1. Introduction

2. miRNA production and dynamics

3. miRNAs as biomarkers in AKI

4. miRNAs as therapeutic targets for AKI

4.2. Therapeutic targeting of miRNAs for AKI

Some of the most important evidence regarding miRNAs in AKI was reported in 2010, where knockouts of tubular miRNAs were analyzed in the rodent model [74]. The loxp-cre system was used to produce mice lacking Dicer, a key enzyme for miRNA production, in proximal tubular cells. The mice showed global downregulation of miRNAs in the kidney cortex and had normal kidney function and histology under normal conditions, while they were resistant to renal ischemia-reperfusion (I/R) injury, demonstrating the involvement of miRNAs under AKI. Since then, there has been accumulating evidence supporting miRNAs as potential therapeutic targets in AKI. Several miRNAs have been reported to have protective and/or pathogenic roles in AKI, regulating tubular apoptosis, tubulointerstitial fibrosis and inflammation in a variety of etiologies of AKI, including ischemia, drug and sepsis. Some miRNAs have common gene targets. For example, miR-30 and miR-26a target Snai1, regulating epithelial–mesenchymal transition (EMT), while miR-21, miR-17, miR-188 and miR-378 target PTEN, which is implicated in cell apoptosis, proliferation, inflammation and fibrosis [27]. The potential therapeutic targets of miRNAs for AKI are summarized in Table 2.

miR-21 is one of the most analyzed miRNAs, described as having double-edged-sword effects in kidneys [67] and both protective and pathogenic effects in kidney diseases. As a protective effect, miR-21 ameliorates I/R-induced AKI by inhibiting tubular cell apoptosis in I/R and LPS-induced AKI mice [71,75], targeting PTEN/Akt/mammalian target of rapamycin (mTOR) signaling and Cyclin-dependent kinase 6 (CDK6). miR-21 is also shown to inhibit maturation of dendritic cells through the PDCD4/ NFκ-B pathway [71] and CCR7 [76], thereby mediating anti-inflammatory effects in I/R-induced AKI mice. In addition, miR-21 is also shown to target mitogen-activated protein kinase kinase 3 (MKK3), inhibiting the downstream factors IL-6 and TNF-α levels, mediating anti-inflammatory effects [77]. On the other hand, as a pathogenic effect, miR-21 inhibits autophagy in I/R-AKI rats by targeting Rab11a [78]. In addition, long-term elevation of miR-21 might promote kidney fibrosis, including via PPARα [79]. Furthermore, miR-21 was shown to regulate energy metabolism via AKT/Cyclin-dependent kinase 2 (CDK2)-FOXO1 in a sepsis-induced rat AKI model, while it was unclear whether the regulation was protective or harmful for long-term prognosis [80]. Taken together, these data suggest that miR-21 may target several signaling pathways, involving cell apoptosis, inflammation and autophagy as well as energy metabolism under AKI. Similarly to miR-21, miR-30, miR-181, miR-22 and miR-590 are also reported to mediate anti-apoptosis and anti-inflammation effects in I/R-AKI rodent models. It was reported that MSC-derived extracellular vesicles ameliorated rat I/R-induced AKI by inhibiting cell apoptosis through miR-30, which targeted dynamin-related protein 1 (DRP1), thereby inhibiting mitochondrial fission [81]. In addition, injection of miR-30c-5p agomir, chemically modified double-stranded small RNA that mimics the miR-30c-5p, ameliorated rat I/R-induced AKI through transformation of M1 to M2 macrophages, mediating an anti-inflammatory effect via changes in inflammatory cytokines [82]. miR-181d-5p overexpression in the mouse model of I/R-induced AKI ameliorated kidney injury by reducing inflammatory mediators and apoptosis through targeting Krueppel-like factor 6 (KLF6) [83]. A renoprotective effect of miR-181 was also shown in a cisplatin-induced mouse AKI model by targeting PTEN [84] in LPS-induced AKI model through an anti-apoptosis effect by targeting GJB2 [85]. In addition, miR-181a-5p is reported to target NIMA-related kinase 7 (NEK7), thereby inhibiting pyroptosis in sepsis-induced AKI mice [86]. It was reported that in a sepsis-induced AKI mouse model, lncRNA TCONS_00016233, targeting miR-22, was upregulated in plasma, and TCONS_00016233 overexpression worsened sepsis-induced mouse AKI by downregulating miR-22, thus increasing the mir-22 target, with apoptosis-inducing factor mitochondrion-associated 1 (AIFM1) leading to apoptosis [87], indicating an anti-apoptosis effect of miR-22. In addition, miR-22 attenuated sepsis-induced rat AKI, targeting High Mobility Group Box 1 (HMGB1) and inhibiting the HMGB1/TLR4/NF-kB pathway [88]. Adenovirus expressing miR-590-3p via tail-vein injection in LPS-induced septic AKI mice ameliorated cell apoptosis and inflammation by targeting tumor necrosis factor receptor-associated factor 6 (TRAF6) [89]. miR-590-3p was also shown to oissibly increase autophagy and protect kidney injury by targeting TRAF6, which was evaluated in an in vitro I/R model using renal tubular epithelial cell line (HK-2 cells) [90].

