
Figure 1
Canonical biosynthesis of miRNAs/mitomiRs (adopted from 29, 43, 45). Mature miRNA can be transported into any part of the cell; but miRNA/mitomiR regulation is possible only after incorporation into RISC. (AGO2 = argonaute 2; DGCR8 = DiGeorge critical region 8; EXP5 = exportin 5; GTP = guanosine triphosphate; IMP8 = importin 8; mRISC = RNA induced silencing complex loaded with mature miRNA; POLII = DNA polymerase II; RANGTP = binding nuclear protein RAN; RISC = RNA-induced silencing complex; TRBP2 = RISC-loading complex subunit TRBP2).
Table 1
Summary of microRNAs and their roles in mitochondria
| miR | miR accession number | Target genes | Gene accession number | Function | Functional pathway | Location | Species | References |
|---|---|---|---|---|---|---|---|---|
| miR-378 | MI0000795 | Crat | ENSMUSG00000026853 | Downregulation | Mitochondrial oxidative metabolism | Mitochondria in liver cells | Mouse | Carrer et al., 201259 |
| miR-24 | MI0000080 | H2ax | ENSMUSG00000049932 | Downregulation | Insulin pathway signaling | Mitochondria liver cells in | Human | Jeong et al., 201761 |
| miR-23b | MI0000439 | GLS | ENSG00000115419 | Downregulation | Glutamine metabolism | LMitochondria liver cells in | Human | Gao et al., 200960 |
| SLC2A1 | ENSG00000117394 | Downregulation | ||||||
| miR-1291 | MI0006353 | CPT1C ESRRA | ENSG00000169169 ENSG00000173153 | Downregulation Downregulation | Mitochondrial | Mitochondria in | Human | Yamasaki et al., 2013; Chen et al., |
| ASS1 | ENSG00000130707 | Downregulation | metabolism | renal cells | 2020, Tu et al., 202063, 64, 65 | |||
| GLUT1 | ENSG00000117394 | Downregulation | ||||||
| miR-138 | MI0000455 | PDK1 | ENSG00000152256 | Downregulation | Glucose metabolism | Mitochondria in cardiac cells | Human | Zhu et al., 201766 |
| miR-150 | MI0000920 MI0000479 | Slc2a4 SLC2A1 | ENSRNOG00000017226 ENSG00000117394 | Downregulation Downregulation | Metabolism | Mitochondria in cardiac cells | Rat Human | Ju et al., 202067 Li et al., 201768 |
| Slc2a4 | ENSRNOG00000017226 | Upregulation | Mitochondria in | |||||
| miR-199a | MI0000941 | Hk2 | ENSRNOG00000006116 | Upregulation | Expression of | muscle cells | Rat | Esteves et al., 2018, Yan et al., 2014, Guo |
| MI0000242 | HK2 | ENSG00000159399 | Upregulation | glucose transporters | Mitochondria in liver cells | Human | et al., 20154, 69, 70 | |
| miR-532-5p | MI0006154 | SlcHk22 a4 | ENSRNOGENSRNOG00000017226 00000006116 | Upregulation Upregulation | Expression glucose transporters of | Mitochondria muscle cells in | Rat | Esteves et al., 201870 |
| Mitochondrial | Mitochondria in | |||||||
| miR-29a | MI0000576 | Slc16a1 | ENSMUSG00000032902 | Downregulation | oxidative metabolism | pancreatic betacells | Mouse | Pullen et al., 201171 |
| miR-29b | MI0000143 | Slc16a1 | ENSMUSG00000032902 | Downregulation | Mitochondrial oxidative metabolism | Mitochondria in pancreatic betacells | Mouse | Pullen et al., 201171 |
| Mitochondrial | Mitochondria in | |||||||
| miR-124 | MI0000716 | Slc16a1 | ENSMUSG00000032902 | Downregulation | oxidative | pancreatic beta- | Mouse | Pullen et al., 201171 |
| metabolism | cells | |||||||
| CROT | ENSANAG00000028065 | Downregulation | ||||||
| CPT1A | ENSANAG00000017356 | Downregulation | ||||||
| HADHB | ENSANAG00000027802 | Downregulation | ||||||
| PRKAA1 | ENSANAG00000032687 | Downregulation | ||||||
