drives mucus viscoelasticity, Adv. Colloid Interface Sci, vol.252, pp.69-82, 2018.

J. Desseyn and A. Laine, Characterization of mouse muc6 and evidence of conservation of the gel-forming mucin gene cluster between human and mouse, Genomics, vol.81, pp.36-39, 2003.
URL : https://hal.archives-ouvertes.fr/hal-02177647

J. L. Desseyn, D. Tetaert, and V. Gouyer, Architecture of the large membrane-bound mucins, Gene, vol.410, pp.215-237, 2008.

D. Dursun, M. Wang, D. Monroy, D. Q. Li, B. L. Lokeshwar et al., A mouse model of keratoconjunctivitis sicca, Invest. Ophthalmol. Vis. Sci, vol.43, pp.632-638, 2002.

A. M. Flores, S. D. Casey, C. M. Felix, P. W. Phuan, A. S. Verkman et al., Smallmolecule CFTR activators increase tear secretion and prevent experimental dry eye disease, FASEB J, vol.30, pp.1789-1797, 2016.

A. M. Floyd, X. Zhou, C. Evans, O. J. Rompala, L. Zhu et al., Mucin deficiency causes functional and structural changes of the ocular surface, PLoS One, vol.7, 2012.

I. K. Gipson, Distribution of mucins at the ocular surface, Exp. Eye Res, vol.78, pp.379-388, 2004.

I. K. Gipson, S. Spurr-michaud, and A. Tisdale, Human conjunctival goblet cells express the membrane associated mucin MUC16: Localization to mucin granules, Exp. Eye Res, vol.145, pp.230-234, 2016.

I. K. Gipson and A. S. Tisdale, Visualization of conjunctival goblet cell actin cytoskeleton and mucin content in tissue whole mounts, Exp. Eye Res, vol.65, pp.407-415, 1997.

D. Gupta, S. A. Harvey, N. Kaminski, and S. K. Swamynathan, Mouse conjunctival forniceal gene expression during postnatal development and its regulation by krüppel-like factor 4, Investig. Ophthalmol. Vis. Sci, vol.52, pp.4951-4962, 2011.

Y. Hayashi, W. W. Kao, .. Kohno, N. Nishihara-hayashi, M. Shiraishi et al., Expression patterns of sialylated epitope recognized by KL-6 monoclonal antibody in ocular surface epithelium of normals and dry eye patients, Investig. Ophthalmol. Vis. Sci, vol.45, pp.2212-2217, 2004.

J. S. House, S. Zhu, R. Ranjan, K. Linder, and R. C. Smart, C/EBPalpha and C/EBPbeta are required for Sebocyte differentiation and stratified squamous differentiation in adult mouse skin, PLoS One, vol.5, 2010.

J. Huang, L. K. Dattilo, R. Rajagopal, Y. Liu, V. Kaartinen et al., FGF-regulated BMP signaling is required for eyelid closure and to specify conjunctival epithelial cell fate, Development, vol.136, pp.1741-1750, 2009.

T. Inaba, C. Hisatsune, Y. Sasaki, Y. Ogawa, E. Ebisui et al., Mice lacking inositol 1,4,5-trisphosphate receptors exhibit dry eye, PLoS One, vol.9, 2014.

T. Inaba, Y. Tanaka, S. Tamaki, T. Ito, J. M. Ntambi et al., Compensatory increases in tear volume and mucin levels associated with meibomian gland dysfunction caused by stearoyl-CoA desaturase-1 deficiency, Sci. Rep, vol.8, 2018.

M. C. Jong, M. J. Gijbels, V. E. Dahlmans, P. J. Gorp, S. J. Koopman et al., Hyperlipidemia and cutaneous abnormalities in transgenic mice overexpressing human apolipoprotein C1, J. Clin. Invest, vol.101, pp.145-152, 1998.

R. Kardon, R. E. Price, J. Julian, E. Lagow, S. C. Tseng et al., Bacterial conjunctivitis in Muc1 null mice, Invest. Ophthalmol. Vis. Sci, vol.40, pp.1328-1335, 1999.

D. Kenchegowda, S. Swamynathan, D. Gupta, H. Wan, J. Whitsett et al., Conditional disruption of mouse Klf5 results in defective eyelids with malformed meibomian glands, abnormal cornea and loss of conjunctival goblet cells, Dev. Biol, vol.356, pp.5-18, 2011.

