TISIDB

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Basic function annotation. > Subcellular Location, Domain and Function > Gene Ontology > KEGG and Reactome Pathway

> Subcellular Location, Domain and Function [ TOP ]
Subcellular Location	Serine/threonine-protein kinase PAK 2: Cytoplasm. Note=MYO18A mediates the cellular distribution of the PAK2-ARHGEF7-GIT1 complex to the inner surface of the cell membrane.; SUBCELLULAR LOCATION: PAK-2p34: Nucleus. Cytoplasm, perinuclear region. Membrane; Lipid-anchor. Note=Interaction with ARHGAP10 probably changes PAK-2p34 location to cytoplasmic perinuclear region. Myristoylation changes PAK-2p34 location to the membrane.
Domain	PF00786 P21-Rho-binding domain PF00069 Protein kinase domain
Function	Serine/threonine protein kinase that plays a role in a variety of different signaling pathways including cytoskeleton regulation, cell motility, cell cycle progression, apoptosis or proliferation. Acts as downstream effector of the small GTPases CDC42 and RAC1. Activation by the binding of active CDC42 and RAC1 results in a conformational change and a subsequent autophosphorylation on several serine and/or threonine residues. Full-length PAK2 stimulates cell survival and cell growth. Phosphorylates MAPK4 and MAPK6 and activates the downstream target MAPKAPK5, a regulator of F-actin polymerization and cell migration. Phosphorylates JUN and plays an important role in EGF-induced cell proliferation. Phosphorylates many other substrates including histone H4 to promote assembly of H3.3 and H4 into nucleosomes, BAD, ribosomal protein S6, or MBP. Additionally, associates with ARHGEF7 and GIT1 to perform kinase-independent functions such as spindle orientation control during mitosis. On the other hand, apoptotic stimuli such as DNA damage lead to caspase-mediated cleavage of PAK2, generating PAK-2p34, an active p34 fragment that translocates to the nucleus and promotes cellular apoptosis involving the JNK signaling pathway. Caspase-activated PAK2 phosphorylates MKNK1 and reduces cellular translation.