There are several reports indicating an anti-apoptosis effect from miRNAs. miR-124 was shown to ameliorate I/R-induced mouse AKI, where miR-124 mimics reduced endoplasmic reticulum stress (ERS)-mediated apoptosis [91]. In addition to the anti-apoptosis mechanism, miR-124 also inhibited necroptosis by targeting PARP1 in an I/R-induced mouse AKI model [92]. miR-489 also targeted PARP1, mediating an anti-apoptosis effect, and miR-489 mimics protected against I/R mouse AKI [73]. miR-17-5p mimics suppressed death receptor 6 (DR6), mediating anti-apoptosis in an I/R-induced mouse AKI model [93]. Likewise, miR-424 mimics inhibited its target gene DR6, mediating anti-apoptosis in an I/R-induced mouse AKI model [94]. miR-5100 mimic injection into I/R-mice ameliorated kidney injury by inhibiting several apoptotic pathways [55]. miR-486-5p was shown to target PTEN, thereby mediating an anti-apoptosis effect in a mouse I/R-AKI model [95]. miR-191-5p mimic injection could inhibit cell apoptosis by targeting Oxidative stress responsive 1 (OXSR1) [96]. miR-290-5p activated by propofol ameliorated a sepsis-induced mouse AKI model by targeting C-C motif chemokine ligand 2 (CCL2), thereby mediating an anti-apoptosis effect [97]. In addition to the anti-apoptosis effect, an anti-inflammation effect by miRNAs has also been explored. miR-204/miR-211 mimics, targeting H6 Family Homeobox 1 (Hmx1), reduced kidney injury via immune suppression in candidemia-induced AKI mice [98]. miR-195-5p mimic injection ameliorated rat I/R-AKI by targeting vascular endothelial growth factor A (VEGFA) via anti-inflammatory and anti-oxidative stress [99]. miR-140-5p up-regulation by apigenin ameliorated I/R-induced AKI mice through targeting Chemokine (C-X-C Motif) Ligand 12 (CXCL2), thereby reducing inflammation [100]. In addition, miR-140-5p was shown to activate nuclear factor erythroid 2-related factor (Nrf2) pathway, thereby mediating anti-oxidative stress in cisplatin-induced AKI mice [101]. miR-27a targeting Toll-like receptor 4 (TLR4) inhibited inflammation in I/R-induced AKI [102]. In addition, overexpression of LINC00520, targeting miR-27b-3p, activated Oncostatin M Receptor (OSMR), leading to the PI3K/AKT pathway to aggravate kidney injury in I/R-induced AKI, while up-regulation of miR-27b-3p could accelerate recovery from AKI [103]. miR-146a-5p derived from human urine-derived stem cells protected against rat I/R-induced AKI by targeting interleukin 1 receptor associated kinase 1 (IRAK1), thereby inhibiting NF-κB signaling and infiltration of inflammatory cells [104]. Mice lacking miR-146 showed more extensive tubular injury, inflammatory infiltrates, and fibrosis than wild-type mice after I/R-induced AKI [105]. Furthermore, the renoprotective effect of miRNAs involves vascular, mitochondrial and podocyte protection. miR-210 was shown to activate VEGF signaling to regulate angiogenesis in I/R-induced AKI mice [106]. Overexpression of miR-126 in the hematopoietic compartment promoted vascular integrity and supported recovery of kidney injury after I/R-induced AKI mice [107]. miR-668 was shown to target mitochondrial protein 18 kDa (MTP18) to preserve mitochondrial dynamics and tubular cell survival in I/R-AKI mice [72]. miR-187 agomir injection in I/R protected against I/R-induced AKI and mediated podocyte protection, evaluated by nephrin expression by targeting acetylcholinesterase (AChE) [108].