C. E. Kim, H. K. Kleinman, G. Sosne, G. W. Ousler, K. Kim et al., RGN-259 (thymosin ?4) improves clinically important dry eye efficacies in comparison with prescription drugs in a dry eye model, Sci. Rep, vol.8, p.10500, 2018.

C. S. Kim, K. Jo, I. S. Lee, and J. Kim, Topical application of apricot kernel extract improves dry eye symptoms in a unilateral exorbital lacrimal gland excision mouse, Nutrients, vol.8, 2016.

D. W. Kim, S. H. Lee, S. K. Ku, S. H. Cho, S. W. Cho et al., Transduced PEP-1-FK506BP ameliorates corneal injury in Botulinum toxin A-induced dry eye mouse model, BMB Rep, vol.46, pp.124-129, 2013.

T. Kojima, M. Dogru, O. M. Ibrahim, T. Nagata, K. Higa et al., The effects of 3% diquafosol sodium application on the tear functions and ocular surface of the Cu,Zn-superoxide dismutase-1 (Sod1)-knockout mice, Mol. Vis, vol.20, pp.929-938, 2014.

T. Kojima, T. H. Wakamatsu, M. Dogru, Y. Ogawa, A. Igarashi et al., Age-related dysfunction of the lacrimal gland and oxidative stress: evidence from the Cu,Zn-superoxide dismutase-1 (Sod1) knockout mice, Am. J. Pathol, vol.180, pp.1879-1896, 2012.

C. Lange, J. Fernandez, D. Shim, S. Spurr-michaud, A. Tisdale et al., Mucin gene expression is not regulated by estrogen and/or progesterone in the ocular surface epithelia of mice, Exp. Eye Res, vol.77, pp.59-68, 2003.

T. N. Lavoie, B. H. Lee, and C. Q. Nguyen, Current concepts: mouse models of Sjögren's syndrome, J. Biomed. Biotechnol, pp.1-14, 2011.

H. Lee, S. Jeon, C. E. Kim, Y. J. Park, and J. Yang, A new ophthalmic pharmaceutical formulation, topical sulglycotide, enhances the ocular mucin secretion in desiccation stressmediated dry eye disease, Investig. Ophthalmol. Vis. Sci, vol.60, pp.1076-1087, 2019.

M. J. Lee, D. H. Kim, J. S. Ryu, A. Y. Ko, J. H. Ko et al., Topical TSG-6 administration protects the ocular surface in two mouse models of inflammation-related dry eye, Invest. Ophthalmol. Vis. Sci, vol.56, pp.5175-5181, 2015.

M. J. Lee, A. Y. Ko, J. H. Ko, H. J. Lee, M. K. Kim et al., Mesenchymal stem/stromal cells protect the ocular surface by suppressing inflammation in an experimental dry eye, Mol. Ther, vol.23, pp.139-146, 2015.

M. H. Lin, F. F. Hsu, and J. H. Miner, Requirement of fatty acid transport protein 4 for development, maturation, and function of sebaceous glands in a mouse model of ichthyosis prematurity syndrome, J. Biol. Chem, vol.288, pp.3964-3976, 2013.

,

Z. Lin, X. Liu, T. Zhou, Y. Wang, L. Bai et al., A mouse dry eye model induced by topical administration of benzalkonium chloride, Mol. Vis, vol.17, pp.257-264, 2011.

Y. Machida, J. Shoji, N. Harada, and N. Inada, Two patients with dry eye disease followed up using an expression assay of ocular surface mucin, Case Rep. Ophthalmol, vol.7, pp.208-215, 2016.

F. Malet, M. Le-goff, J. Colin, C. Schweitzer, M. N. Delyfer et al., Dry eye disease in French elderly subjects: the Alienor Study, Acta Ophthalmol, vol.92, pp.429-436, 2014.

F. Mantelli, M. Massaro-giordano, I. Macchi, A. Lambiase, and S. Bonini, The cellular mechanisms of dry eye: from pathogenesis to treatment, J. Cell. Physiol, vol.228, pp.2253-2256, 2013.

C. K. Marko, B. B. Menon, G. Chen, J. A. Whitsett, H. Clevers et al., Spdef null mice lack conjunctival goblet cells and provide a model of dry eye, Am. J. Pathol, vol.183, pp.35-48, 2013.

C. K. Marko, A. S. Tisdale, S. Spurr-michaud, C. Evans, and I. K. Gipson, The ocular surface phenotype of Muc5ac and Muc5b null mice, Invest. Ophthalmol. Vis. Sci, vol.55, pp.291-300, 2014.

