Significance of microRNAs in Cardiovascular Diseases: Comparison
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The microRNAs (miRNAs), lncRNAs (long ncRNAs), and circRNAs (circular RNAs) with significant regulatory and structural roles make up approximately 99% of the human genome, which does not contain proteins. Non-coding RNAs (ncRNA) have been discovered to be essential novel regulators of cardiovascular risk factors and cellular processes, making them significant prospects for advanced diagnostics and prognosis evaluation. Cases of cardiovascular diseases (CVDs) are rising due to limitations in the existing therapeutic approach; most of the treatment options are based on the coding transcripts that encode proteins. Various investigations have shown the role of nc-RNA in the early diagnosis and treatment of CVDs. Furthermore, the development of novel diagnoses and treatments based on miRNAs, lncRNAs, and circRNAs could be more helpful in the clinical management of patients with CVDs. 

  • cardiovascular disease
  • microRNAs
  • diagnosis
  • long noncoding RNA
  • ncRNAs
  • therapy

1. Introduction

Cardiovascular diseases (CVDs) are caused by a reduced flow of oxygenated blood in the human body. CVDs are classified into various types of heart diseases, including cardiac hypertrophy (CH), heart failure (HF), rheumatic heart disease (RHD), acute coronary syndrome (ACS), myocardial infarction (MI), atherosclerosis (AS), myocardial fibrosis (MF), arrhythmia (ARR), and pulmonary arterial hypertension (PAH) [1][2]. CVDs are the major cause of death worldwide, according to the World Health Organisation; 17.9 million deaths were reported in 2019, accounting for 32% of all deaths at the global level. Early diagnosis and targeted treatment of CVDs remain challenges [3][4]. Current treatment options are available, with their limitations, and can reduce disease development. Novel treatment options are required to target cellular events during disease progression to facilitate the timely management of the patient’s clinical conditions [5]. Multiple coding genes are involved in the development of CVDs. In recent years, it has been discovered that non-coding RNA (ncRNA) regulates disease development [6]. The ncRNAs are 200-nt base sequences that regulate the genetic, epigenetic, and cell signalling mechanisms, as well as gene expression [7]. ncRNAs are used as biomarkers for diagnosis and treatment due to their involvement in disease severity. In recent years, ncRNA has been investigated in CVDs [8]. ncRNAs have great importance in clinical applications and are classified into various categories, including miRNAs, lncRNAs, and circRNAs [9]. SiRNAs can be used to target ncRNAs. RNAi-mediated siRNAs are highly adaptable and are used to silence their mRNA’s protein-encoding gene. MiRNAs are 22-nucleotide RNA molecules that regulate cell signalling and downregulate the expression of specific genes by modifying the translation process [10]. Researchers herein discuss the biological and clinical importance of miRNAs, lncRNAs, and circRNAs and their expression profiles and manipulation of non-coding transcripts in CVDs, which will deliver an in-depth knowledge of the role of ncRNAs in CVDs for progressing new clinical diagnosis and treatment.

2. miRNAs and CVDs

The miRNA lin-4 was identified in 1993 in Caenorhabditis [11]. The synthesis of miRNA occurs in the nucleus and is transcribed by RNA polymerase II into coding and noncoding, capping polyadenylated pri-miRNAs. The pre-miRNA produces a hairpin-like structure and is timed by the Drosha nuclear enzyme, then transported to the cytoplasm [12]. The DICER removes the terminal loop of the pri-miRNAs, resulting in a 20-25 nucleotide base pair dsRNA complex. The dsRNA, as attached to the miRNA-linked RISC (RNA-induced silencing complex), targets mRNAs and results in mRNA de-adenylation and translational repression (Figure 1) [13]. In vivo and in vitro studies have shown that miRNA plays a critical role in the regulation of CVDs such as CH, HF, ARR, ACS, MI, AS, RHD, and PAH (Table 1) [14].
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Figure 1. Synthesis and mechanism of miRNA.
Table 1. Regulations of miRNA and their clinical importance in CVDs.
S.N. Type of Disease miRNA Regulation Importance Reference
1 Cardiac hypertrophy (CH) miR-208a Up-regulation Cardiac remodelling [13]
2 CH miR-19a/b Up-regulation Cardiac remodelling in response to angiotensin II infusion [14]
3 CH miR-155 Up-regulation Cardiac remodelling [15]
4 CH miR-199a Up-regulation Maintenance of cell size in cardiomyocytes [16]
5 CH miR-1, Down-regulation Induces cardiac hypertrophy. [17]
6 CH miR-101 Down-regulation Inhibit cardiac hypertrophy signalling [18]
7 CH miR-185 Down-regulation Inhibit CH hypertrophy signalling [19]
8 CH miR-34a Down-regulation Regulation of Ang II-induced cardi myocyte hypertrophy [20]
Cardiomyocyte


