Table 1.

Examples of heparin/HS-binding proteins and their biological activity.

Class	Examples	Physiological Function	Pathophysiology	Refs
Growth factors	FGF2, VEGF165, neuropilin-1	mitogenesis, development, wound healing, angiogenesis, axon guidance	cancer	8b, 19
ECM proteins	collagen, fibrinogen, laminin	cell adhesion, migration, differentiation, blood coagulation	cancer, Knobloch syndrome	20
Cell adhesion proteins	P-selectin, L-selectin, integrins	cell adhesion, inflammation	cancer	21
Morphogens	activin, BMP-2, sonic hedgehog	development, regeneration, bone formation	multiple hereditary exostoses, osteoarthritis	22
Chemokines	platelet factor 4, IL-8, TNF-α, CXCL12	chemotaxis, cell migration, immune response, angiogenesis	inflammation, arteriosclerosis, cancer	23
Blood coagulation factors	antithrombin III, factor Xa, leuserpin-2	regulation of clotting cascade	heparin-induced thrombocytopenia	24
Lipoproteins	ApoE, ApoB, lipoprotein lipase	lipid metabolism, cell membrane functions	atherosclerosis, Alzheimer’s disease, AA amyloidosis	25
Nuclear proteins	histones, transcription factors	unknown	cancer	26
Amyloid proteins	APP, Aβ, tau protein, α-synuclein, PrPSc	synapse organization, brain development, memory, circadian rhythm	Alzheimer’s/Parkinson’s disease, prion disease	27
Viral proteins	gB, gC, gD, gp120, Tat, E protein, L1 capsid protein	-----	HIV-1, HPV, HSV-1, HCMV, Dengue virus	28
Microbial proteins	M protein, PfEMP1, Opa, circumsporozoite protein	-----	bacterial/parasite infection, inflammation	29