Table 2 Interaction between miRNAs and KRAS in the context of cancer (ANTs: adjacent normal tissues)

miR-217

Pancreatic ductal adenocarcinoma

Down

21 pairs of PDAC specimens and ANTs samples

PANC-1, MIAPaCa-2, AsPC-1 and BxPC-3 cells

KRAS, AKT

RAS signaling pathway

MiR-217 inhibited tumor cell growth, anchorage-independent colony formation and in vivo xenograft tumor growth

[26]

miR-96

Pancreatic cancer

Down

10 pairs of pancreatic cancer tissues and ANTs

Six-week-old male nude mice

MIA PaCa-2, PANC-1, and BxPC-3 and the cervical adenocarcinoma cell line HeLa

KRAS

KRAS/Akt signaling pathway

miR-96 in pancreatic cancer cells suppressed cell proliferation, migration, and invasion

[27]

miR-96

Pancreatic carcinogenesis

Down

156 PDACs, 64 IPMNs and 5 MCNs

HPDE, BxPC-3, PANC-1, PK-1, PK-8, PK-9, PK-45H, PK-45P, KLM-1 and BxPC-3

EVI1

KRAS/p27Kip1 pathway

miR-96 potently suppresses KRAS and serves as tumor suppressor in pancreatic cancer

[28]

miR-96

Colon cancer

–

APCCKO/Krasmut mice

HCT116 and SW480

KRAS,

–

Resveratrol has been shown to prevent colorectal carcinogenesis in an animal model of Kras activated cancer possibly through up-regulation of miR-96

[29]

miR‑193b

Esophageal squamous cell carcinoma

Down

53 pairs of esophageal squamous cell carcinoma tissues and ANTs

KYSE450 and TE1, and normal epithelial cell line, Het-1A

KRAS

–

miR-193b inhibited the cell growth, cell proliferation, migration and invasion, and increased the level of apoptotic cells

[36]

miR-873

Pancreatic ductal adenocarcinoma

Down

45 pancreatic tumor tissues and 45 normal tissues

MCF10A; MDA-MB-436, MDA-MB-231, MDA-MB-453, BT-20, HCC1937, SKBR3, T47D, and HEK293; PANC1, BxPC-3, MiaPaCa-2, and Capan-2

KRAS

KRAS/Akt signaling pathway

miR-873 inhibited proliferation, migration, invasion, and colony formation of PDAC cells, and induces cell apoptosis

[37]

miR-31

Colorectal cancer

Up

30 tumor specimens

Caco2, Caco2-BRAFV600E and Caco2-KRASG12V

KRAS and BRAF

–

miR-31 induced cell proliferation and migration

[38]

miR-373

Colorectal cancer

Down

30 tumor specimens

Caco2, Caco2-BRAFV600E and Caco2-KRASG12V

KRAS and BRAF

–

miR-373 inhibited cell proliferation and migration

[38]

miR-30c and miR-21

Non-small-cell lung cancer

Up

44 normal lung samples, 150 lung adenocarcinoma KRAS WT samples and 5 lung adenocarcinoma KRAS G12D samples and KRASLSL-G12D mice

H1299, A549, Calu-6, H1703, H292

NF1 and RASA1/ELK1

KRAS and NF-κB signaling

miR-30c and miR-21 promoted drug resistance and induced cell migration/invasion

[39]

miR-30c

Colorectal cancer

Down

CRC samples from 14 patients

HCT116, DLD1, SW48, HT29 and RKO CRC cells, and HEK-293

KRAS, ME1/P65

miR-30a inhibited tumor growth, migration and invasion

[40]

miR-27b-3p, miR-191-5p, miR-let7d-5p, miR-15b-5p, miR-98-5p, miR-10a-5p, and miR-149-5p

Colorectal cancer

Down

26 tumoral and 30 ANTs

–

KRAS

–

The presence of a different set of miRNAs in KRAS mutated CRC tissues could suggest their putative role as responsive molecular targets

