Original Article
Copyright ©2013 Baishideng. All rights reserved.
World J Transplant. Sep 24, 2013; 3(3): 36-47
Published online Sep 24, 2013. doi: 10.5500/wjt.v3.i3.36
mTOR signaling in liver regeneration: Rapamycin combined with growth factor treatment
Suomi MG Fouraschen, Petra E de Ruiter, Jaap Kwekkeboom, Ron WF de Bruin, Geert Kazemier, Herold J Metselaar, Hugo W Tilanus, Luc JW van der Laan, Jeroen de Jonge
Suomi MG Fouraschen, Petra E de Ruiter, Ron WF de Bruin, Geert Kazemier, Hugo W Tilanus, Luc JW van der Laan, Jeroen de Jonge, Department of Surgery and Laboratory of Experimental Transplantation and Intestinal Surgery, Erasmus MC-University Medical Center, 3015 CE Rotterdam, The Netherlands
Jaap Kwekkeboom, Herold J Metselaar, Department of Gastroenterology and Hepatology, Erasmus MC-University Medical Center, 3015 CE Rotterdam, The Netherlands
Author contributions: van der Laan LJW and de Jonge J contributed equally to this study; Fouraschen SMG designed the study, performed the experiments, collected and analyzed the data and wrote the manuscript; de Ruiter PE performed experiments, collected and analyzed data and edited the manuscript; Kwekkeboom J and de Bruin RWF provided scientific input as well as analytic tools and edited the manuscript; Kazemier G, Metselaar HJ and Tilanus HW provided clinical and scientific input and edited the manuscript; van der Laan LJW and de Jonge J designed the study, analyzed data and wrote the manuscript.
Supported by Erasmus MC Grant and the Liver Research Foundation (SLO) Rotterdam
Correspondence to: Dr. Luc JW van der Laan, Department of Surgery and Laboratory of Experimental Transplantation and Intestinal Surgery, Erasmus MC-University Medical Center, ‘s Gravendijkwal 230, 3015 CE Rotterdam, The Netherlands. l.vanderlaan@erasmusmc.nl
Telephone: +31-10-7032759 Fax: +31-10-7032793
Received: March 7, 2013
Revised: May 28, 2013
Accepted: June 18, 2013
Published online: September 24, 2013
Processing time: 170 Days and 0.1 Hours
Abstract

AIM: To investigate the effects of mammalian target of rapamycin (mTOR) inhibition on liver regeneration and autophagy in a surgical resection model.

METHODS: C57BL/6 mice were subjected to a 70% partial hepatectomy (PH) and treated intraperitoneally every 24 h with a combination of the mTOR inhibitor rapamycin (2.5 mg/kg per day) and the steroid dexamethasone (2.0 mg/kg per day) in phosphate buffered saline (PBS) or with PBS alone as vehicle control. In the immunosuppressant group, part of the group was treated subcutaneously 4 h prior to and 24 h after PH with a combination of human recombinant interleukin 6 (IL-6; 500 μg/kg per day) and hepatocyte growth factor (HGF; 100 μg/kg per day) in PBS. Animals were sacrificed 2, 3 or 5 d after PH and liver tissue and blood were collected for further analysis. Immunohistochemical staining for 5-Bromo-2’-deoxyuridine (BrdU) was used to quantify hepatocyte proliferation. Western blotting was used to detect hepatic microtubule-associated protein 1 light chain 3 (LC3)-II protein expression as a marker for autophagy. Hepatic gene expression levels of proliferation-, inflammation- and angiogenesis-related genes were examined by real-time reverse transcription-polymerase chain reaction and serum bilirubin and transaminase levels were analyzed at the clinical chemical core facility of the Erasmus MC-University Medical Center.

RESULTS: mTOR inhibition significantly suppressed regeneration, shown by decreased hepatocyte proliferation (2% vs 12% BrdU positive hepatocyte nuclei at day 2, P < 0.01; 0.8% vs 1.4% at day 5, P = 0.02) and liver weight reconstitution (63% vs 76% of initial total liver weight at day 3, P = 0.04), and furthermore increased serum transaminase levels (aspartate aminotransferase 641 U/L vs 185 U/L at day 2, P = 0.02). Expression of the autophagy marker LC3-II, which was reduced during normal liver regeneration, increased after mTOR inhibition (46% increase at day 2, P = 0.04). Hepatic gene expression showed an increased inflammation-related response [tumor necrosis factor (TNF)-α 3.2-fold upregulation at day 2, P = 0.03; IL-1Ra 6.0-fold upregulation at day 2 and 42.3-fold upregulation at day 5, P < 0.01] and a reduced expression of cell cycle progression and angiogenesis-related factors (HGF 40% reduction at day 2; vascular endothelial growth factor receptor 2 50% reduction at days 2 and 5; angiopoietin 1 60% reduction at day 2, all P≤ 0.01). Treatment with the regeneration stimulating cytokine IL-6 and growth factor HGF could overcome the inhibitory effect on liver weight (75% of initial total liver weight at day 3, P = 0.02 vs immunosuppression alone and P = 0.90 vs controls) and partially reversed gene expression changes caused by rapamycin (TNF-α and IL-1Ra levels at day 2 were restored to control levels). However, no significant changes in hepatocyte proliferation, serum injury markers or autophagy were found.

CONCLUSION: mTOR inhibition severely impairs liver regeneration and increases autophagy after PH. These effects are partly reversed by stimulation of the IL-6 and HGF pathways.

Keywords: Hepatocyte proliferation, Autophagy, Microtubule-associated protein 1 light chain 3, Partial hepatectomy, Rapamycin

Core tip: Interference of immunosuppressive medication with liver regeneration is a highly relevant issue for transplantation of small-for-size liver grafts. Inhibition of mammalian target of rapamycin (mTOR) represents an important immunosuppressive strategy after transplantation, yet as mTOR regulates cell proliferation and autophagy, concerns remain regarding a negative impact on regeneration. The exact role of mTOR signaling after living-donor liver transplantation is largely unknown. Here we report that mTOR inhibition by rapamycin severely impairs liver regeneration and increases autophagy after liver resection in mice. The most novel finding of this study is that this impaired regeneration can be partly reversed by treatment with exogenous growth factors.