Downstream Targets: Pdk4, Sgk1
[
56
]
74
MI
miR-21
Down-regulated
Fibroblast, Downstream Targets Pten; Sprouty-1, collagens
[
57
]
75
MI
miR-24
Up-regulated
Anti-apoptosis in Cardiomyocyte, fibroblast, endothelial cell;


Downstream Targets Bim; Furin; Gata2, Pak4
[
58
]
76
MI
miR-29
Down-regulated
Cardiomyocyte, fibroblast


Downstream Targets: Mcl-1; Collagens
[
59
]
77
MI
miR-92a
Up-regulated
Endothelial cell


Downstream Targets: Itga5 [60]
78 MI miR-101 Down-regulated Cardiac remodelling

Downstream Targets: Collagens		 [61]
79 MI miR-126 Down-regulated Protects against myocardial ischemia-reperfusion injury [62]
Myocardial hypertrophy (MH) is caused by the development of CVDs, including stenosis of the heart valve and hypertension, and causes HF and death [71]. Several miRNAs, including miR-208a, miR-19a/b, miR-34a, miR-145, miR-150, miR-378, and others, are involved in the development of MH [72]. MiR-378 is an anti-MH miRNA and regulates the Igf1r (insulin-like growth factor receptor), Grb2 (growth factor receptor binding protein 2), and Ksr1 (Ras kinase inhibitor 1) [73]. MiR-185 regulates cardiac cell proliferation and is related to the signal transduction mechanism. MiR-34a regulates the Agt9a gene, which is involved in autophagy. The transcription activator p300 is regulated by miR-150 [74]. MiR-1 is involved in the growth and development of cardiomyocytes by reducing the expression of GATA-binding protein 4 (GATA4) and calmodulin Mef2a, which regulate the calcium signal pathway and protein expression and could be targeted for diagnosis and therapy [75].

2.1. miRNAs and HF

HF is caused by a failure of the regulatory mechanism in the heart [76]. Many different forms of miRNA, such as miR-320a, miR-423-5p, miR-200b, miR-622, miR-1228, miR-208b, miR-499, miR-223, miR-1254, miR-1306, miR-18a, miR-26b, miR-27a, miR-30e, miR-106a, miR-199a, are crucial in the development of HF conditions [77]. Early hypertrophic growth in the left ventricle may be caused by miR-125b and lead to HF. The expression of brain natriuretic peptide (BNP) is regulated by miR-200b, miR-622, and miR-1228. HF may also be caused by increased expression of miR-208b and miR-499. These miRNA regulations in HF could be targeted for diagnosis and therapeutic approaches [78].

2.2. Arrhythmias

Arrhythmias (AR) are mainly caused by imbalances of the ion channel and dysregulations of conduction in cardiac muscles. Atrial fibrillation (AF) is a severe AR observed in CVDs that can lead to HF, stroke, and death [79]. There are various types of miRNA involved in the development of AR in CVD patients, including miR-664, miR-133, miR-590, miR-130a, miR-21, miR-208b, miR-483, miR-1, and miR-150. In addition, the AF is controlled by the miRNAs miR-328, miR-2, miR-664, miR-483, miR-133, miR-1, miR-208b, miR-590, miR-328, and miR-223 [80]. The overexpression of miR-130a is linked with cx43 (protein connexin 43). MiR-150 regulates the platelet count in patients with AF, which plays a major role in fibrosis and inflammation and is involved in the development of AF [81].