[30]

let-7

Non-small cell lung cancer

–

74 NSCLC cases

–

KRAS

–

let-7 miRNAs is in lung cancer susceptibility

[41]

miR‐127‐3p and miR‐92a

Colorectal carcinoma

Up

Primary tumor of 60 patients with metastatic CRC

–

RSG3 and TOB1

–

Deregulated miRNAs played roles in nicotinamide adenine dinucleotide phosphate (NADPH) regeneration and G protein‐coupled receptor signaling pathways

[42]

miR-18a*

Squamous carcinoma, colon carcinoma

Down

–

Squamous carcinoma A431 cells, colon carcinoma HT-29 cells and fetal hepatic WRL-68 cells

KRAS

–

miR-18a* decreased proliferation and inhibited anchorage-independent growth of cells

[43]

miR-31-3p

Colorectal cancer

Down

Primary tumors from 149 KRAS WT patients

–

KRAS

–

miR-31-3p is a prognostic marker in patients treated with chemotherapy plus cetuximab

[44]

miR-31

Pancreatic and colorectal cancer

Up

–

HPNE cells, HCT116

KRAS, RASA1

MAPK pathway

miR-31 induced invasion and migration in PDAC lines through activation of Rho

[45]

miR-143

Colorectal cancer

Down

13 pairs of matched CRC and ANTs

Lovo cells

KRAS

ERK pathway

miR-143 functions as a tumor suppressor

[31]

miR-143-3p

Pancreatic ductal adenocarcinoma

Down

37 pairs of PDAC tissues and ANTs

MIA PaCa-2, PANC-1 and HPDE

KRAS

ERK pathway

miR-143-3p inhibited cell proliferative, migratory and invasive capacities in PDAC cells

[32]

miR-143

Colorectal cancer

Down

77 pairs of matched CRC and ANT samples

–

KRAS

–

miR-143 expression levels serve as an independent prognostic biomarker for CRC in KRAS wild-type patients

[33]

miR‐143

Colon cancer

Down

BALB/cSlc‐nu/nu (nude) mice

DLD‐1, SW48, HT29 and SW480

K‐Ras, Sos1

K‐Ras/ EGFR

miR‐143 is a tumor suppressive that inhibited proliferation and growth

[34]

miR-143

Prostate cancer

Down

Nine prostate cancer tissues

DU145 and PC3

KRAS and Cyclin D1

EGFR/RAS/MAPK pathway

miR-143 inhibits cell proliferation, migration, and improvement of chemosensitivity to docetaxel

[35]

miR-155

Colon carcinomas

Down

Cbx7+/+, Cbx7+/−, Cbx7−/− and transgenic (TG) Cbx7 mice

–

CBX7/KRAS

–

miR‐155 is a tumor suppressive gene

[46]

miR-193a-3p

Lung cancer

Down

8 pairs of NSCLC tissues and ANTs

A549 and H1975

KRAS

KRAS pathway

miR-193a-3p functions as a tumor suppressor that inhibited proliferation, viability and migration

[47]

miR-200c and miR-221/222

Colorectal cancer

Up

–

HCT116 cells and HKe3 cells

KRAS/PTEN

–

Oncogenic KRAS regulates 3D-specific molecules via miR-200c and miR-221/222

[48]

miR-29b

Colon cancer

Down

40 pairs of tumor tissues and ANTs

HT29, DLD1 and SW480

KRAS

NF-κB signaling

miR-29b-1-5p significantly suppressed cell proliferation

[49]

miR-126

Colorectal cancer

Down

–

HCT116 KRAS-WT and HCT116 KRAS-Mutant

KRAS

–

miR-126 as a selective inhibitor of the viability of KRAS-mutant cells

[50]

miR-126-3p and miR-126-5p

Colorectal cancer

Down

63 pairs of tumor tissues and ANTs

–

KRAS

–

Role of miR-126-3p and miR-126-5p related to regulation of angiogenesis, in patients with CRC treated with bevacizumab

[51]

miR-126

Colorectal cancer

Down

Colorectal tissues from 245 patients (42 noncancer:40 adenoma; 163 primary adenocarcinomas

SW480 and SW48

KRAS

–

miR-126 reduced cell proliferation, increased apoptosis and decreased accumulation of cells in the G0–G1 phase of the colon cancer cells