> Gene Ontology [ TOP ]
Biological Process	GO:0001933 negative regulation of protein phosphorylation GO:0002218 activation of innate immune response GO:0002220 innate immune response activating cell surface receptor signaling pathway GO:0002223 stimulatory C-type lectin receptor signaling pathway GO:0002429 immune response-activating cell surface receptor signaling pathway GO:0002694 regulation of leukocyte activation GO:0002696 positive regulation of leukocyte activation GO:0002697 regulation of immune effector process GO:0002757 immune response-activating signal transduction GO:0002758 innate immune response-activating signal transduction GO:0002764 immune response-regulating signaling pathway GO:0002768 immune response-regulating cell surface receptor signaling pathway GO:0002831 regulation of response to biotic stimulus GO:0006469 negative regulation of protein kinase activity GO:0007159 leukocyte cell-cell adhesion GO:0009615 response to virus GO:0010466 negative regulation of peptidase activity GO:0010951 negative regulation of endopeptidase activity GO:0016358 dendrite development GO:0018105 peptidyl-serine phosphorylation GO:0018108 peptidyl-tyrosine phosphorylation GO:0018209 peptidyl-serine modification GO:0018212 peptidyl-tyrosine modification GO:0022407 regulation of cell-cell adhesion GO:0022409 positive regulation of cell-cell adhesion GO:0031294 lymphocyte costimulation GO:0031295 T cell costimulation GO:0031349 positive regulation of defense response GO:0033673 negative regulation of kinase activity GO:0033674 positive regulation of kinase activity GO:0038093 Fc receptor signaling pathway GO:0038095 Fc-epsilon receptor signaling pathway GO:0042110 T cell activation GO:0042326 negative regulation of phosphorylation GO:0043154 negative regulation of cysteine-type endopeptidase activity involved in apoptotic process GO:0043281 regulation of cysteine-type endopeptidase activity involved in apoptotic process GO:0043900 regulation of multi-organism process GO:0043903 regulation of symbiosis, encompassing mutualism through parasitism GO:0045088 regulation of innate immune response GO:0045089 positive regulation of innate immune response GO:0045785 positive regulation of cell adhesion GO:0045860 positive regulation of protein kinase activity GO:0045861 negative regulation of proteolysis GO:0046777 protein autophosphorylation GO:0048010 vascular endothelial growth factor receptor signaling pathway GO:0050688 regulation of defense response to virus GO:0050690 regulation of defense response to virus by virus GO:0050730 regulation of peptidyl-tyrosine phosphorylation GO:0050731 positive regulation of peptidyl-tyrosine phosphorylation GO:0050792 regulation of viral process GO:0050851 antigen receptor-mediated signaling pathway GO:0050852 T cell receptor signaling pathway GO:0050863 regulation of T cell activation GO:0050865 regulation of cell activation GO:0050867 positive regulation of cell activation GO:0050870 positive regulation of T cell activation GO:0051249 regulation of lymphocyte activation GO:0051251 positive regulation of lymphocyte activation GO:0051346 negative regulation of hydrolase activity GO:0051348 negative regulation of transferase activity GO:0051607 defense response to virus GO:0052547 regulation of peptidase activity GO:0052548 regulation of endopeptidase activity GO:0060996 dendritic spine development GO:0061097 regulation of protein tyrosine kinase activity GO:0061098 positive regulation of protein tyrosine kinase activity GO:0070486 leukocyte aggregation GO:0070489 T cell aggregation GO:0071407 cellular response to organic cyclic compound GO:0071593 lymphocyte aggregation GO:0097191 extrinsic apoptotic signaling pathway GO:0097194 execution phase of apoptosis GO:0098542 defense response to other organism GO:1900117 regulation of execution phase of apoptosis GO:1900118 negative regulation of execution phase of apoptosis GO:1903037 regulation of leukocyte cell-cell adhesion GO:1903039 positive regulation of leukocyte cell-cell adhesion GO:2000116 regulation of cysteine-type endopeptidase activity GO:2000117 negative regulation of cysteine-type endopeptidase activity GO:2001233 regulation of apoptotic signaling pathway GO:2001235 positive regulation of apoptotic signaling pathway GO:2001236 regulation of extrinsic apoptotic signaling pathway GO:2001238 positive regulation of extrinsic apoptotic signaling pathway GO:2001270 regulation of cysteine-type endopeptidase activity involved in execution phase of apoptosis GO:2001271 negative regulation of cysteine-type endopeptidase activity involved in execution phase of apoptosis
Molecular Function	GO:0004674 protein serine/threonine kinase activity GO:0004702 receptor signaling protein serine/threonine kinase activity GO:0005057 receptor signaling protein activity GO:0008047 enzyme activator activity GO:0019207 kinase regulator activity GO:0019209 kinase activator activity GO:0019887 protein kinase regulator activity GO:0030295 protein kinase activator activity GO:0030296 protein tyrosine kinase activator activity GO:0045296 cadherin binding GO:0050839 cell adhesion molecule binding GO:0098631 protein binding involved in cell adhesion GO:0098632 protein binding involved in cell-cell adhesion GO:0098641 cadherin binding involved in cell-cell adhesion
Cellular Component	GO:0005913 cell-cell adherens junction

> KEGG and Reactome Pathway [ TOP ]
KEGG	hsa04010 MAPK signaling pathway hsa04012 ErbB signaling pathway hsa04014 Ras signaling pathway hsa04360 Axon guidance hsa04510 Focal adhesion hsa04660 T cell receptor signaling pathway hsa04810 Regulation of actin cytoskeleton
Reactome	R-HSA-428540: Activation of Rac R-HSA-1280218: Adaptive Immune System R-HSA-109581: Apoptosis R-HSA-75153: Apoptotic execution phase R-HSA-422475: Axon guidance R-HSA-5621481: C-type lectin receptors (CLRs) R-HSA-5621575: CD209 (DC-SIGN) signaling R-HSA-389356: CD28 co-stimulation R-HSA-389359: CD28 dependent Vav1 pathway R-HSA-388841: Costimulation by the CD28 family R-HSA-1266738: Developmental Biology R-HSA-1643685: Disease R-HSA-2682334: EPH-Ephrin signaling R-HSA-3928664: Ephrin signaling R-HSA-2871796: FCERI mediated MAPK activation R-HSA-2454202: Fc epsilon receptor (FCERI) signaling R-HSA-202433: Generation of second messenger molecules R-HSA-162906: HIV Infection R-HSA-162909: Host Interactions of HIV factors R-HSA-168256: Immune System R-HSA-5663205: Infectious disease R-HSA-168249: Innate Immune System R-HSA-5683057: MAPK family signaling cascades R-HSA-5687128: MAPK6/MAPK4 signaling R-HSA-164944: Nef and signal transduction R-HSA-5357801: Programmed Cell Death R-HSA-195258: RHO GTPase Effectors R-HSA-5627123: RHO GTPases activate PAKs R-HSA-169911: Regulation of Apoptosis R-HSA-211728: Regulation of PAK-2p34 activity by PS-GAP/RHG10 R-HSA-211733: Regulation of activated PAK-2p34 by proteasome mediated degradation R-HSA-399954: Sema3A PAK dependent Axon repulsion R-HSA-373755: Semaphorin interactions R-HSA-162582: Signal Transduction R-HSA-194315: Signaling by Rho GTPases R-HSA-376176: Signaling by Robo receptor R-HSA-194138: Signaling by VEGF R-HSA-211736: Stimulation of the cell death response by PAK-2p34 R-HSA-202403: TCR signaling R-HSA-164952: The role of Nef in HIV-1 replication and disease pathogenesis R-HSA-4420097: VEGFA-VEGFR2 Pathway R-HSA-5218920: VEGFR2 mediated vascular permeability