There are several reports indicating MSC-related miRNAs as providing renoprotection via anti-apoptosis. MSC-derived exosome containing miR-199a-3p targeted semaphorin3A (SEMA3A), leading to the AKT and Extracellular signal-regulated kinase (ERK) pathway activation and ameliorated apoptosis [109]. Bone-marrow-derived MSCs ameliorate I/R-induced mouse AKI via miR-223, targeting NLR family pyrin domain containing 3 (NLRP3), thereby inhibiting apoptosis [110]. Human umbilical-cord-derived MSCs containing miR-125b-5p suppressed p53, leading to an anti-apoptosis effect [111]. Bone-marrow-derived MSCs reduced miR-107, increasing the expression of the miR-107 target, Ribosomal Protein S19 (RPS19), and protected from cisplatin-induced apoptosis [112]. In contrast, miR-107 was shown to induce TNF-α by targeting dual specificity phosphatase 7 (DUSP7), causing tubular injury [113]. Microvesicles from human Wharton’s Jelly MSCs ameliorated I/R-induced rat AKI by suppressing C-X3-C Motif Chemokine 1 (CXCL1) and ameliorated inflammation and fibrosis partly via miR-16, miR-15a and miR-15b [114]. On the other hand, urine miR-16 transactivated by CCAAT enhancer binding protein beta (C/EBP-β) worsened I/R-induced AKI, causing apoptosis [41]. Bone-marrow-derived MSCs ameliorated cisplatin-induced AKI by inhibiting fibrosis through the regulation of miR-146a-5p and its target transcription Factor Dp-2 (Tfdp2) [115].

In addition to these renoprotective miRNAs, there are several pathogenic miRNAs under AKI, aggravating kidney injury by causing apoptosis, inflammation, mitochondrial damage and fibrosis. It is reported that the administration of antisense oligonucleotide inhibiting miR-182 ameliorated an I/R-induced AKI rat model [116]. In an other report, an miR-182 inhibitor ameliorated I/R-induced rat AKI and apoptosis by regulating the transcription factor 7-like-2 (TCF7L2)/ Wnt/β-catenin pathway [117]. miR-182 was also shown to target Forkhead box O3 (FoxO3), leading to cell apoptosis in an I/R-induced AKI rat model [118]. Similarly, there are several miRNAs reported to promote cell apoptosis under AKI, including miR-122, miR-301, miR-375, miR-188, miR-687, miR-24 and miR-218. miR-122 was suggested to target FoxO3, thereby causing cell apoptosis in cisplatin-induced AKI mice [119]. miR-301a-5p inhibition ameliorated vancomycin-induced AKI by reducing apoptosis [120]. miR-375 was reported to be induced in a cisplatin-induced mouse AKI model to repress hepatocyte nuclear factor 1 homeobox B (HNF-1β), leading to the promotion of tubular cell apoptosis [121]. miR-188 was shown to aggravate contrast-induced AKI by targeting Serine And Arginine Rich Splicing Factor 7 (SRSF7), leading to cell apoptosis [122]. miR-687 was shown to be interfered with by lncRNA TCONS_00016406, leading to an anti-apoptosis effect [123]. miR-24 silencing protected against I/R-induced mouse AKI by inhibiting cell apoptosis by targeting H2A histone family, member X, (H2A.X) and HO-1 [124]. It was also suggested that honokiol inhibited miR-218-5p, leading to an increase in its target HO-1, thereby mediating an anti-apoptosis effect in sepsis-induced mouse AKI [125]. Several miRNAs, namely, miR-494, miR106, miR-155 and miR-152, were shown to promote inflammation as well as apoptosis. It is reported that miR-494 down-regulation by lncRNA TUG1 reduced I/R-induced mouse AKI and cell apoptosis by regulating E-cadherin [126]. In addition, overexpression of miR-494 was shown to reduce activating transcription factor 3 (ATF3), leading to an increase in inflammatory mediators, such as IL-6 and monocyte chemotactic protein-1, exacerbating inflammation in I/R-induced mouse AKI [45]. Injection of miR-494 antagomir, chemically modified miR-494 antagonist, ameliorated LPS-induced mouse AKI via anti-apoptosis and anti-inflammation mechanisms by regulating the NF-κB signaling pathway [127]. It was reported that serum miR-106a was increased in sepsis-induced AKI mice, and miR-106a was suggested to target thrombospondin 2 (THBS2), leading to inflammation and apoptosis [128]. miR-155 inhibitor ameliorated LPS-induced mouse AKI through the reduction of inflammatory cells by regulating the target Suppressor Of Cytokine Signaling 1 (SOCS1) and Signal Transducer And Activator Of Transcription (STAT)1 mRNAs [129]. It was also reported that macrophage-derived exosomal miR-155 promoted tubular injury by targeting SOCS1 [130]. In addition, miR-155^-/- mice were made resistant to cisplatin-induced AKI by reducing tubular cell apoptosis [130]. miR-155 was shown to be up-regulated in rat I/R-induced AKI and suggested to promote kidney injury and apoptosis by targeting Transcription factor 4 (TCF4)/Wnt/β-catenin signaling pathway [131]. miR-152-3p was suggested to promote cell apoptosis by silencing Sirtuin 7 (SIRT7) in I/R-induced rat AKI [132]. miR-152-3p was shown to promote sepsis-induced rat AKI by targeting ERBB receptor feedback inhibitor 1 (ERRFI1), leading to an increase in STAT3 expression, resulting in promoting cell apoptosis and inflammation [133]. miR-709 antagomir injection attenuated cisplatin-induced mouse AKI via a reduction in mitochondrial dysfunction by regulating miR-709 target gene mitochondrial transcriptional factor A (TFAM) [134].