[52]

miR-193a-3p

Colorectal cancer

Down

70 pairs of tumor tissues and ANTs

SW480 and SW48

KRAS

EMT process

miR-193a-3p reduces the proliferation, migration and colony formation

[53]

miR-181a, miR-200c and miR-210

Colorectal cancer

Up

–

DLD-1 and DKO-4

KRAS

–

These miRNAs are possibly associated with CRC development

[54]

miR-134

Majority of glioblastoma

Down

–

U87, U373, A172, T98G, SNB-19, and SF-767

KRAS and STAT5B

KRAS/STAT5B

miR-134 inhibits cancer cell and stem-cell proliferation and survival

[55]

miR-134

Renal cell carcinoma

Down

24 pairs of tumor specimens and ANTs

786-O, caki-1, 769-P, HEK-293T and ACHN

KRAS

KRAS-related MAPK/ERK

miR-134 could also inhibit migration and invasion by blocking EMT

[56]

miR-134

Glioma

Down

63 glioma tissue samples

U251

KRAS

ERK pathway

miR-134 inhibited glioma cell proliferation and invasion

[57]

miR‑98

Retinoblastoma

Down

RB samples from 60 patients

WERI-Rb-1, Y79 and SO-RB50

IGF1R

IGF1R/k‑Ras/Raf/MEK/ERK signaling pathway

miR‑98 suppress cell growth, migration and invasion

[58]

Let-7a

Colorectal carcinomas

Down

Tissue from 172 patients

–

KRAS

–

Higher let-7a levels were significantly associated with better survival outcomes

[59]

miR-193b

Pancreatic ductal adenocarcinoma

Down

Pancreatic tissue samples from 10 patients

MIA PaCa-2, PANC-1, AsPC-1 and BxPC-3, and hTERT-HPNE

KRAS

AKT and ERK pathways

miR-193b inhibits pancreatic cancer cell growth and proliferation

[60]

miR-206

Pancreatic ductal adenocarcinoma

Down

SCID mice

PANC-1, PANC10.05, BxPC-3, MiaPaca-2, CFPAC-1, Colo357 and Capan-1

KRAS and ANXA2

NF-κB signaling

miR-206 inhibits cell cycle progression, cell proliferation, migration and invasion

[61]

miR-21

Non-small-cell lung cancer

Up

Transgenic mice

–

KRAS

Ras/MEK/ERK pathway

MiR-21 drives tumorigenesis through inhibition of negative regulators of the Ras/MEK/ERK pathway and inhibition of apoptosis

[62]

miR-3923

Pancreatic cancer

Down

Pancreatic tissue samples from 30 pancreatic cancers, 10 chronic pancreatitis and 30 noncancerous pancreatic tissues

PANC-1 and BXPC-3

miR-3923/KRAS pathway

miR-3923 inhibits viability, proliferation and invasion

[23]

miR-489

Pancreatic ductal adenocarcinoma

Down

NOD/SCID mice

BxPC-3 and PANC-1

ADAM9 and MMP7

KRAS-NF-κB-YY1

miR-489 inhibits the migration and metastasis

[63]

miR-155

Pancreatic cancer

Up

–

Capan2, Aspc1, Panc1 and BxPC3

KRAS and Foxo3a

MAPK and NF-κB pathway

miR-155 promotes pancreatic cancer cell proliferation

[64]

miR‑337

Colorectal cancer

Down

54 paired CRC tissues and ANTs

LoVo, HCT116, HT29, SW480, SW620

KRAS

AKT and ERK signalling pathways

miR-337 inhibits cell proliferation, invasion and increases apoptosis

[65]

miR-217

Osteosarcoma

Up

–

Human osteosarcoma 143B cell line

KRAS

miR-217-KRAS axis

miR-217 downregulation led to the loss of enhanced cisplatin sensitivity

[66]

miR-193a-3p

Lung cancer

Down

10 female athymic nude mice

MDA-MB-231, HeyA8 and SKOV3.ip1 cells

KRAS

–

miR-193a-3p functions as a tumor suppressor by inhibiting K-Ras

[67]

miR-768-3p

Brain tumor and lung cancer

Down

19 brain tissue from patients

H520, A549, H661, H441, astrocyte cell line

KRAS

–

miRNA-768-3p inhibits K-ras and suppresses metastasis

[68]