> KEGG and Reactome Pathway

[ TOP ]

KEGG

hsa04010 MAPK signaling pathway
hsa04012 ErbB signaling pathway
hsa04014 Ras signaling pathway
hsa04360 Axon guidance
hsa04510 Focal adhesion
hsa04660 T cell receptor signaling pathway
hsa04810 Regulation of actin cytoskeleton

Reactome

R-HSA-428540: Activation of Rac
R-HSA-1280218: Adaptive Immune System
R-HSA-109581: Apoptosis
R-HSA-75153: Apoptotic execution phase
R-HSA-422475: Axon guidance
R-HSA-5621481: C-type lectin receptors (CLRs)
R-HSA-5621575: CD209 (DC-SIGN) signaling
R-HSA-389356: CD28 co-stimulation
R-HSA-389359: CD28 dependent Vav1 pathway
R-HSA-388841: Costimulation by the CD28 family
R-HSA-1266738: Developmental Biology
R-HSA-1643685: Disease
R-HSA-2682334: EPH-Ephrin signaling
R-HSA-3928664: Ephrin signaling
R-HSA-2871796: FCERI mediated MAPK activation
R-HSA-2454202: Fc epsilon receptor (FCERI) signaling
R-HSA-202433: Generation of second messenger molecules
R-HSA-162906: HIV Infection
R-HSA-162909: Host Interactions of HIV factors
R-HSA-168256: Immune System
R-HSA-5663205: Infectious disease
R-HSA-168249: Innate Immune System
R-HSA-5683057: MAPK family signaling cascades
R-HSA-5687128: MAPK6/MAPK4 signaling
R-HSA-164944: Nef and signal transduction
R-HSA-5357801: Programmed Cell Death
R-HSA-195258: RHO GTPase Effectors
R-HSA-5627123: RHO GTPases activate PAKs
R-HSA-169911: Regulation of Apoptosis
R-HSA-211728: Regulation of PAK-2p34 activity by PS-GAP/RHG10
R-HSA-211733: Regulation of activated PAK-2p34 by proteasome mediated degradation
R-HSA-399954: Sema3A PAK dependent Axon repulsion
R-HSA-373755: Semaphorin interactions
R-HSA-162582: Signal Transduction
R-HSA-194315: Signaling by Rho GTPases
R-HSA-376176: Signaling by Robo receptor
R-HSA-194138: Signaling by VEGF
R-HSA-211736: Stimulation of the cell death response by PAK-2p34
R-HSA-202403: TCR signaling
R-HSA-164952: The role of Nef in HIV-1 replication and disease pathogenesis
R-HSA-4420097: VEGFA-VEGFR2 Pathway
R-HSA-5218920: VEGFR2 mediated vascular permeability

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Literatures that report relations between PAK2 and anti-tumor immunity. The specific mechanism were also collected if the literature reports that a gene specifically promotes or inhibits the infiltration or function of T/NK cells.

> Text Mining [ TOP ]
There is no record.

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	High-throughput screening data (e.g. CRISPR-Cas9, shRNA and RNAi) for T cell-mediated killing. Genetic screen techniques can identify mechanisms of tumor cell resistance (e.g., PTPN2) and sensitivity (e.g., APLNR) to killing by cytotoxic T cells, the central effectors of anti-tumor immunity. After comprehensively searching, eight groups of screening data sets were collected in the current database. In this tab, users can check whether their selected genes cause resistance or increase sensitivity to T cell-mediated killing in various data sets.