2.3. miRNAs and ACS and MI

ACS (acute coronary syndrome) is developed by reduced blood flow in the heart, an immediate blockage of the coronary arteries, and localized heart necrosis, all contribute to the development of AMI (acute myocardial infractions) [82]. AMI patients have a high level of miR-1 expression. MiR-1, miR-133a, and miR-208a levels have been found to be higher in AMI patients. Cardiac arrest is regulated by miR-208b and miR-499-5p in patients with coronary artery bypass grafting [83]. These two miRNAs are expressed by dysregulated cardiac muscles. A reduced level of expression has been shown in AMI patients. The expression profile of all these miRNA regulations can be used for early diagnosis and treatment [84]. High miR-208 expression levels have been observed in a mouse model with AMI. High-throughput analysis of miRNA expression in patients with AMI can be explored further for sensitive and specific early diagnosis and treatment [85].

2.4. miRNAs and Atherosclerosis

The miRNA plays an important role in the generation of atherosclerosis by vascular angiogenesis, endothelial dysfunction, lipid accumulation, local inflammation, calcification, thrombosis, and endothelial dysfunction [86]. Play important in the development of CAD (coronary artery disease), causes significant death at global level. Expression profile of miRNA has been investigated in patients with AS. MiR-33 regulates the AS disease progression by involving the inflammatory response, cell cycle progression, lipid metabolism, and proliferation [87]. In patients with AS, miR-122 is substantially expressed. MiR-122 controls the levels of high-density lipoprotein (HDL) and low-density lipoprotein (LDL).Leukocyte aggregation on endothelial cells is triggered by miR-126-mediated upregulation of VCAM-1 (vascular cell adhesion molecule-1) [88]. Mi-R1 regulates the signalling pathways for MLCK (Myosin Light Chain Kinase) and ERK (Extracellular Signal-Regulated Kinase). MiR-221 and miR-222 control the growth and development of vascular smooth muscle cells (VSMCs). In patients with AS, there is generally less miR-126, miR-1, and miR-221/222 expression [89].

2.5. miRNAs and RHD

RHD lesions have primarily been found in the mitral valve. RHD tissue and plasma samples have significant levels of miRNA-1299 and miRNA-1183 expression. MiR-328-3p is found in RHD and AF (atrial fibrillation) patients [90]. MiRNA-432 expression levels have been found to be lower in RHD patients. All these microRNAs could be used for early diagnosis. Further investigations are needed to find out more about miRNA regulations in RHD [91].

2.6. LncRNAs and Cardiovascular Diseases

LncRNAs are more complex and heterogeneous in nature in comparison to miRNAs, which regulate gene expression. LncRNAs are involved in CVDs and categorized into various classes based on their structure and functions, including bidirectional lncRNAs, enhancer lncRNAs, sense lncRNAs, antisense lncRNAs, intergenic lncRNAs, and intron lncRNAs [92]. The gene expression level is changed by the interactions of lncRNA with DNA, RNA, proteins, elements of the chromatin modification complex, and transcription factors. Guided lncRNAs can either activate lncRNA processes or suppress gene expression by delocalizing regulatory elements [93][94][95]. Ribonucleoprotein (RNP) complex formation involves the scaffold lncRNAs (Figure 2). The lncRNAs serve as primary miRNA precursors that are converted into mature miRNAs while the miRNA precursor is suppressed. Long-range gene regulation begins when the lncRNA activates transcription from regulatory areas of the genome. LncRNAs interact with miRNAs and disrupt the RNA molecules’ regulatory system (Figure 3) [93]. lncRNAs also act as a maternal or paternal genomic imprinting expression and help in the development of organisms [96][97]. Regulations and clinical importance of lncRNA in cardiovascular are shown in Table 2.
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Figure 2. lncRNA mechanisms of action. (A) Guide lncRNAs activate or repress gene expression through relocalization of regulatory factors. (B) Scaffold lncRNAs aid in the formation of Ribonucleoprotein (RNP) complexes. (C) Decoy lncRNAs remove the regulatory factor bound to the genome, thereby terminating its regulation. (D) lncRNAs sponge the miRNAs, thus inhibiting the miRNA-mediated gene repression. (E) miRNA precursor lncRNAs function as primary miRNA precursors that are processed into mature miRNAs. (F) lncRNA transcription from regulatory regions of the genome initiates long-range gene regulation.
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Figure 3. lncRNA-miRNA Gene Expression: Effect of lncRNA expression changes on CVDs depends on canonical function of miRNA target gene.
Table 2. Regulations of lncRNA in cardiovascular disorders and their clinical significance.
  Type of Disease lncRNA Regulations Importance References
1 Myocardial infraction (MI) aHIF Regulations of the angiogenesis process and a biomarker Inhibits the autophagy of cardiac cells during MI [98]
2 MI ANRIL Regulates myocardial cell apoptosis in AMI Protection of cardiomyocytes [99]
3 MI APF APF lncRNA regulates autophagy Acting as a sponge for miRNA-

