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Presumably, this occurs as a response to tissue dissection and organ culture, which is well-known to elicit an immediate wound healing response in the epithelium. The latter rapidly starts to migrate over the wound edge in an attempt to enwrap the exposed skin mesenchyme epiboly phenomenon Stenn, ; Brown et al. This constitutive up-regulation of K6 in organ-cultured normal human skin may greatly heighten the sensitivity to K6 expression-modulatory compounds, such as CB ligands, thus making human skin organ culture a particularly sensitive and instructive tool for clinically relevant keratin research.

At the same time, however, caution is advised in extrapolating from these observation made in what essentially reflects a wound healing milieu to healthy, unmanipulated human skin in vivo. Our results suggest that cannabinoids and their receptors constitute a novel, clinically relevant control element of human K6 and K16 expression. The authors thank Motoko Sugawara for her excellent technical assistance, and Dr.

Stephan Tiede for professional advice. The generous professional support of Prof. Masamitsu Ishii and Prof. Hiromi Kobayashi, Osaka, for this work is also gratefully appreciated. Wolfgang Funk is gratefully acknowledged for harvesting human skin samples for the organ culture. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Author Contributions Yuval Ramot and Koji Sugawara conceived and designed the experiments, performed the experiments, analyzed the data, wrote the paper.

Ralf Paus conceived, designed, and supervised the experiments, and wrote the paper. Human Ethics The following information was supplied relating to ethical approvals i. Institutional review board ethics committee of the University of Luebeck, reference number National Center for Biotechnology Information , U. Journal List Peerj v. Published online Feb Author information Article notes Copyright and License information Disclaimer.

Corresponding author. Ralf Paus: ed. Received Nov 30; Accepted Jan This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

This article has been cited by other articles in PMC. Abstract Cannabinoid receptors CB are expressed throughout human skin epithelium. Keywords: Cannabinoid, Keratin, Psoriasis, Wound healing. Immunohistochemistry For the detection of K6 in organ cultured human skin as well as cultured HaCaT KCs, indirect immunofluorescence staining was performed using mouse anti-human K6 antibody Progen, Ks6. Open in a separate window. ACEA also down-regulates K16 protein expression in situ Since K16 is the type I keratin partner of K6 in KCs and is thought to stabilize this keratin protein as a cytoskeletal heteropolymeric intermediate filament Ramot et al.

CB1-mediated signaling also regulates K6 expression in cultured, hyperproliferative human keratinocytes In order to check whether the observed CB1-mediated effects on K6 regulation within intact human skin epithelium depend on intact epithelial-mesenchymal interactions between epidermis and dermis, or are likely to reflect a direct impact of CB1 ligands on epidermal keratinocytes, we next investigated K6 expression in cultured human HaCaT KCs.

The CB1 specific agonist, ACEA significantly decreases human epidermal keratinocyte proliferation in situ However, the observed effects of CB1-mediated signaling on epidermal K6 expression could simply reflect the appreciated anti-proliferative effects of CB1 agonists Casanova et al. The CB1 specific agonist, ACEA, significantly decreases K6 expression in suprabasal cells in a proliferation-independent manner Therefore, it needed to be dissected whether or not CB1 also reduces K6 expression in a proliferation-independent manner.

Conclusion Our results suggest that cannabinoids and their receptors constitute a novel, clinically relevant control element of human K6 and K16 expression. Acknowledgments The authors thank Motoko Sugawara for her excellent technical assistance, and Dr. Yuval Ramot and Koji Sugawara conceived and designed the experiments, performed the experiments, analyzed the data, wrote the paper.

References Akhmetshina et al. The cannabinoid receptor CB2 exerts antifibrotic effects in experimental dermal fibrosis. Arthritis and Rheumatism. IL is secreted by psoriatic keratinocytes and induces pro-inflammatory cytokines via keratinocyte and mast cell activation. Experimental Dermatology. The endocannabinoid system of the skin in health and disease: novel perspectives and therapeutic opportunities.

Trends in Pharmacological Sciences. Thyroid-stimulating hormone, a novel, locally produced modulator of human epidermal functions, is regulated by thyrotropin-releasing hormone and thyroid hormones. Normal keratinization in a spontaneously immortalized aneuploid human keratinocyte cell line.