Furthermore, similarly to miR-21, several miRNAs were shown to mediate both protective and pathogenic effects under AKI. miR-34 agomir injection ameliorated sepsis-induced mouse AKI via anti-inflammatory mechanism by targeting ubiquitin-like protein 4A (UBL4A) [135]. miR-34a induced via p53 was suggested to play a cytoprotective role in cell survival in cisplatin-induced mouse AKI [136]. In contrast to these renoprotective effects of miR-34, miR-34 was also shown to promote kidney injury under AKI. lncRNA HOX transcript antisense RNA (HOTAIR) overexpression ameliorated sepsis-induced rat AKI via an anti-apoptosis mechanism by targeting miR-34a and regulating its target B-cell lymphoma-2 (Bcl-2) [137]. It is also reported that increased miR-34a promoted acetylation of FOXO3 by repressing Sirtuin 1 (SIRT1), leadind to p53 activation and cell apoptosis in the cisplatin-induced mouse AKI model [119]. miR-34a was also suggested to suppress autophagy in kidney tubular cells by targeting autophagy related 4B cysteine peptidase (ATG4B) in I/R-induced mouse AKI [138]. miR-34a mimics prevented Nicotinamide phosphoribosyltransferase (NAMPT) expression, which suggest that they affected oxidized NAD (NAD⁺) metabolism, leading to kidney dysfunction in I/R-induced AKI mice [139]. In contrast to the anti-apoptosis effect of miR-125b-5p from MSCs in I/R-induced AKI mice [111], miR-125 was also shown to disrupt mitochondrial dynamics by targeting modulate mitofusin1 (MFN1) in cisplatin-induced AKI mice, where anti-miR-125b treatment reduced cisplatin-induced mitochondrial fragmentation and kidney injury [140]. miR-150-5p agomar was shown to ameliorate sepsis-induced mouse AKI via an anti-apoptosis mechanism by targeting mitogen-activated protein kinase kinase kinase 3 (MEKK3)/JNK pathway [141]. On the other hand, it was also reported that deletion of miR-150 in mice protected against myocardial-infarction-induced AKI through anti-apoptosis and anti-fibrosis mechanism by targetinginsulin-like growth factor-1 receptor (IGF-1R) [142]. In addition, pretreatment with exosomes enriched in miR-150 worsened kidney fibrosis in I/R-induced AKI mice [143]. Similarly, kidney fibrosis was reduced by miR-150-5p-deficient tubular cell-derived exosome in I/R-induced AKI mice by regulating the miR-150 target gene, SOCS1 [144]. miR-214 is also shown to mediate both protective and pathogenic effects under AKI. Regarding renoprotective effects, miR-214 injection in I/R-induced mice ameliorated kidney injury by inhibiting apoptosis through targeting dickkopf WNT signaling pathway inhibitor 3 (Dkk3) [145]. Adenovirus-mediated miR-214 treatment inhibited excess autophagy through regulation of the PTEN/AKT/mTOR pathway, thereby limiting kidney injury in sepsis-induced AKI mice [146]. On the other hand, regarding pathogenic effects, it is reported that kidney proximal-tubular-cell-specific miR-214 knockout mice showed less kidney damage and less apoptosis after I/R-induced AKI by targeting mitofusin-2 (Mfn2) [147]. miR-214-5p antagomir ameliorated LPS-induced kidney inflammation and oxidative stress, while miR-214-5p agomir aggravated kidney injury, presumably by targeting glucagon-like peptide-1 receptor (GLP-1R) [148]. miR-214-30 antagomir protected against cisplatin-induced AKI in mice by inhibiting tubular cell ferroptosis by targeting Glutathione Peroxidase 4 (GPX4) [149]. Collectively, miRNAs have several gene targets, thus regulating a variety of mechanisms that may mediate protective and/or pathogenic effects under AKI, and it may depend on the etiologies of AKI and evaluating methods.