188-3p.		 [98]
4 MI CARL Regulates mitochondrial fission and apoptosis Acting as a sponge for

miRNA-539.		 [98]
5 MI CDR1AS Inhibiting the autophagy of cardiac cells during MI Biomarker. [98]
6 MI FTX Regulates cardiomyocytes Act as a sponge for miRNA-

29b-1-5.		 [100]
7 MI GAS5 Regulates the protection of cardiomyocytes against hypoxic injury Act as a sponge

for miRNA-142; improves apoptosis by

negatively regulating sema3a.		 [98]
8 MI H19 Regulates autophagy Induction of cardiac remodeling, autophagy, and biomarker. [98]
9 CH miR-145 Down-regulation Inhibits isoproterenol-induced cardiomyocyte hypertrophy
9 MI HOTAIR[ Regulates cardioprotective21]
Act as a sponge for miRNA-1 and as a

biomarker. [101] 10 CH miR-150 Down-regulation Reduces the immunosuppression function of Myeloid-derived suppressor cells (MDSCs) [22]
10 MI KCNQ1OT1 Down-regulation of lncRNA KCNQ1OT1 protects against myocardial ischemia/reperfusion injury Biomarker for left ventricular dysfunction. [102] 11 CH miR-378 Down-regulation Act as negative regulator for CH [23]
11 MI LIPCAR Down-regulated Biomarker for cardiac remodelling. [103] 12 Heart failure (HF) miR-125b Up-regulation Conduction of Cardiac fibrosis (CF) [24]
12 MI Lnc-Ang362 Upregulation Promotion of CF [104] 13 HF miR-22, Up-regulation Regulator for cardiac remodelling  
  MI MALAT1 Down-regulation Regulation of cardiomyocytes apoptosis and autophagy through miRNA-

558; and biomarker.		 [105] 14 HF miR-92b Up-regulation Related to the left atrium diameter, left ventricular end-diastolic dimension [25]
15
13 MI MDRL Regulates mitochondrial fission Reduction of mitochondrial fission and apoptosis acting as a sponge for

miRNA-361.		 [106] HF miR-320a Up-regulation CF through activation of the IL6/STAT3 axis. [26]
14 MI MEG3 Regulates cardiomyocytes Regulation of cardiomyocytes apoptosis. [107] 16 HF miR-423-5p Up-regulation Upregulated in human failing myocardium [27]
15 MI MHRT Regulates cardiomyocytes Regulation of cardiomyocytes apoptosis and biomarker. [108] 17 HF miR-200b Up-regulation Regulation of multiple cellular pathways in HF
16 MI MIAT[ Regulates CF28]
Regulation of cardiac hypertrophy and fibrosis acting as a sponge for