Journal of Cell Biology. Vitronectin: effects on keratinocyte motility and inhibition of collagen-induced motility. Journal of Investigative Dermatology. Inhibition of skin tumor growth and angiogenesis in vivo by activation of cannabinoid receptors.

Journal of Clinical Investigation. Journal of Biological Chemistry. Endocannabinoids regulate growth and survival of human eccrine sweat gland-derived epithelial cells. Endocannabinoids enhance lipid synthesis and apoptosis of human sebocytes via cannabinoid receptormediated signaling. Etanercept restores a differentiated keratinocyte phenotype in psoriatic human skin: a morphological study.

The cannabinoid receptor CB 1 inverse agonist AM potentiates the anxiogenic activity of urocortin I in the basolateral amygdala. Inflammation Research. The cannabinoid receptor inverse agonist AM regulates the expression of the EGF receptor and its ligands via destabilization of oestrogen-related receptor alpha protein.

British Journal of Pharmacology. Evaluation of a range of anti-proliferative assays for the preclinical screening of anti-psoriatic drugs: a comparison of colorimetric and fluorimetric assays with the thymidine incorporation assay. Assay and Drug Development Technologies. Pathogenesis and clinical features of psoriasis. The effect of pimecrolimus on expression of genes associated with skin barrier dysfunction in atopic dermatitis skin lesions.

Contrasting protective effects of cannabinoids against oxidative stress and amyloid-beta evoked neurotoxicity in vitro. Cannabinoids, endocannabinoids, and cancer. Cancer and Metastasis Reviews. Development of quantitative molecular clinical end points for siRNA clinical trials. Unique keratinization process in psoriasis: late differentiation markers are abolished because of the premature cell death. Journal of Dermatology. Attenuation of allergic contact dermatitis through the endocannabinoid system.

Cannabinoid type-1 receptor reduces pain and neurotoxicity produced by chemotherapy. Journal of Neuroscience. Klein Klein TW. Cannabinoid-based drugs as anti-inflammatory therapeutics. Nature Reviews Immunology. Thyrotropin-releasing hormone controls mitochondrial biology in human epidermis. Journal of Clinical Endocrinology and Metabolism. Assessment of epidermal subpopulations and proliferation in healthy skin, symptomless and lesional skin of spreading psoriasis. British Journal of Dermatology.

Involvement of the endocannabinoid system in periodontal healing. Biochemical and Biophysical Research Communications. Cannabinoid system in the skin — a possible target for future therapies in dermatology. Thyrotropin-releasing hormone and oestrogen differentially regulate prolactin and prolactin receptor expression in female human skin and hair follicles in vitro. Keratins of the human hair follicle. International Review of Cytology.

Falcarinol is a covalent cannabinoid CB1 receptor antagonist and induces pro-allergic effects in skin. Biochemical Pharmacology. Towards the development of a simplified long-term organ culture method for human scalp skin and its appendages under serum-free conditions.

The endocannabinoid system in human keratinocytes. Evidence that anandamide inhibits epidermal differentiation through CB1 receptor-dependent inhibition of protein kinase C, activation protein-1, and transglutaminase. The human keratins: biology and pathology.

Histochemistry and Cell Biology. Changes in keratin 6 and keratin 10 co- expression in lesional and symptomless skin of spreading psoriasis. Namazi Namazi MR. Cannabinoids, loratadine and allopurinol as novel additions to the antipsoriatic ammunition. Journal of the European Academy of Dermatology and Venereology. Elements controlling the expression and induction of the skin hyperproliferation-associated keratin K6.

Onset of re-epithelialization after skin injury correlates with a reorganization of keratin filaments in wound edge keratinocytes: defining a potential role for keratin Anandamide regulates keratinocyte differentiation by inducing DNA methylation in a CB1 receptor-dependent manner. International Union of Basic and Clinical Pharmacology.

Thyrotropin powers human mitochondria. Endocannabinoids stimulate human melanogenesis via type-1 cannabinoid receptor. For all experiments, control and treated sections were stained and later evaluated on the same day by the same investigator.

