Table 1.

Summary of examples of virus–lipid interactions discussed in this review.

Stage	Lipid class	Mode of action	Virus examples
Docking/fusion	Phosphatidylserines	PS addition causes a “glycosylation-specific enhancement” of virus entry	HIV (Coil and Miller, 2005b, Coil and Miller, 2005a)
		Annexins are hijacked as co-receptors by PS expressed in the virus envelope	Influenza virus (Huang et al., 1996), HBV (Hertogs et al., 1993), HCMV (Raynor et al., 1999), RSVa (Malhotra et al., 2003), HIV (Callahan et al., 2003, Ma et al., 2004)
		Molecular mimicry of apoptotic bodies facilitates macropinocytosis/virus entry	VACV (Mercer and Helenius, 2008)
	Sterols/Sterol Esters	Association with apolipoproteins facilitates virus docking and subsequent entry steps	HCV (Prince et al., 1996, Agnello et al., 1999, Molina et al., 2007)
		Virus surface glycoproteins and receptors are clustered in “lipid rafts”	HCV (Kapadia et al., 2007, Burlone and Budkowska, 2009), HIV (Sorice et al., 2000, Sorice et al., 2001, Nguyen and Taub, 2002, Popik et al., 2002, Bhattacharya et al., 2004, Nguyen et al., 2005), RSVa (Fleming et al., 2006), influenza virus (Takeda et al., 2003)
	Sphingolipids	Glycan–glycan interactions between host and virus stabilize virus attachment	Influenza virus (Huang, 1976, Leskawa et al., 1986, Kasson and Pande, 2008)
		GalCer, GM3 and Gb3 serve as auxiliary receptors for virus entry	HIV (Fantini et al., 1993, Seddiki et al., 1996, Hammache et al., 1999, Hug et al., 2000, Magerus-Chatinet et al., 2007)
		GSLs aggregate viral receptors which have low endogenous cellular levels	EBOV, Marburg virus, VACV, HSV (reviewed in Teissier and Pecheur, 2007)
		Gangliosides serve as main receptors for virus entry	SeV (Markwell et al., 1981, Hansson et al., 1984, Markwell et al., 1984, Muthing and Unland, 1994, Kaneda et al., 1997), NDV (Ferreira et al., 2004a, Ferreira et al., 2004b, Villar and Barroso, 2006)
Replication/translation	Phosphatidylserines	PS enriched in endosomal membranes is required by NSP1 for capping of viral mRNAs	SFV (Ahola et al., 1999)
	Sterols/sterol esters	Nef accessory protein enhances cholesterol biosynthesis and impairs ABCA1 dependent cholesterol efflux	HIV (Mujawar et al., 2006)
	Sphingolipids	Enhanced levels of cholesterol are required to support formation of “lipid-raft” structures	RSVa (Money et al., 2009)
		Increased activity of SphK1 promotes accumulation of transcriptional activator IE1	HCMV (Machesky et al., 2008)
	Fatty acids	Genome replication is stimulated by saturated or monounsaturated fatty acids. FAS is highly enriched in the supernatant of infected cells	HCV (Su et al., 2002)
Assembly/budding	Phosphoinositides	Gag binds specifically to PI(4,5)P2 causing a myristic switch in MA which increases membrane partitioning of Gag	HIV (Zhou et al., 1994, Ono et al., 2004, Freed, 2006)
	Sterols	It is also possible that PI(4,5)P2 interactions with cellular proteins contribute to membrane curvature during virus assembly and budding	HIV (Chan et al., 2008)
		Surface exposed polybasic domains of similar matrix proteins also use phosphoinositides or other anionic phospholipids for “lipid-raft” targeting/induction of membrane curvature	Influenza virus (Ruigrok et al., 2000, Thaa et al., 2009), EBOV (Timmins et al., 2004), VSV (Gaudier et al., 2002), RSVa (Money et al., 2009)
		Virus assembly occurs on lipid droplets	HCV (Miyanari et al., 2007)
	Sphingolipids	Nef contains cholesterol binding sequence in its C terminus which allows transport to and association with lipid rafts	HIV (Zheng et al., 2003)
		Cholesterol depletion enhances virus budding	Influenza virus (Barman and Nayak, 2007)
		Cholesterol depletion inhibits virus production	HIV (Ono and Freed, 2001, Pickl et al., 2001)
		“Lipid-raft” dependent viruses do not co-localize on the plasma membrane	HIV, Influenza virus, EBOV (Khurana et al., 2007, Leung et al., 2008)
	Fatty acids	Several viral proteins have myristoyl and palmitoyl covalent modifications for “lipid-raft” targeting	HIV, Influenza virus, EBOV, Marburg virus, measles virus, NDV, RSVa (reviewed in Hruby and Franke, 1993, Maurer-Stroh and Eisenhaber, 2004)
Extracellular	Sterols/sterol esters	Cholesterol depletion of virus envelopes leads to virus inactivation due to a loss of protein core and genome integrity	HIV and SIV (Graham et al., 2003), influenza virus (Barman and Nayak, 2007, Ambrose et al., 2008)
	Lipid molecular geometry	Inverted-cone shaped lipids like lyso PC inhibits virus envelope fusion with target membrane while cone shaped lipids like pPE, DAG and ceramide may facilitate membrane dynamics	HIV (Martin and Ruysschaert, 1995, Gunther-Ausborn and Stegmann, 1997, Brugger et al., 2006, Chan et al., 2008), TBEV (Stiasny and Heinz, 2004), influenza virus (Gunther-Ausborn et al., 1995, Chernomordik et al., 1998)