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SMAD2

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GENERAL INFORMATION

Gene name

 SMAD2

Gene description

 SMAD family member 2

Protein class

 Cancer-related genes
Plasma proteins
Predicted intracellular proteins
Predicted membrane proteins
Transcription factors

Predicted localization

 Intracellular,Membrane

Number of transcripts

 10
SHOW MORE

HUMAN PROTEIN ATLAS INFORMATION

Summary

 Localized to the nucleus, nucleoli & cytosol.

RNA cell category

 Expressed in all

Protein evidence

 Evidence at protein level

Main location

 Localized to the Nucleus (supported), Nucleoli (supported), Cytosol (supported)

DATA RELIABILITY

Reliability score

 Supported based on 1 antibody.
CAB073546
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ASSAYS
RNA expression
Human cells

RNA EXPRESSION OVERVIEW
RNA cell line category: Expressed in all
Organ
Origin
Category
Expression
Alphabetical
Cell lines sorted after organ of phenotypic resemblance.

HUMAN CELLS

Summary

 Localized to the nucleus, nucleoli & cytosol.

Main location

 Nucleus (supported), Nucleoli (supported), Cytosol (supported)

 

 

 
Toggle channels
Low
High
G1
S
G2
M
N/A

Thumbnail

Antibody

Cell line

Location

Single-cell variation

Cell cycle dependent variation
CAB073546 MCF7 Nucleus
Nucleoli
Cytosol
CAB073546 U-2 OS Nucleus
Nucleoli
Cytosol

GENE INFORMATION

Gene name

 SMAD2 (HGNC Symbol)

Synonyms

 JV18-1, MADH2, MADR2

Description

 SMAD family member 2 (HGNC Symbol)

Entrez gene summary

 The protein encoded by this gene belongs to the SMAD, a family of proteins similar to the gene products of the Drosophila gene 'mothers against decapentaplegic' (Mad) and the C. elegans gene Sma. SMAD proteins are signal transducers and transcriptional modulators that mediate multiple signaling pathways. This protein mediates the signal of the transforming growth factor (TGF)-beta, and thus regulates multiple cellular processes, such as cell proliferation, apoptosis, and differentiation. This protein is recruited to the TGF-beta receptors through its interaction with the SMAD anchor for receptor activation (SARA) protein. In response to TGF-beta signal, this protein is phosphorylated by the TGF-beta receptors. The phosphorylation induces the dissociation of this protein with SARA and the association with the family member SMAD4. The association with SMAD4 is important for the translocation of this protein into the nucleus, where it binds to target promoters and forms a transcription repressor complex with other cofactors. This protein can also be phosphorylated by activin type 1 receptor kinase, and mediates the signal from the activin. Alternatively spliced transcript variants have been observed for this gene. [provided by RefSeq, May 2012]

Chromosome

 18

Cytoband

 q21.1

Chromosome location (bp)

 47808957 - 47931146

Protein evidence

 Evidence at protein level (all genes)

Ensembl

 ENSG00000175387 (version 83.38)

Entrez gene

 4087

UniProt

 Q15796 (UniProt - Evidence at protein level)

neXtProt

 NX_Q15796

Antibodypedia

 SMAD2 antibodies

PROTEIN BROWSER
SMAD2-001
SMAD2-002
SMAD2-003
SMAD2-004
SMAD2-005
SMAD2-006
SMAD2-007
SMAD2-008
SMAD2-009
SMAD2-201

PROTEIN INFORMATION

Splice variant

UniProt

Protein class

Gene ontology

Length & mass

Signal peptide
(predicted)