Table 2. Potential therapeutic targets of miRNAs for AKI.

Table 2. Potential therapeutic targets of miRNAs for AKI.

miRNA	Effect	Target	Model	Species	Function	Ref.
miR-21	Protective	PTEN/Akt/mTOR Pdcd4/NFκ-B	I/R	Mice	Anti-apoptosis Anti-inflammation	[71]
	Protective	MKK3	I/R	Mice	Anti-inflammation	[77]
	-	AKT/CDK2-FOXO1	Sepsis	Rats	Metabolism alteration	[80]
	Protective	CDK6	LPS	Mice	Anti-apoptosis	[75]
	Protective	CCR7	I/R	Mice	Anti-inflammation	[76]
	Pathogenic	Rab11a	I/R	Rats	Anti-autophagy	[78]
miR-30	Protective	DRP1	I/R	Rats	Anti-apoptosis	[81]
	Protective	M1-M2 macrophage transition	I/R	Rats	Anti-inflammation	[82]
miR-17	Protective	DR6	I/R	Mice	Anti-apoptosis	[93]
miR-5100	Protective	Apoptotic pathway	I/R	Mice	Anti-apoptosis	[55]
miR-187	Protective	AChE	I/R	-	Podocyte protection	[108]
miR-182	Pathogenic	-	I/R	Rats	-	[116]
	Pathogenic	TCF7L2/Wnt/β-catenin	I/R	Rats	Apoptosis	[117]
	Pathogenic	FoxO3	I/R	Rats	Apoptosis	[118]
miR-489	Protective	PARP1	I/R	Mice	Anti-apoptosis	[73]
miR-668	Protective	MTP18	I/R	Mice	Mitochondrial dynamics	[72]
miR-27	Protective	OSMR	I/R	Rats	PI3K/AKT signal	[103]
	Protective	TLR4	I/R	Rats	Anti-inflammation	[102]
miR-140	Protective	Nrf2	Cisplatin	Mice	Anti-oxidative stress	[101]
	Protective	CXCL12	I/R	Mice	Anti-inflammation	[100]
miR-125	Pathogenic	MFN1	Cisplatin	Mice	Mitochondrial damage	[140]
	Protective	P53	I/R	Mice	Anti-apoptosis	[111]
miR-122	Pathogenic	FoxO3	Cisplatin	Mice	Apoptosis	[119]
miR-150	Pathogenic	IGF-1R	Ischemic	Mice	Apoptosis Fibrosis	[142]
	Pathogenic	-	I/R	Mice	Fibrosis	[143]
	Pathogenic	SOCS1	I/R	Mice	Fibrosis	[144]
	Protective	MEKK3/JNK	LPS	Mice	Anti-apoptosis	[141]
miR-218	Pathogenic	HO-1	Sepsis	Mice	Apoptosis	[125]
miR-126	Protective	-	I/R	Mice	Vascular protection	[107]
miR-195	Protective	VEGFA	I/R	Rats	Anti-inflammation Anti-oxidative stress	[99]
miR-181	Protective	KLF6	I/R	Mice	Anti-apoptosis Anti-inflammation	[83]
	Protective	PTEN	Cisplatin	Mice	-	[84]
	Protective	GJB2	LPS	Mice	Anti-apoptosis	[85]
	Protective	NEK7	Sepsis	Mice	Anti-pyroptosis	[86]
miR-301	Pathogenic	-	Vancomycin	Mice	Apoptosis	[120]
miR-709	Pathogenic	TFAM	Cisplatin	Mice	Mitochondrial damage	[134]
mR-375	Pathogenic	HNF1b	Cisplatin	Mice	Apoptosis	[121]
miR-204	Protective	Hmx1	Candidemia	Mice	Anti-inflammation	[98]
miR-211	Protective	Hmx1	Candidemia	Mice	Anti-inflammation	[98]
miR-590	Protective	TRAF6	LPS	Mice	Anti-apoptosis Anti-inflammation	[89]