B. B. Menon, C. Kaiser-marko, S. Spurr-michaud, A. S. Tisdale, and I. K. Gipson, Suppression of Toll-like receptor-mediated innate immune responses at the ocular surface by the membrane-associated mucins MUC1 and MUC16, Mucosal Immunol, vol.8, pp.1000-1008, 2015.

K. Miyashita, N. Azuma, and T. Hida, Morphological and histochemical studies of goblet cells in developing human conjunctiva, Jpn. J. Ophthalmol, vol.36, pp.169-174, 1992.

M. Miyazaki, W. C. Man, and J. M. Ntambi, Targeted disruption of stearoyl-CoA desaturase1 gene in mice causes atrophy of sebaceous and meibomian glands and depletion of wax esters in the eyelid, J. Nutr, vol.131, pp.2260-2268, 2001.

I. Moon, H. G. Kang, A. Yeo, H. Noh, H. C. Kim et al., Comparison of ocular surface mucin expression after topical ophthalmic drug administration in dry eye-induced mouse model, J. Ocul. Pharmacol. Ther, vol.34, pp.612-620, 2018.

J. Murube and L. Rivas, Impression cytology on conjunctiva and cornea in dry eye patients establishes a correlation between squamous metaplasia and dry eye clinical severity, Eur. J. Ophthalmol, vol.13, pp.115-127, 2003.

A. Naito, H. Yoshida, E. Nishioka, M. Satoh, S. Azuma et al., TRAF6-deficient mice display hypohidrotic ectodermal dysplasia, Proc. Natl. Acad. Sci. U. S. A, vol.99, pp.8766-8771, 2002.

T. Ohguchi, T. Kojima, O. M. Ibrahim, T. Nagata, T. Shimizu et al., The effects of 2% rebamipide ophthalmic solution on the tear functions and ocular surface of the superoxide dismutase-1 (sod1) knockout mice, Investig. Ophthalmol. Vis. Sci, vol.54, pp.7793-7802, 2013.

,

A. Pajoohesh-ganji, S. Pal-ghosh, G. Tadvalkar, and M. A. Stepp, K14 + compound niches are present on the mouse cornea early after birth and expand after debridement wounds, Dev. Dyn, vol.245, pp.132-143, 2016.

L. T. Pallesen, L. Berglund, L. K. Rasmussen, T. E. Petersen, and J. T. Rasmussen, Isolation and characterization of MUC15, a novel cell membrane-associated mucin, Eur. J. Biochem, vol.269, pp.2755-2763, 2002.

G. J. Parfitt, D. J. Brown, and J. Jester, Transcriptome analysis of aging mouse meibomian glands, Mol. Vis, vol.22, pp.518-527, 2016.

F. Paulsen, G. Langer, W. Hoffmann, and M. Berry, Human lacrimal gland mucins, Cell Tissue Res, vol.316, pp.167-177, 2004.

S. C. Pflugfelder, Z. Liu, D. Monroy, D. Q. Li, M. E. Carvajal et al., Detection of sialomucin complex (MUC4) in human ocular surface epithelium and tear fluid, Investig. Ophthalmol. Vis. Sci, vol.41, pp.1316-1326, 2000.

M. Plikus, W. P. Wang, J. Liu, X. Wang, T. X. Jiang et al., Morpho-regulation of ectodermal organs: integument pathology and phenotypic variations in K14-Noggin engineered mice through modulation of bone morphogenic protein pathway, Am. J. Pathol, vol.164, issue.10, pp.63197-63202, 2004.

C. Portal, V. Gouyer, F. Gottrand, and J. Desseyn, Preclinical mouse model to monitor live Muc5b-producing conjunctival goblet cell density under pharmacological treatments 12, 2017.

C. Portal, V. Gouyer, M. Magnien, S. Plet, F. Gottrand et al., In vivo imaging of the Muc5b gel-forming mucin, Sci. Rep, vol.7, 2017.

L. W. Reneker, L. Wang, R. T. Irlmeier, and A. J. Huang, Fibroblast growth factor receptor 2 (FGFR2) is required for meibomian gland homeostasis in the adult mouse, Investig. Ophthalmol. Vis. Sci, vol.58, pp.2638-2646, 2017.