> High-throughput Screening

[ TOP ]

Statistical results of PAK2 in screening data sets for detecting immune reponses.

PMID	Screening System	Cancer Type	Cell Line	Data Set	Statistical Results	Relation to immunity
29301958	CRISPR-Cas9	melanoma	B16F10	Pmel-1 T cell	logFC: -2.30; FDR: 0.04630	Resistant to T cell-mediated killing
29301958	CRISPR-Cas9	melanoma	B16F10	OT-1 T cell	logFC: -1.86; FDR: 0.04220	Resistant to T cell-mediated killing
28783722	CRISPR-Cas9	melanoma	Mel624	2CT-CRISPR	NA/NS	NA/NS
28723893	CRISPR-Cas9	melanoma	B16	GVAX+Anti-PD1	NA/NS	NA/NS
28723893	CRISPR-Cas9	melanoma	B16	GVAX	NA/NS	NA/NS
25691366	RNAi	Breast cancer	MCF7	Luc-CTL assay	NA/NS	NA/NS
24476824	shRNA	melanoma	B16	Primary screen	NA/NS	NA/NS
24476824	shRNA	melanoma	B16	Secondary screen	NA/NS	NA/NS

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Transcriptomic and genomic profiling of pre-treated tumor biopsies from responders and non-responders to immunotherapy. These data were used to identify signatures and mechanisms of response to checkpoint blockade (e.g., anti-PDL1 and anti-PD1). One example is that mutations in the gene PBRM1 benefit clinical survival of patients with clear cell renal cell carcinoma. After comprehensively searching, we collected 5 and 6 of transcriptomic and genomic data sets, respectively. In this tab, users can check whether their selected genes have significant difference of expression or mutation between responders and non-responders in various data sets. > Expression difference between responders and non-responders > Mutation difference between responders and non-responders

> Expression difference between responders and non-responders

[ TOP ]

Points in the above scatter plot represent the expression difference of PAK2 in various data sets.

No	PMID	Cancer type	Group	Drug	# Res	# NRes	Log2 (Fold Change)	P value
1	26997480	Melanoma	all	Anti-PD-1 (pembrolizumab and nivolumab)	14	12	-0.07	0.734
2	26997480	Melanoma	MAPKi	Anti-PD-1 (pembrolizumab and nivolumab)	6	5	0.155	0.941
3	26997480	Melanoma	non-MAPKi	Anti-PD-1 (pembrolizumab and nivolumab)	8	7	-0.234	0.879
4	28552987	Urothelial cancer	all	Anti-PD-L1 (atezolizumab)	9	16	0.163	0.609
5	28552987	Urothelial cancer	smoking	Anti-PD-L1 (atezolizumab)	5	9	-0.008	0.997
6	28552987	Urothelial cancer	non-smoking	Anti-PD-L1 (atezolizumab)	4	7	0.381	0.882
7	29033130	Melanoma	all	Anti-PD-1 (nivolumab)	26	23	0.241	0.601
8	29033130	Melanoma	NIV3-PROG	Anti-PD-1 (nivolumab)	15	11	0.309	0.879
9	29033130	Melanoma	NIV3-NAIVE	Anti-PD-1 (nivolumab)	11	12	0.189	0.932
10	29301960	Clear cell renal cell carcinoma (ccRCC)	all	Anti-PD-1 (nivolumab)	4	8	0.383	0.789
11	29301960	Clear cell renal cell carcinoma (ccRCC)	VEGFRi	Anti-PD-1 (nivolumab)	2	0	0	1
12	29301960	Clear cell renal cell carcinoma (ccRCC)	non-VEGFRi	Anti-PD-1 (nivolumab)	2	8	0.807	0.709
13	29443960	Urothelial cancer	all	Anti-PD-L1 (atezolizumab)	68	230	0.068	0.22

> Mutation difference between responders and non-responders

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Points in the above scatter plot represent the mutation difference of PAK2 in various data sets.