miRNA-150 and -93. [98] 18 HF miR-622 Up-regulation Improves blood vessel growth [29]
17 MI Mirt1/2 Regulates cardiomyocytes Regulation of cardiac remodelling. [98] 19 HF miR-1228 Up-regulation Marker for systolic HF [30]
18 MI n379519 Regulates CF Promotion of cardiac fibrosis through miRNA-30. [109] 20 HF miR-208b Up-regulation pathogenesis of DCM [31]
21 HF miR-499 Up-regulation Cardiac development [32]
22 HF miR-223 Up-regulation Altered in post-MI HF in humans [33]
23 HF miR-1254 Up-regulation Altered in post-MI HF in humans [34]
24 HF miR-1306 Up-regulation Elected to explore novel circulating markers for HF [35]
25 HF miR-18a Down-regulation CF through the Notch2 pathway. [36]
26 HF miR-26b Down-regulation Controlling critical signalling pathways, such as BMP/ SMAD1 signalling [37]
27 HF miR-27a Down-regulation Inhibiting miR-27a-3p mitigated CH phenotype induced by Ang II (Angiotensin -II) [38]
28 HF miR-30e Down-regulation The overexpression of miR-30c reduces the level of connective tissue growth [39]
29 HF miR-106a Down-regulation Notch 3 pathway in ischemic heart injury. [40]
30 HF miR-199a Down-regulation Improves contractile function [41]
31 HF miR-652 Down-regulation Marker for predicting acute coronary syndrome [42]
32 HF miR-1 Down-regulation Systolic HF [43]
33 HF miR-126 Down-regulation Activation of the vascular endothelial growth factor (VEGM) signalling pathway in the endothelium. [44]
34 HF miR-423 Down-regulation It is a circulating biomarker for heart failure. [45]
35
19 MI NONRATT021972 Regulates cardiomyocytes Promotion of cardiac function. [110]
20 MI NRF Regulates cardiomyocyte necrosis. Regulation of cardiomyocyte necrosis. [111]
21 MI NRON Up-regulated Wisper in cardiac fibroblast; Biomarker [98]
22 MI PCFL Up-regulated Promotion of cardiac fibrosis through miRNA-378. [98]
23 MI TTTY15 Up-regulated Induction of cardiomyocyte injury by hypoxia targeting miRNA-455. [98]
24 MI UCA1 Regulates cardiomyocytes Regulated ischemia and hypoxia of cardiomyocytes; Biomarker. [111]
25 MI UIHTC Regulates cardiomyocytes against MI Promotion of mitochondrial function. [98]
26 MI Wisper Regulates cardiac fibroblast MI-induced fibrosis and cardiac dysfunction [112]
27 MI ZFAS1 Regulates cardiomyocyte Induction of cardiomyocyte apoptosis, cardiac contractility reduction, and

biomarker.		 [113]
28 Coronary heart disease aHIF Up-regulated Biomarker. [114]
29 Coronary heart disease ANRIL Down-regulates Diagnostic and prognostic indicator for CHD [114]
30 Coronary heart disease APOA1-AS Up-regulations increase the risk of CHD Biomarker. [115] Cardiac electrical and structural remodelling (CE and SR) miR-1
31 Coronary heart disease AWPPH Regulates apoptosis. Promotion of ECs apoptosis. [116]
32 Coronary heart disease BACE1-AS dysregulation dysregulation of the BACE1/BACE1-AS/Aβ axis is associated with HF. [Down-regulated Increased altered conduction

Increased CF		 [46]
117]
33 Coronary heart disease BANCR Differentially expressed Promotion of VSMCs proliferation and migration. [118]
34 Coronary heart disease CHROME Up-regulated Regulation of cellular cholesterol homeostasis [119] 36 CE and SR miR-26 Down-regulated Increase inwardly rectifying channel [47]
35 Coronary heart disease CoroMarker Differentially expressed novel biomarker for the diagnosis [120] 37 CE and SR miR-29 Down-regulated Increased CF [48]
36 Coronary heart disease EGOT Differentially expressed Biomarker. [121] 38 CE and SR miR-30 Down-regulated Increased CF [49
37]
Coronary heart disease H19 Differentially expressed Biomarker. [122] 39 CE and SR miR-133 Down-regulated Increased CF
38 Coronary heart disease[ HOTTIP50]
Up-regulates Promotes ECs proliferation and migration [123] 40 CE and SR miR-328 Up-regulated Shortened atrial action potential duration by targeting [51]
39 Coronary heart disease HRCR Regulates hypertrophic Ca2+ signaling pathway Regulation of cardiomyocytes apoptosis and proliferation. [124] 41 CE and SR miR-499 Up-regulated Altered conduction
40

by targeting [52]
Coronary heart disease LIPCAR Differentially expressed Biomarker. [103] 42 CE and SR miR-21 Up-regulated Inhibition of fibroblast proliferation
41 Coronary heart disease[ lincRNA-p2153]
Regulates cardiac remodelling and heart failure Regulation of cardiomyocytes apoptosis and proliferation. [125] 43 Acute coronary syndrome (ACS) and myocardial infarction (MI) miR-1 Up-regulated
42marker of cardiomyocyte injury Coronary heart disease[54]
LINC00968 Up-regulated Promotion of ECs proliferation and migration acting as a sponge for

