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. 2021 May 14;12:661955. doi: 10.3389/fneur.2021.661955

Table 2.

Inflammatory markers associated with HT.

Early HT (first 18 h of stroke onset)
Inflammatory marker Source/mechanism Role in HT
ROS (increase) Intracellular mitochondria, nicotinamide adenine dinucleotide phosphate (NADPH)-oxidase xanthine oxidases, cellular membrane receptors inflammatory mediators, astrocytes Disrupt the NVU (endothelial-pericyte-astrocyte) leading to increase BBB degradation
HMGB1 Microglia, astrocytes Upregulates MMP-9 via TLR4. TNF, IL-1β
Peroxynitrite Micro vessels, neurons and astrocytic end feet Activate MMPs, disrupting vascular integrity
NF-κB Astrocytes, microglia, and endothelial cells Part of neutrophil infiltration pathway, upregulate cell adhesion molecules and inflammatory cytokines.
Leukocytes (increase) Circulating leukocytes adhere to vascular endothelial cells following ischemia. Leukocyte adhesion and migration across the vasculature activates a number of signaling cascades (protein kinase C, focal adhesion kinase) that increase BBB permeability Signaling cascade increased BBB permeability through ROS and MMP-9 expression.
MMP-9 (blood derived) (increase) Leukocytes (Neutrophils) (not primary source). Mechanisms for MMP-9 activation following ischemia include: (1) ROS (2) TNF, IL-1β, and other cytokines that activate MMP-3 which converts proMMP-9 to active MMP-9 (3) actions of high mobility group box protein1 (HMGB1) on TLR4 receptors that then induce MMP-9 or (4) NF-κB induction of MMP-9 Luminal side: acts on TJP (tight junction proteins) (claudin-5, occludin, ZO-1) and basal lamina proteins (fibronectin, lamina, collagen), taken into endothelial cells or acts on basal lamina to open the BBB. Inside brain: Neutrophils can release MMPs that act directly on TJPs and/or basal lamina to disrupt the NVU (endothelial-pericyte-astrocyte)
MMP-2 (brain derived) (increase) remains elevated for days post-stroke Astrocytes, endothelial cells and leukocytes MMP-2 mediates degradation of occludin (tight junction protein)
Leukocyte gene expression Six genes identified through mRNA expression: amphiregulin (AREG), membrane-associated ring finger (C3HC4) 7 (MARCH7), SMAD family member 4 (SMAD4), inositol polyphosphate-5-phosphatase (INPP5D), multiple coagulation factor deficiency 2 (MCFD2) and vascular endothelial growth inhibitor (VEGI)
Late HT (18–24 h of stroke onset)
Source Role
MMP-9 (Brain derived) (Increase) MMP-9: astrocytes, neurons, microglia and endothelial cells. Activated by from ROS, TNF and IL-1β, HMGB1, NF-κB induction Disruption of BBB
MMP-3 (Brain derived) (Increase) MMP-3: pericytes and endothelial cells. MMP-3 acts on proMMP-9 to produce active MMP-9 and thus may promote HT Disruption of BBB
Role of Vascular Remodeling • A number of growth factors, MMPs and other molecules form new vessels and NVU
• VEGF plays an important role in vascular remodeling and angiogenesis
VEGF: early, promotes BBB disruption; later promotes BBB integrity and vascular function. MMPs: Promote new vessel formation and increased pericyte/endothelial expression of tight junction proteins (ZO-1, occludin, claudin-5) HMGB1: acts on endothelial progenitor cells to promote peri-infarct angiogenesis (beneficial role)
ROS Act as signaling molecules to regulate cell growth, differentiation and angiogenesis.

BBB, blood-brain barrier; HMGB1, high mobility group box protein 1; IL, interleukin; MMP, matrix metalloproteinase; NF-κB, nuclear factor-κB; NVU, neurovascular unit; ROS, reactive oxygen species; TLR, toll like receptor; TNF, tumor necrosis factor; TJP, tight junction protein; VEGF, vascular endothelial growth factor.