| miR-33a/b | a-MI0002684, b-MI0007603 | ABCA1 | ENSANAG00000033387 | Downregulation | Lipid metabolism | Mitochondria in liver cells | Monkey | Rayner et al., 201172 |
| SREBF1 | ENSANAG00000021477 | Upregulation | ||||||
| FASN | ENSANAG00000032055 | Upregulation | ||||||
| ACLY | ENSANAG00000036009 | Upregulation | ||||||
| ACACA | ENSANAG00000035253 | Upregulation | ||||||
| miR-143 | MI0000916 | Map2k5 | ENSRNOG00000007926 | Downregulation | Adipogenesis | Mitochondria in adipose cells | Rat | Chen et al., 201473 |
| MI0000459 | APOL6 | ENSG00000221963 | Downregulation | Adpiogenesis | Mitochondria in adipose cells | Human | Ye et al., 201374 | |
| miR-204 | MI0000284 | ACACB | ENSG00000076555 | Downregulation | Lipid metabolism | Mitochondria adipose cells in | Human | Civelek et al., 201375 |
| miR-200 | MI0000737 | ZEBZEB1 2 | ENSGENSG00000148516 00000169554 | Upregulation Upregulation | Lipid metabolism | Mitochondria breast cells in | Human | Ahmad et al., 201176 |
| miR-338 | MI0000618 | COXIV | ENSRNOG00000007827 | Downregulation | Mitochondria oxidative metabolism | Mitochondria in neural cells | Rat | Aschrafi et al., 200877 |
| miR-181c | MI0000924 | COX1 | ENSRNOG00000034234 | Downregulation | Mitochondria oxidative metabolism | Mitochondria in cardiac cells | Rat | Das et al., 201288 |
| miR-210 | MI0000268 | ISCU | ENSG00000136003 | Downregulation | Mitochondria oxidative metabolism | Mitochondria in placenta cells | Human | Colleoni et al., 2013; Qiao et al., 201381, 85 |
| miR-15b | MI0000843 | Arl2 | ENSRNOG00000021010 | Downregulation | ATP production | Mitochondria in cardiac cells | Rat | Nishi et al., 201082 |
| Bcl2 | ENSRNOG00000002791 | Downregulation | ||||||
| miR-16 | MI0000844 | Bcl2 | ENSRNOG00000002791 | Downregulation | ATP production | Mitochondria in cardiac cells | Rat | Nishi et al., 201082 |
| Arl2 | ENSRNOG00000021010 | Downregulation | ||||||
| miR-195 | MI0000939 | Arl2 | ENSRNOG00000021010 | Downregulation | ATP production | Mitochondria in cardiac cells | Rat | Nishi et al., 201082 |
| miR-29a-3p | MI0000576 | Foxa2 | ENSMUSG00000037025 | Upregulation | Lipid metabolism | Mitochondria liver cells in | Mouse | Kurtz et al., 201486 |
| miR-19b | MI0000074 | MFN1 | ENSG00000171109 | Downregulation | Apoptosis | Mitochondria in bone cells | Human | Li et al., 201487 |
| miR-101-3p | MI0000103 | ATP5B | ENSG00000110955 | Silencing | Mitochondria metabolism | Mitochondria heLa cells in | Human | Zheng et al., 201184 |

Figure 2
The network of the mitomiRs and their target genes (grey rectangle) in primates (data from Table 1). Blue arrows present downregulation, green arrows present upregulation and black T-line present silencing. Purple octagon shape presents monkey miRNA and cyan hexagon presents human miRNAs (figure constructed with Cytoscape Network Data Integration, Analysis, and Visualization in a Box V3.8.2).

Figure 3
The network of the mitomiRs and their target genes (grey rectangle) in rodents (data from Table 1). Blue arrows present downregulation and green arrows present upregulation. Orange diamond shape presents rat miRNAs, yellow rectangle presents mouse miRNAs and cyan hexaon presents human miRNAs. miR-199a and miR-143 show that this two miRNAs regulate (figure constructed with Cytoscape Network Data Integration, Analysis, and Visualization in a Box V3.8.2).

