Herpesvirus Nuclear Egress across the Outer Nuclear Membrane.
Herpesvirus Nuclear Egress across the Outer Nuclear Membrane.
复制标题
疱疹病毒核流横跨外核膜。
DOI：
10.3390/v13122356
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发表时间：
2021-11-24
期刊：
影响因子：
--
通讯作者：
Johnson DC
中科院分区：
文献类型：
--
作者：
Roller RJ;Johnson DC
疱疹病毒衣壳在细胞核中组装，并经过两步过程穿过核膜。衣壳在核出口复合物（NEC）蛋白UL 31/34的帮助下发芽进入内核膜（INM）。在这个阶段的出口，包膜病毒粒子被发现在核周空间的短时间。在核出口的第二步中，核周包膜病毒体（PEV）与外核膜（ONM）融合，将衣壳递送到细胞质中。一旦在细胞质中，衣壳在高尔基体/反高尔基体中经历再结晶，产生成熟的病毒粒子。核出口的第二步被称为去核化，这也是本综述的重点。与INM的疱疹病毒感染相比，对疱疹病毒感染的了解要少得多。我们提出了一个模型，其中去病毒化涉及两个阶段：（i）PEV膜与ONM的融合和（ii）融合孔的扩张导致病毒衣壳释放到细胞质中。第一个阶段的去除，膜融合，涉及四个单纯疱疹病毒（HSV）蛋白：gB，gH/gL，gK和UL 20。gB是病毒融合蛋白，似乎起扰乱膜和促进融合的作用。gH/gL也可能具有类似的性质，并且似乎能够在没有gB的情况下发挥作用。gK和UL 20负调控这些融合蛋白。在脱膜的第二阶段（孔扩张和衣壳释放），α-疱疹病毒蛋白激酶US 3使NEC蛋白磷酸化，其通常在脱膜期间产生膜弯曲。NEC蛋白的磷酸化逆转紧密的膜曲率，引起膜融合孔的扩张并促进衣壳释放到细胞质中。
Herpesvirus capsids are assembled in the nucleus and undergo a two-step process to cross the nuclear envelope. Capsids bud into the inner nuclear membrane (INM) aided by the nuclear egress complex (NEC) proteins UL31/34. At that stage of egress, enveloped virions are found for a short time in the perinuclear space. In the second step of nuclear egress, perinuclear enveloped virions (PEVs) fuse with the outer nuclear membrane (ONM) delivering capsids into the cytoplasm. Once in the cytoplasm, capsids undergo re-envelopment in the Golgi/trans-Golgi apparatus producing mature virions. This second step of nuclear egress is known as de-envelopment and is the focus of this review. Compared with herpesvirus envelopment at the INM, much less is known about de-envelopment. We propose a model in which de-envelopment involves two phases: (i) fusion of the PEV membrane with the ONM and (ii) expansion of the fusion pore leading to release of the viral capsid into the cytoplasm. The first phase of de-envelopment, membrane fusion, involves four herpes simplex virus (HSV) proteins: gB, gH/gL, gK and UL20. gB is the viral fusion protein and appears to act to perturb membranes and promote fusion. gH/gL may also have similar properties and appears to be able to act in de-envelopment without gB. gK and UL20 negatively regulate these fusion proteins. In the second phase of de-envelopment (pore expansion and capsid release), an alpha-herpesvirus protein kinase, US3, acts to phosphorylate NEC proteins, which normally produce membrane curvature during envelopment. Phosphorylation of NEC proteins reverses tight membrane curvature, causing expansion of the membrane fusion pore and promoting release of capsids into the cytoplasm.
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