miR-152	Pathogenic	ERRFI1	Sepsis	Rats	Inflammation Apoptosis	[133]
	Pathogenic	SIRT7	I/R	Rats	Apoptosis	[132]
miR-155	Pathogenic	SOCS1/STAT1	LPS	Mice	Inflammation	[129]
	Pathogenic	TCF4/Wnt/β-catenin	I/R	Rats	Apoptosis	[131]
	Pathogenic	SOCS1	I/R	Mice	Tubular injury	[130]
	Pathogenic	-	Cisplatin	Mice	Apoptosis	[150]
miR-106	Pathogenic	THBS2	Sepsis	Mice	Inflammation Apoptosis	[128]
miR-22	Protective	AIFM1	Sepsis	Mice	Anti-apoptosis	[87]
	Protective	HMGB1	Sepsis	Rats	Anti-inflammation	[88]
miR-107	Protective	RPS19	I/R	Rats	Anti-apoptosis	[112]
	Pathogenic	DUSP7	Sepsis	Mice	Tubular injury	[113]
miR-290	Protective	CCL2	Sepsis	Mice	Anti-apoptosis	[97]
miR-34	Pathogenic	NAMPT	I/R	Mice	NAD depletion	[139]
	Pathogenic	Atg4B	I/R	Mice	Reduce autophagy	[138]
	Pathogenic	Bcl2	Sepsis	Rats	Apoptosis	[137]
	Pathogenic	SIRT1	Cisplatin	Mice	Apoptosis	[119]
	Protective	UBL4A	Sepsis	Mice	Anti-inflammation	[135]
	Protective	-	Cisplatin	Mice	Cytoprotective	[136]
miR-188	Pathogenic	SRSF7	Contrast	Rat	Apoptosis	[122]
miR-214	Pathogenic	Mfn2	I/R	Mice	Apoptosis	[147]
	Pathogenic	GLP-1R	LPS	Mice	Inflammation	[148]
	Pathogenic	GPX4	Cisplatin	Mice	Ferroptosis	[149]
	Protective	PTEN/AKT/mTOR	Sepsis	Mice	Autophagy regualation	[146]
	Protective	Dkk3	I/R	Mice	Anti-apoptosis	[145]
miR-124	Protective	PARP1	I/R	Mice	Anti-necroptosis	[92]
	Protective	-	I/R	Mice	Anti-apoptosis	[91]
miR-146	Protective	IRAK1	I/R	Rats	Anti-inflammation	[104]
	Protective	-	I/R	Mice	Anti-inflammation	[105]
	Protective	Tfdp2	Cisplatin	Mice	Anti-fibrosis	[115]
miR-687	Pathogenic	-	LPS	Mice	Apoptosis	[123]
miR-24	Pathogenic	H2A.X/HO-1	I/R	Mice	Apoptosis	[124]
miR-494	Pathogenic	ATF3	I/R	Mice	Inflammation	[45]
	Pathogenic	E-cadherin	I/R	Mice	Apoptosis	[126]
	Pathogenic	NF-κB signaling	LPS	Mice	Inflammation Apoptosis	[127]
miR-223	Protective	NLRP3	I/R	Mice	Anti-apoptosis	[110]
miR-15	Protective	CX3CL1	I/R	Rats	Anti-inflammation Anti-fibrosis	[114]
miR-16	Protective	CX3CL1	I/R	Rats	Anti-inflammation Anti-fibrosis	[114]
	Pathogenic	-	I/R	Mice	Anti-apoptosis	[41]
miR-199	Protective	Sema3A/AKT/ERK	I/R	Mice	Anti-apoptosis	[109]
miR-210	Protective	VEGF pathway	I/R	Mice	Angiogenesis	[106]
miR-486	Protective	PTEN	I/R	Mice	Anti-apoptosis	[95]
miR-424	Protective	DR6	I/R	Mice	Anti-apoptosis	[94]
miR-191	Protective	OXSR1	Sepsis	Rat	Anti-apoptosis	[96]

miRNA: microRNA; I/R: ischemia-reperfusion; LPS: lipopolysaccharide.

5. Conclusions

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