miRNA-9 [126] 44 ACS and MI miR-133a Up-regulated Development of VF (Ventricular fibrillation) [55]
43 Coronary heart disease MALAT1 Differentially expressed Biomarker. [127] 45 ACS and MI miR-208a Up-regulated Regulates the cardiac stress response. [56
44]
Coronary heart disease MIAT Differentially expressed Biomarker [128] 46 ACS and MI miR-499-5p Up-regulated Associated with cardiac injury and also with cardio protection [57]
45 Coronary heart disease NEXN-AS1 Differentially expressed Mitigation of atherosclerosis. [129] 47 ACS and MI miR-126, Down-regulated downregulated in the region adjacent to MI areas [58]
46 Coronary heart disease SMILR Differentially expressed Biomarker. [130] 48 ACS and MI miR-221/222 Down-regulated Severity of the coronary artery lesions [59]
47 Arterial Hypertension AK098656 Up-regulated Regulation of arteries of resistance and a biomarker [131] 49 ACS and MI miR-29 Dysregulation Involved in CF multiple collagens, fibrillin’s, and elastin [60]
48 Arterial Hypertension ANRIL Regulates endothelial cell activities Increase of susceptibility to higher systolic blood pressure conferred by

polymorphisms.		 [132] 50 ACS and MI miR-145 Up-regulated Significantly upregulated in mice in response to chronic hypoxia and that genetic ablation
49 Arterial Hypertension[ GAS561]
Regulates ECs and VSMCs function Regulation of ECs and VSMCs function acting as endogenous RNA

