Macdonald Laura J, Graham Joseph G, Kurten Richard C, Voth Daniel E
Department of Microbiology and Immunology, University of Arkansas for Medical Sciences, Little Rock, AR, USA.
Cell Microbiol. 2014 Jan;16(1):146-59. doi: 10.1111/cmi.12213. Epub 2013 Oct 9.
Intracellular bacterial pathogens often subvert apoptosis signalling to regulate survival of their host cell, allowing propagation of the bacterial population. Coxiella burnetii, the intracellular agent of human Q fever, inhibits host cell apoptosis through several mechanisms, including prevention of mitochondrial cytochrome c release, triggering of an anti-apoptotic transcriptional programme, and activation of pro-survival kinases. To control host cell survival, C. burnetii delivers effector proteins to the eukaryotic cytosol using a specialized Dot/Icm type IV secretion system (T4SS). Effectors are predicted to regulate activity of pro-survival host signalling proteins, such as Akt and cAMP-dependent protein kinase (PKA), to control infection. Here, we show that host PKA activity is required for C. burnetii inhibition of macrophage apoptosis. PKA is activated during infection and inhibits activity of the pro-apoptotic protein Bad via phosphorylation. Bad is also phosphorylated at an Akt-specific residue, indicating C. burnetii uses two kinases to fully inactivate Bad. Additionally, Bad and the tethering protein 14-3-3β colocalize at the C. burnetii parasitophorous vacuole (PV) membrane during infection, an event predicted to alter Bad promotion of apoptosis. Inhibiting PKA activity prevents Bad recruitment to the PV, but the protein is retained at the membrane during induction of apoptosis. Finally, PKA regulatory subunit I (RI) traffics to the PV membrane in a T4SS-dependent manner, suggesting a C. burnetii effector(s) regulates PKA-dependent activities. This study is the first to demonstrate subversion of host PKA activity by an intracellular bacterial pathogen to prevent apoptosis and survive within macrophages.
细胞内细菌病原体常常破坏凋亡信号传导以调控其宿主细胞的存活,从而使细菌群体得以繁殖。人类Q热的细胞内病原体伯氏考克斯体通过多种机制抑制宿主细胞凋亡,包括阻止线粒体细胞色素c释放、触发抗凋亡转录程序以及激活促存活激酶。为了控制宿主细胞的存活,伯氏考克斯体利用一种特殊的Dot/Icm IV型分泌系统(T4SS)将效应蛋白递送至真核细胞胞质溶胶中。预计效应蛋白会调节促存活宿主信号蛋白(如Akt和环磷酸腺苷依赖性蛋白激酶(PKA))的活性,以控制感染。在此,我们表明宿主PKA活性是伯氏考克斯体抑制巨噬细胞凋亡所必需的。PKA在感染过程中被激活,并通过磷酸化抑制促凋亡蛋白Bad的活性。Bad在一个Akt特异性残基处也被磷酸化,这表明伯氏考克斯体利用两种激酶来完全使Bad失活。此外,在感染期间,Bad和锚定蛋白14-3-3β在伯氏考克斯体吞噬泡(PV)膜上共定位,这一事件预计会改变Bad对凋亡的促进作用。抑制PKA活性可阻止Bad募集至PV,但在诱导凋亡期间该蛋白会保留在膜上。最后,PKA调节亚基I(RI)以T4SS依赖性方式转运至PV膜,这表明一种伯氏考克斯体效应蛋白调节PKA依赖性活动。本研究首次证明细胞内细菌病原体对宿主PKA活性的破坏,以防止凋亡并在巨噬细胞内存活。