No	PMID	Cancer type	Group	Drug	# Res	# NRes	% Mut/Res	% Mut/NRes	% Diff (R vs NR)	Pval
1	25765070	Non-small cell lung cancer (NSCLC)	all	Anti-PD-1 (pembrolizumab)	14	17	0	0	0	1
2	25765070	Non-small cell lung cancer (NSCLC)	smoking	Anti-PD-1 (pembrolizumab)	10	3	0	0	0	1
3	25765070	Non-small cell lung cancer (NSCLC)	non-smoking	Anti-PD-1 (pembrolizumab)	4	14	0	0	0	1
4	26359337	Melanoma	all	Anti-CTLA-4 (ipilimumab)	27	73	0	1.4	-1.4	1
5	26359337	Melanoma	BRAFi	Anti-CTLA-4 (ipilimumab)	0	14	0	0	0	1
6	26359337	Melanoma	non-BRAFi	Anti-CTLA-4 (ipilimumab)	27	59	0	1.7	-1.7	1
7	26997480	Melanoma	all	Anti-PD-1 (pembrolizumab and nivolumab)	21	17	0	5.9	-5.9	0.447
8	26997480	Melanoma	MAPKi	Anti-PD-1 (pembrolizumab and nivolumab)	8	6	0	0	0	1
9	26997480	Melanoma	non-MAPKi	Anti-PD-1 (pembrolizumab and nivolumab)	13	11	0	9.1	-9.1	0.458
10	28552987	Urothelial cancer	all	Anti-PD-L1 (atezolizumab)	9	16	22.2	0	22.2	0.12
11	28552987	Urothelial cancer	smoking	Anti-PD-L1 (atezolizumab)	5	9	20	0	20	0.357
12	28552987	Urothelial cancer	non-smoking	Anti-PD-L1 (atezolizumab)	4	7	25	0	25	0.364
13	29033130	Melanoma	all	Anti-PD-1 (nivolumab)	38	27	13.2	11.1	2.1	1
14	29033130	Melanoma	NIV3-PROG	Anti-PD-1 (nivolumab)	22	13	13.6	15.4	-1.8	1
15	29033130	Melanoma	NIV3-NAIVE	Anti-PD-1 (nivolumab)	16	14	12.5	7.1	5.4	1
16	29301960	Clear cell renal cell carcinoma (ccRCC)	all	Anti-PD-1 (nivolumab)	11	13	0	0	0	1
17	29301960	Clear cell renal cell carcinoma (ccRCC)	VEGFRi	Anti-PD-1 (nivolumab)	6	1	0	0	0	1
18	29301960	Clear cell renal cell carcinoma (ccRCC)	non-VEGFRi	Anti-PD-1 (nivolumab)	5	12	0	0	0	1

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Relations between abundance of tumor-infiltrating lymphocytes (TILs) and expression, copy number, methylation, or mutation of PAK2. The immune-related signatures of 28 TIL types from Charoentong's study, which can be viewed in the download page. For each cancer type, the relative abundance of TILs were inferred by using gene set variation analysis (GSVA) based on gene expression profile. In this tab, users can examine which kinds of TILs might be regulated by the current gene.

> Lymphocyte [ TOP ]

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Relations between three kinds of immunomodulators and expression, copy number, methylation, or mutation of PAK2. These immunomo-dulators were collected from Charoentong's study. In this tab, users can examine which immunomodulators might be regulated by PAK2. > Immunoinhibitor > Immunostimulator > MHC molecule

> Immunoinhibitor [ TOP ]

> Immunostimulator [ TOP ]

> MHC molecule [ TOP ]

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Relations between chemokines (or receptors) and expression, copy number, methylation, or mutation of PAK2. In this tab, users can examine which chemokines (or receptors) might be regulated by the current gene. > Chemokine > Receptor

> Chemokine [ TOP ]

> Receptor [ TOP ]

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Distribution of PAK2 expression across immune and molecular subtypes. > Immune subtype > Molecular subtype

> Immune subtype [ TOP ]

> Molecular subtype [ TOP ]

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Associations between PAK2 and clinical features. > Overall survival analysis > Cancer stage > Tumor grade

> Overall survival [ TOP ]

> Stage [ TOP ]

> Grade [ TOP ]

Summary
Symbol	PAK2
Name	p21 protein (Cdc42/Rac)-activated kinase 2
Aliases	PAK65; PAKgamma; S6/H4 kinase; p21 (CDKN1A)-activated kinase 2; gamma-PAK; p21-activated kinase 2; Serine/th ......
Chromosomal Location	3q29
External Links	HGNC, NCBI, Ensembl, Uniprot, GeneCards
Content	Drugs targeting PAK2 collected from DrugBank database.

> Drugs from DrugBank database [ TOP ]
There is no record.

Browse PAK2