J. Ricciuto, S. R. Heimer, M. S. Gilmore, and P. Argüeso, Cell surface O-glycans limit Staphylococcus aureus adherence to corneal epithelial cells, Infect. Immun, vol.76, pp.5215-5220, 2008.

E. M. Rocha, A. P. Cotrim, C. Zheng, P. P. Riveros, B. J. Baum et al., Recovery of radiation-induced dry eye and corneal damage by pretreatment with adenoviral vectormediated transfer of erythropoietin to the salivary glands in mice, Hum. Gene Ther, vol.24, pp.417-423, 2013.

I. M. Shih, M. Nesbit, M. Herlyn, and R. J. Kurman, A new Mel-CAM (CD146)-specific monoclonal antibody, MN-4, on paraffin-embedded tissue, Mod. Pathol, vol.11, pp.1098-1106, 1998.

S. Shimazaki-den, M. Dogru, K. Higa, and J. Shimazaki, Symptoms, visual function, and mucin expression of eyes with tear film instability, Cornea, vol.32, pp.1211-1218, 2013.

P. Song, W. Xia, M. Wang, X. Chang, . Wang et al., Variations of dry eye disease prevalence by age, sex and geographic characteristics in China: a systematic review and meta-analysis, J. Glob. Health, vol.8, p.20503, 2018.

X. J. Song, D. Q. Li, W. Farley, L. H. Luo, R. O. Heuckeroth et al., Neurturin-deficient mice develop dry eye and keratoconjunctivitis sicca, Invest. Ophthalmol. Vis. Sci, vol.44, pp.4223-4229, 2003.

S. Spurr-michaud, P. Argüeso, and I. Gipson, Assay of mucins in human tear fluid, Exp. Eye Res, vol.84, pp.939-950, 2007.

F. Stapleton, M. Alves, V. Y. Bunya, I. Jalbert, K. Lekhanont et al., TFOS DEWS II Epidemiology Report. Ocul. Surf, vol.15, pp.334-365, 2017.

D. N. Stephens and N. A. Mcnamara, Altered mucin and glycoprotein expression in dry eye disease, Optom. Vis. Sci, vol.92, pp.931-938, 2015.

W. Stevenson, Y. Chen, S. Lee, H. S. Lee, J. Hua et al., Extraorbital lacrimal gland excision: a reproducible model of severe aqueous tear-deficient dry eye disease, Cornea, vol.33, pp.1336-1341, 2014.

O. Suwan-apichon, M. Rizen, R. Rangsin, S. Herretes, J. M. Reyes et al., Botulinum toxin B-induced mouse model of keratoconjunctivitis sicca, Invest. Ophthalmol. Vis. Sci, vol.47, pp.133-139, 2006.

S. K. Swamynathan, J. P. Katz, K. H. Kaestner, R. Ashery-padan, M. A. Crawford et al., Conditional deletion of the mouse Klf4 gene results in corneal epithelial fragility, stromal edema, and loss of conjunctival goblet cells, Mol. Cell. Biol, vol.27, pp.182-194, 2007.

M. Takahashi, N. Ishimaru, K. Yanagi, N. Haneji, I. Saito et al., High incidence of autoimmune dacryoadenitis in male non-obese diabetic (NOD) mice depending on sex steroid, Clin. Exp. Immunol, vol.109, pp.555-561, 1997.

O. Y. Tekta?, A. Yadav, F. Garreis, U. Schlötzer-schrehardt, M. Schicht et al., Characterization of the mucocutaneous junction of the human eyelid margin and meibomian glands with different biomarkers, Ann. Anat, vol.194, pp.436-445, 2012.

D. J. Thornton, K. Rousseau, and M. A. Mcguckin, Structure and function of the polymeric mucins in airways mucus, Annu. Rev. Physiol, vol.70, pp.459-486, 2008.

,

P. M. Treuting, R. Wong, D. C. Tu, and I. Phan, Special Senses, Comparative Anatomy and Histology, pp.395-418, 2012.

C. Tsau, M. Ito, A. Gromova, M. P. Hoffman, R. Meech et al., Barx2 and Fgf10 regulate ocular glands branching morphogenesis by controlling extracellular matrix remodeling, Development, vol.138, pp.3307-3317, 2011.

M. Uchino, N. Yokoi, Y. Uchino, M. Dogru, M. Kawashima et al., Prevalence of dry eye disease and its risk factors in visual display terminal users: The Osaka study, Am. J. Ophthalmol, vol.156, pp.759-766, 2013.