Figure 4
Traditional cancer traits.89
Table 2
Summary of mitomiRs with roles in cancer
| miR | miR accession number | Target genes | Gene accession number | Function | Functional pathway | Type of cancer | Species | References |
|---|---|---|---|---|---|---|---|---|
| HIF-1 | ENSG00000258777 | Upregulation | ||||||
| ISCU | ENSG00000136003 | Upregulation | ||||||
| miR-210 | MI0000286 | COX10 | ENSG00000006695 | Upregulation | Hypoxia | Breast cancer, neck and head cancer, lung cancer | Human | Qin et al., 2014; Gee et al., 2010; Puissegur et al., 2011109, 110, 111 |
| SDHD | ENSG00000204370 | Upregulation | ||||||
| NDUFA4 | ENSG00000189043 | Upregulation | ||||||
| miR-200a | MI0000342 | TFAM | ENSG00000108064 | Downregulation | Mitochondrial biogenesis, cancer metabolism | Breast cancer | Human | Yao et al., 2014112 |
| miR-155 | MI0000681 | HK2 | ENSG00000159399 | Upregulation | phosphorylation Glucose | Breast cancer | Human | Jiang Fang et et al.al., , 20122012; 104 |
| miR-124 | MI0000443 | PKM | ENSG00000067225 | Upregulation | metabolism Glucose | Colorectal cancer | Human | Sun et al., 2012105 |
| miR-137 | MI0000454 | PKM | ENSG00000067225 | Upregulation | metabolism Glucose | Colorectal cancer | Human | Sun et al., 2012105 |
| miR-340 | MI0000802 | PKM | ENSG00000067225 | Upregulation | metabolism Glucose | Colorectal cancer | Human | Sun et al., 2012105 |
| miR-326 | MI0000808 | PKM2 | ENSG00000067225 | Downregulation | metabolism Glucose | Glioblastoma | Human | Kefas et al., 2010106 |
| miR-181-5p | MIMAT0000256 | RASSF6 INPP5A | ENSG00000169435 ENSG00000068383 | Downregulation Downregulation | Mitogen-activated protein kinase (MAPK) signaling pathway | Gastric cancer, cervical cancer | Human | Mi et al., 2017; Zhuang et al., 2017108, 113 |
| miR-92a-1 | MI0000093 | BCL2L11 | ENSG00000153094 | Downregulation | Apoptosis | Lymphoma | Human | Mogilyansky Rigoutsos, 2013 and 94 |
| PIK3R2 | ENSG00000105647 | Downregulation | Breast cancer cells | Zhu et al., 2011114 | ||||
| PLK2 | ENSG00000260410 | Downregulation | Acute leukaemia cells | Li et al., 2008115 | ||||
| EGFL7 | ENSG00000172889 | Downregulation | Oral squamous cells | Sasahira et al., 2012116 | ||||
| CRK | ENSG00000167193 | Downregulation | Lung cancer cells | Crawford 2008117 et al., | ||||
| ADAM9 | ENSG00000168615 | Downregulation | Melanoma cells cancer | Felli et al., 2013118 | ||||
| miR-126 | MI0000471 | HOXA9 | ENSG00000078399 | Downregulation | Inflammation, angiogenesis | Acute leukaemia cells | Human | Shen et al., 2008119 |
| IRS1 | ENSG00000169047 | Downregulation | Breast cancer cells | Zhang et al., 2008120 | ||||
| SOX-2 | ENSG00000242808 | Downregulation | Gastric cells cancer | Otsubo et al., 2011121 | ||||
| SLC7A5 | ENSG00000103257 | Downregulation | Lung cancer cells | Miko et al., 2011122 | ||||
| VEGFA | ENSG00000150630 | Downregulation | Oral squamous cells | Sasahira et al., 2012116 | ||||
| MMP7 | ENSG00000137673 | Downregulation | Melanoma cells cancer | Felli et al., 2013118 | ||||
| BCL-2 | ENSG00000171791 | Downregulation | ||||||
| Gao et al., 2010101 | ||||||||
| MCL-1 | ENSG00000143384 | Downregulation | ||||||
| COX4I2 | ENSG00000131055 | Downregulation | ||||||
| miR-15a | MI0000069 | COX6A2 | ENSG00000156885 | Downregulation | Apoptosis, ATP production | B-cell chronic lymphocytic leukemia | Human | Siengdee et al., |
| NDUFB7 | ENSG00000099795 | Downregulation | 2010102 | |||||
| NDUFV1 | ENSG00000167792 | Downregulation | ||||||
| NDUFS4 | ENSG00000164258 | Downregulation | ||||||
| COX4I2 | ENSG00000131055 | Downregulation | ||||||
| COX6A2 | ENSG00000156885 | Downregulation | ||||||
| miR-16a | MI0000070 | NDUFB7 | ENSG00000099795 | Downregulation | Apoptosis, ATP production | B-lymphocytic cell chronic leukemia | Human | Siengdee et al., 2010102 |
| NDUFV1 | ENSG00000167792 | Downregulation | ||||||
| NDUFS4 | ENSG00000164258 | Downregulation |