Cardiovascular Division, University of Minnesota Medical School, Minneapolis, MN, USA.
Am J Physiol Heart Circ Physiol. 2013 Mar 1;304(5):H749-58. doi: 10.1152/ajpheart.00935.2011. Epub 2013 Jan 11.
Cell hypertrophy requires increased protein synthesis and expansion of the cytoskeletal networks that support cell enlargement. AMPK limits anabolic processes, such as protein synthesis, when energy supply is insufficient, but its role in cytoskeletal remodeling is not known. Here, we examined the influence of AMPK in cytoskeletal remodeling during cardiomyocyte hypertrophy, a clinically relevant condition in which cardiomyocytes enlarge but do not divide. In neonatal cardiomyocytes, activation of AMPK with 5-aminoimidazole carboxamide ribonucleotide (AICAR) or expression of constitutively active AMPK (CA-AMPK) attenuated cell area increase by hypertrophic stimuli (phenylephrine). AMPK activation had little effect on intermediate filaments or myofilaments but dramatically reduced microtubule stability, as measured by detyrosinated tubulin levels and cytoskeletal tubulin accumulation. Importantly, low-level AMPK activation limited cell expansion and microtubule growth independent of mTORC1 or protein synthesis repression, identifying a new mechanism by which AMPK regulates cell growth. Mechanistically, AICAR treatment increased Ser-915 phosphorylation of microtubule-associated protein 4 (MAP4), which reduces affinity for tubulin and prevents stabilization of microtubules (MTs). RNAi knockdown of MAP4 confirmed its critical role in cardiomyocyte MT stabilization. In support of a pathophysiological role for AMPK regulation of cardiac microtubules, AMPK α2 KO mice exposed to pressure overload (transverse aortic constriction; TAC) demonstrated reduced MAP4 phosphorylation and increased microtubule accumulation that correlated with the severity of contractile dysfunction. Together, our data identify the microtubule cytoskeleton as a sensitive target of AMPK activity, and the data suggest a novel role for AMPK in limiting accumulation and densification of microtubules that occurs in response to hypertrophic stress.
细胞肥大需要增加蛋白质合成和扩展细胞骨架网络,以支持细胞增大。当能量供应不足时,AMPK 限制合成代谢过程,如蛋白质合成,但它在细胞骨架重塑中的作用尚不清楚。在这里,我们研究了 AMPK 在心肌细胞肥大过程中细胞骨架重塑的影响,心肌细胞肥大是一种临床相关的情况,即心肌细胞增大但不分裂。在新生心肌细胞中,用 5-氨基咪唑甲酰胺核苷酸(AICAR)激活 AMPK 或表达组成型激活的 AMPK(CA-AMPK)可减轻肥大刺激(苯肾上腺素)引起的细胞面积增加。AMPK 激活对中间丝或肌丝几乎没有影响,但大大降低了微管稳定性,如通过去酪氨酸化微管蛋白水平和细胞骨架微管蛋白积累来衡量。重要的是,低水平的 AMPK 激活限制了细胞扩张和微管生长,而不依赖于 mTORC1 或蛋白质合成抑制,确定了 AMPK 调节细胞生长的新机制。从机制上讲,AICAR 处理增加了微管相关蛋白 4(MAP4)的丝氨酸-915 磷酸化,这降低了与微管蛋白的亲和力,并防止了微管的稳定(MTs)。MAP4 的 RNAi 敲低证实了其在心肌细胞 MT 稳定中的关键作用。支持 AMPK 调节心脏微管的病理生理作用,暴露于压力超负荷(横主动脉缩窄;TAC)的 AMPK α2 KO 小鼠显示 MAP4 磷酸化减少和微管积累增加,这与收缩功能障碍的严重程度相关。总之,我们的数据确定微管细胞骨架是 AMPK 活性的敏感靶点,并且数据表明 AMPK 在限制肥大应激时发生的微管积累和密集化方面具有新的作用。