Y. Uchino, T. Kawakita, T. Ishii, N. Ishii, and K. Tsubota, A new mouse model of dry eye disease: oxidative stress affects functional decline in the lacrimal gland, Cornea, vol.31, issue.1, pp.63-67, 2012.

Y. Uchino, M. Uchino, N. Yokoi, M. Dogru, M. Kawashima et al., Alteration of tear mucin 5AC in office workers using visual display terminals: The Osaka study, JAMA Ophthalmol, vol.132, pp.985-992, 2014.

M. Uhlén, L. Fagerberg, B. M. Hallström, C. Lindskog, P. Oksvold et al., Tissue-based map of the human proteome, vol.347, p.1260419, 2015.

A. Ushio, R. Arakaki, H. Eguchi, F. Hotta, A. Yamada et al., Pathological analysis of ocular lesions in a murine model of Sjögren's syndrome, Int. J. Mol. Sci, vol.18, 1209.

T. Vijmasi, F. Y. Chen, Y. T. Chen, M. Gallup, and N. Mcnamara, Topical administration of interleukin-1 receptor antagonist as a therapy for aqueous-deficient dry eye in autoimmune disease, Mol. Vis, vol.19, pp.1957-1965, 2013.

I. Wang, C. Yu, and F. Hu, Alteration of ocular surface mucins in MUC5AC-DTA transgenic mice, Mol. Vis, vol.15, pp.108-119, 2009.

Y. C. Wang, S. Li, X. Chen, B. Ma, H. He et al., Meibomian gland absence related dry eye in ectodysplasin a mutant mice, Am. J. Pathol, vol.186, pp.32-42, 2016.

M. D. Willcox, P. Argüeso, G. A. Georgiev, J. M. Holopainen, G. W. Laurie et al., TFOS DEWS II Tear Film Report. Ocul. Surf, vol.15, pp.366-403, 2017.

A. M. Woodward and P. Argüeso, Expression analysis of the transmembrane mucin MUC20 in human corneal and conjunctival epithelia, Investig. Ophthalmol. Vis. Sci, vol.55, pp.6132-6138, 2014.

H. Xu, Y. Zhao, J. Li, M. Wang, F. Lian et al., Loss of NHE8 expression impairs ocular surface function in mice, Am. J. Physiol. Cell Physiol, vol.308, pp.79-87, 2015.

H. Yagyu, T. Kitamine, J. Osuga, R. Tozawa, Z. Chen et al., Absence of ACAT-1 attenuates atherosclerosis but causes dry eye and cutaneous xanthomatosis in mice with congenital hyperlipidemia, J. Biol. Chem, vol.275, pp.21324-21330, 2000.

D. F. Yu, Y. Chen, J. M. Han, H. Zhang, X. P. Chen et al., MUC19 expression in human ocular surface and lacrimal gland and its alteration in Sjögren syndrome patients, Exp. Eye Res, vol.86, pp.403-411, 2008.

,

X. Zhang, W. Chen, C. S. De-paiva, R. M. Corrales, E. A. Volpe et al., Interferon-? exacerbates dry eye-induced apoptosis in conjunctiva through dual apoptotic pathways, Investig. Ophthalmol. Vis. Sci, vol.52, pp.6279-6285, 2011.

X. Zhang, C. S. De-paiva, Z. Su, E. A. Volpe, D. Li et al., Topical interferon-gamma neutralization prevents conjunctival goblet cell loss in experimental murine dry eye, Exp. Eye Res, vol.118, pp.117-124, 2014.

Z. Zhang, W. Z. Yang, Z. Z. Zhu, Q. Q. Hu, Y. F. Chen et al., Therapeutic effects of topical doxycycline in a benzalkonium chloride-induced mouse dry eye model, Investig. Ophthalmol. Vis. Sci, vol.55, pp.2963-2974, 2014.

A. ?iniauskait?, S. Ragauskas, J. J. Hakkarainen, C. C. Rich, R. Baumgartner et al., Efficacy of trabodenoson in a mouse keratoconjunctivitis sicca (KCS) model for dry-eye syndrome, Investig. Ophthalmol. Vis. Sci, vol.59, pp.3088-3093, 2018.

D. Zoukhri, E. Macari, and C. L. Kublin, A single injection of interleukin-1 induces reversible aqueous-tear deficiency, lacrimal gland inflammation, and acinar and ductal cell proliferation, Exp. Eye Res, vol.84, pp.894-904, 2007.