competing of miRNA-21; and a biomarker. GAS5 Targets miR-194-3p. miR-194-3 [133] 51 ACS and MI miR-21 Up-regulated Up-regulated in the hypoxia [62]
50 Arterial Hypertension Giver Regulates VSMCs dysfunction. Promotion of VSMCs dysfunction. [134] 52 ACS and MI miR-206 Up-regulated Normal and hypertensive mouse PASMCs. [63]
51 Arterial Hypertension Lnc-Ang362 Regulates VSMCs Regulation of VSMCs proliferation through miRNA-221 and -222. 53 ACS and MI miR-328 Down-regulated Regulates Hypoxic Pulmonary Hypertension [64]
[135] 54 Pulmonary arterial hypertension (PAH) miR-204 Down-regulated Hypoxia related to pulmonary arterial hypertension [65]
55 Atherosclerosis (AS) miR-33 Dysregulation Promising strategy to reverse autophagy dysfunction in atherosclerosis. [66]
56
52 Arterial Hypertension NR_027032 Differentially expressed Biomarker. [136]
53 Arterial Hypertension NR_034083 Differentially expressed Biomarker. [137]
54 Arterial Hypertension NR_104181 Differentially expressed Biomarker. [136](AS) miR-122 Up Significantly up-regulated in patients with atherosclerotic lesion [67]
55 Heart failure ANRIL Differentially expressed Biomarker. [138] 57 (AS) miR-126 Down-regulated   [68]
56 Heart failure BACE1-AS Regulates apoptosis. Promotion of ECs apoptosis. 58 (AS) miR-1 Down-regulated Downregulation of miR-10a enhances IκB/NF-κB activation [69]
[139] 59 (AS) miR-221/222 Down-regulated Down-regulatesSuppression of PGC-1α (peroxisome proliferator-activated receptor gamma coactivator 1-alpha) in the progression of atherosclerosis Induction of Pathological cardiac remodelling.[70]
[141] 60 Congenital heart diseases (CHDs) miR-1275, miR-27b, miR-421 Up-regulated usually developing hearts [13]
61
59 Heart failure CHRF Up-regulated Endogenous sponge to miRNA-489 activity. [142] CHD
60miR-122, miR-1201 Down-regulated developing hearts [14]
Heart failure HEAT2 Up-regulated Biomarker. [143] 62 CHD miR-222, miR-337-5p, miR-363, miR-424, miR-424, miR-660, miR-708, miR-421, miR-19a, miR-130b, miR-146b-5p, miR-154, miR-155, miR-181c, miR-181d and miR-192, Up-regulated tetralogy of Fallot [15]
61 Heart failure HOTAIR 63 CHDs miR-181a, miR-720, miR-29c and miR-940 Down-regulated tetralogy of Fallot [16]
64 CHD miR-181c Up-regulated ventricular septal defect [17]
  CHD miR-1-1 Down-regulated ventricular septal defect [18]
65
57 Heart failure Chaer Dysregulation Induction of Pathological cardiac remodelling.Up-regulated LncRNA HOTAIR may function as a miR-19-sponge to modulate PTEN levels Biomarker. [144]
62 Heart failure LIPCAR Up-regulated Biomarker. [145]
63 Heart failure lincRNA-ROR Regulates CH Regulation of cardiac hypertrophy acting as a sponge for miRNA-133. [146]
64 Heart failure CHD miR-106a, miR-144, miR-451, miR-486-3p, miR-486-5p, hsa-let-7e, miR-16, miR-18a, miR-25, miR-93, and miR-505 Up-regulated transposition of the great arteries [19]
66 CHD miR-873 Up-regulated Cyanotic CHD [20]
[140]
58 Heart failure Chast LOC285194 Up-regulated overexpression suppressed MKN45 and HGC-27 cell proliferation and promoted cell apoptosis; Biomarker. [144]
65 Heart failure MEG3 Regulates CF Regulation of cardiac fibrosis and diastolic dysfunction [147]
66 Heart failure MHRT Regulates of chromatin re-modellers Regulation of chromatin remodels and biomarker. [148] 67 CHD miR-182 Down-regulated Cyanotic CHD [21]
67 Heart failure MIAT Regulates CH Regulation of cardiac hypertrophy acting as a sponge for miRNA-150. [149] 68 CHD
68miR-498 Heart failure NRONUp-regulated Ventricular septal defect  
Upregulated Biomarker. [150] 69 CHD miR-379-5p, miR-409-3p, miR-433, hsa-let-7e-5p, miR-155-5p, miR-222-3p, and miR-487b Down-regulated Ventricular septal defect [22]
69 Heart failure RNY5 Dysregulation Biomarker. [151] 70 CHD hsa-let-7b, hsa-let-7a, and miR-486 Up-regulated Atrioventricular septal defect and atrial septal defect [23]
71
70 Heart failure SOX2-OT Dysregulation Biomarker. [152] CHD miR-19b, miR-22, miR-29c, miR-375 Up-regulated Atrioventricular septal defect and atrial septal defect [22][23
71]
Heart failure SRA1 Dysregulation Biomarker. [153] 72 Myocrdial infraction (I) miR-1   Cardiomyocyte

Downstream Targets: Ncx-1; KCNJ2, GJA1; IGF-1		 [55]
73 MI miR-15 Up-regulated
The integration of various types of cells, the vascular system, and blood vessels are all involved in the generation of the heart [94]. lncRNAs, also known as super-enhancer lncRNAs (SE-lncRNAs), control transcription at the tissue and cell levels. MyoD is an important transcription factor that involves muscle cell differentiation along with other core transcription factors [95]. The CE (core enhancer element) is produced by CERNA, which acts as a positive feedback regulator. It has been recently observed that various types of lncRNA are involved in the development of CVDs, including CHRF, Myh7, LIPCAR, MIAT, Carl, LIPCAR, ASB9P1, RP11-218 M11.6, G078882, G064270, G000678, G030563, H19, TUG1, PFL, MIAT, AK081284, HOXA11-ASz, NRON, and GAS5 [92]. H19 is expressed during embryogenesis and CVD but is repressed after birth. miRNA-675 acts as a negative regulator in cardiac hypertrophy. miR-675-3p and miR-675-5p are upregulated in cardiac hypertrophy [96]. Some pro-hypertrophic factors are also involved in CH and are mediated by Ca/calmodulin-dependent protein kinase IIδ (CaMKIIδ). The lncRNA–miRNA–mRNA axis can be a potential target for therapeutic approaches [97]. All these investigations have confirmed that lncRNAs play major roles in cardiovascular biology and diseases (Table 23) [98].

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