The beneficial effects of OSK-induced reprogramming in axon regeneration and vision require the DNA demethylases TET1 and TET2. Using the eye as a model CNS tissue, here we show that ectopic expression of Oct4 (also known as Pou5f1), Sox2 and Klf4 genes (OSK) in mouse retinal ganglion cells restores youthful DNA methylation patterns and transcriptomes, promotes axon regeneration after injury, and reverses vision loss in a mouse model of glaucoma and in aged mice. Over time, the central nervous system (CNS) loses function and regenerative capacity 5– 7. Changes to DNA methylation patterns over time form the basis of ageing clocks 4, but whether older individuals retain the information needed to restore these patterns-and, if so, whether this could improve tissue function-is not known. A proposed cause of ageing is the accumulation of epigenetic noise that disrupts gene expression patterns, leading to decreases in tissue function and regenerative capacity 1– 3. jointly supervised this work.Īgeing is a degenerative process that leads to tissue dysfunction and death. helped with grant applications and project management. helped with plasmid constructs and AAV9 production. M.S.B., X.T., M.B.S., A.E.K., L.A.R., helped with systemic AAV9 experiments. helped with transgenic mouse fibroblasts work. conducted human methylation clock analysis. performed the RNA-seq and gene association analysis. conducted ribosomal DNAm age analysis for mouse RGCs. contributed to the glaucoma and ageing studies. contributed to the optic nerve crush studies and imaging. was involved in all experiments and analyses. wrote the manuscript with input from all co-authors.
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Before advancing, the player must choose a politician and each can provide free units or crates of resources. In Age of Empires III, these phases are called “Ages”, and represent historical time periods. AgesĪs with most RTS games, the player can advance through technological phases, which provide access to greater improvements, units, or buildings. Then as their economy and military improves they can destroy the enemy colonies or the enemy may offer to resign. When the experience points reach a certain threshold the players earn shipment cards that can be shipped from the Home City to the player. The actions such as training units, constructing buildings, and killing enemy units can give the player experience points. Which are used to train more units, create more buildings, and upgrade their technology. Players must explore the map and begin gathering the natural resources, food, coin, or wood. If the player has teammates, they and the land they visit are visible. When the game starts the exact terrain of the map which includes, the position of the enemy, native tribes, and trade post sites are blacked out. Unlike the other Age of Empires games, this game includes much prominent gunpowder warfare, including artillery and rifle companies. The player starts the game with a Town Center, an Explorer, and several Settlers or only a Covered Wagon and an Explorer depending on the type of game selected. Two expansion packs have been released: the first, Age of Empires III: The WarChiefs, was released on October 17, 2006, and introduced three Native American civilizations the second, Age of Empires III: The Asian Dynasties, was released on October 23, 2007, and included three Asian civilizations. Age of Empires III has made several innovations in the series, in particular with the addition of the “Home City”, which combines real-time strategy and role-playing features. There are fourteen civilizations to play within the game. 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