To avoid staining biases, we calculated the relative staining intensity arbitrary intensity; 1 as control group among treatment groups per each individual and then pooled data from all of the experiments. For epidermal evaluation, staining intensity was evaluated in the suprabasal cells, and for HaCaT cells, staining intensity was measured in the colonies formed. Each treatment group was compared to the control group average value , and relative change in expression was calculated.

Highly comparable results were obtained from different sections from different individuals. This down-regulation was abrogated in part by the co-administration with the CB1-specific antagonist, AM Pertwee et al. Therefore, intraepidermal K6 protein expression in normal human skin in situ is down-regulated in a CB1-specific manner.

Since K16 is the type I keratin partner of K6 in KCs and is thought to stabilize this keratin protein as a cytoskeletal heteropolymeric intermediate filament Ramot et al. In order to check whether the observed CB1-mediated effects on K6 regulation within intact human skin epithelium depend on intact epithelial-mesenchymal interactions between epidermis and dermis, or are likely to reflect a direct impact of CB1 ligands on epidermal keratinocytes, we next investigated K6 expression in cultured human HaCaT KCs.

This transformed human KC line is well-appreciated to constitutively express K6 and to be hyperproliferative just like human wounded and psoriatic KC Balato et al. Unexpectedly, though, AM alone had a partial inhibitory effect on K6 expression, although not significant. Therefore, while HaCaT cells may exhibit relatively low CB expression levels, at least under our assay conditions, they showed a vigorous response to CB ligands.

However, the observed effects of CB1-mediated signaling on epidermal K6 expression could simply reflect the appreciated anti-proliferative effects of CB1 agonists Casanova et al. Just as we had seen before with the non-selective endocannabinoid, AEA Toth et al.

This effect was abrogated by the CB1-specific antagonist, AM assessed by quantitative Ki immunomorphometry, Figs. Therefore, it needed to be dissected whether or not CB1 also reduces K6 expression in a proliferation-independent manner. This was done by selectively assessing K6 expression in non-proliferative i. Kinegative epidermal KCs in situ. Furthermore, K6-expressing cells in the epidermis co-expressed CB1 in situ Fig. Thus, CB1 stimulation may affect K6 expression both, by reducing KC proliferation and by down-regulating K6 expression directly via CB1 in a proliferation-independent manner.

Specifically, we show that CB1 stimulation down-regulates expression of the hyper-proliferation-associated human keratins K6 in vitro and in situ , and inhibits human epidermal KC proliferation in situ. The effect of CB-mediated signaling in human KC biology remains to be fully explored.

As we have also observed in isolated human skin Fig. Therefore, it was important to clarify whether specific CB1 stimulation inhibits human epidermal KC proliferation in situ. By using CB1-specific agonists and antagonists we clearly demonstrate that exclusive CB1 stimulation inhibited KC proliferation.

Thus, CB1 is an important key regulator of human KC proliferation. Given the role of epidermal hyperproliferation in the pathobiology of psoriasis Donetti et al. Recently, we have shown that CB1 activation limits excessive mast cell activity and even inhibits mast cell maturation of resident, intracutaneous progenitors Sugawara et al. It should be noted, that the constitutive level of K6 expression in organ-cultured human skin fragments is considerably higher than normal scalp skin in vivo.

Presumably, this occurs as a response to tissue dissection and organ culture, which is well-known to elicit an immediate wound healing response in the epithelium. The latter rapidly starts to migrate over the wound edge in an attempt to enwrap the exposed skin mesenchyme epiboly phenomenon Stenn, ; Brown et al. This constitutive up-regulation of K6 in organ-cultured normal human skin may greatly heighten the sensitivity to K6 expression-modulatory compounds, such as CB ligands, thus making human skin organ culture a particularly sensitive and instructive tool for clinically relevant keratin research.

At the same time, however, caution is advised in extrapolating from these observation made in what essentially reflects a wound healing milieu to healthy, unmanipulated human skin in vivo. Our results suggest that cannabinoids and their receptors constitute a novel, clinically relevant control element of human K6 and K16 expression. Abbreviations ACEA arachidonoyl-chloro-ethanolamide. Common use cases Typos, corrections needed, missing information, abuse, etc.

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