Table 1.

Interactions of selected (+) strand RNA viruses with the nucleolus.

Virus family	Virus Name	Viral factorsa	Cellular proteinsb	Effects on the host and/or the virus	References
Flaviviridae	DENV	Capsid			[41], [42]
	WNV	Capsid (123 aa)	HDM2, DDX56, Jab1	DDX56 important for virus infectivity HDM2 localized in the nucleolus and induction of p53-dpt apoptosis Jab1 protects cells against the capsid cytoxic effects	[43], [44], [60], [61]
	JEV	Capsid	B23	B23 important for virus replication	[45], [130]
	Kunjin virus	Capsid			[46]
	HCV	Core, NS5B?	Nucleolin, B23, PKR	Upregulation of B23 synthesis via reduction of YY1 repressive activity on B23 promoter NCL is important for virus replication NCL found at HCV IRES	[36], [37], [38], [53], [62], [63], [70], [131]
Coronaviridae	SARS-CoV	3b, Nucleocapsid	B23	Inhibition of B23 phosphorylation	[31], [32], [39], [40], [132]
	Avian IBV	Nucleocapsid	Nucleolin, p53	Alteration of fibrillarin localization p53 delocalized in the perinuclear region Inhibition of cell growth	[47], [48], [133], [134]
Arteriviridae	PPRSV	Nucleocapsid	Fibrillarin, HIC	Mutation of N prevents its nucleolar localization reduces viral replication in vitro, delays viremia and increases NAb titers in vivo.	[17], [49], [51], [52], [135]
Togaviridae	SFV	nsP2, capsid		Mutation of nsP2 prevents its nuclear localization and reduces viral spread and neurovirulence in vivo	[34], [35], [57], [73], [74], [75], [76], [136]
	GETV M1		P21waf	S-phase arrest and apoptosis in glioma cells	[72]

^aViral proteins shown to localize in the nucleolus.

^bNucleolar proteins interacting with viral factor or cellular, non nucleolar proteins shown to be translocated into the nucleolus by the virus or the viral proteins.