Transmembrane regions
(predicted)
SMAD2-001
ENSP00000262160
ENST00000262160
Q15796 [Direct mapping]
Mothers against decapentaplegic homolog 2
Q53XR6 [Target identity:100%; Query identity:100%]
Mothers against decapentaplegic homolog
Show all
Predicted intracellular proteins
Plasma proteins
Transcription factors
   beta-Hairpin exposed by an alpha/beta-scaffold
Cancer-related genes
   Candidate cancer biomarkers
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Kim et al 2014)
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0000122 [negative regulation of transcription from RNA polymerase II promoter]
GO:0000790 [nuclear chromatin]
GO:0000978 [RNA polymerase II core promoter proximal region sequence-specific DNA binding]
GO:0001077 [transcriptional activator activity, RNA polymerase II core promoter proximal region sequence-specific binding]
GO:0001657 [ureteric bud development]
GO:0001701 [in utero embryonic development]
GO:0001706 [endoderm formation]
GO:0001707 [mesoderm formation]
GO:0003677 [DNA binding]
GO:0003682 [chromatin binding]
GO:0003690 [double-stranded DNA binding]
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005160 [transforming growth factor beta receptor binding]
GO:0005515 [protein binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005654 [nucleoplasm]
GO:0005667 [transcription factor complex]
GO:0005737 [cytoplasm]
GO:0005829 [cytosol]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0006367 [transcription initiation from RNA polymerase II promoter]
GO:0006468 [protein phosphorylation]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
GO:0007182 [common-partner SMAD protein phosphorylation]
GO:0007183 [SMAD protein complex assembly]
GO:0007352 [zygotic specification of dorsal/ventral axis]
GO:0007369 [gastrulation]
GO:0007389 [pattern specification process]
GO:0007492 [endoderm development]
GO:0007507 [heart development]
GO:0008134 [transcription factor binding]
GO:0008285 [negative regulation of cell proliferation]
GO:0009749 [response to glucose]
GO:0009791 [post-embryonic development]
GO:0009880 [embryonic pattern specification]
GO:0009952 [anterior/posterior pattern specification]
GO:0010467 [gene expression]
GO:0010628 [positive regulation of gene expression]
GO:0010629 [negative regulation of gene expression]
GO:0010718 [positive regulation of epithelial to mesenchymal transition]
GO:0017015 [regulation of transforming growth factor beta receptor signaling pathway]
GO:0019902 [phosphatase binding]
GO:0023019 [signal transduction involved in regulation of gene expression]
GO:0030073 [insulin secretion]
GO:0030324 [lung development]
GO:0030512 [negative regulation of transforming growth factor beta receptor signaling pathway]
GO:0030513 [positive regulation of BMP signaling pathway]
GO:0030618 [transforming growth factor beta receptor, pathway-specific cytoplasmic mediator activity]
GO:0031016 [pancreas development]
GO:0031053 [primary miRNA processing]
GO:0031625 [ubiquitin protein ligase binding]
GO:0032444 [activin responsive factor complex]
GO:0032924 [activin receptor signaling pathway]
GO:0033613 [activating transcription factor binding]
GO:0034713 [type I transforming growth factor beta receptor binding]
GO:0035019 [somatic stem cell population maintenance]
GO:0035265 [organ growth]
GO:0035326 [enhancer binding]
GO:0035556 [intracellular signal transduction]
GO:0038092 [nodal signaling pathway]
GO:0042803 [protein homodimerization activity]
GO:0043234 [protein complex]
GO:0045165 [cell fate commitment]
GO:0045892 [negative regulation of transcription, DNA-templated]
GO:0045893 [positive regulation of transcription, DNA-templated]
GO:0045944 [positive regulation of transcription from RNA polymerase II promoter]
GO:0046332 [SMAD binding]
GO:0046872 [metal ion binding]
GO:0046982 [protein heterodimerization activity]
GO:0048340 [paraxial mesoderm morphogenesis]
GO:0048589 [developmental growth]
GO:0048617 [embryonic foregut morphogenesis]
GO:0048701 [embryonic cranial skeleton morphogenesis]
GO:0051098 [regulation of binding]
GO:0060021 [palate development]
GO:0060039 [pericardium development]
GO:0060395 [SMAD protein signal transduction]
GO:0070410 [co-SMAD binding]
GO:0070411 [I-SMAD binding]
GO:0070412 [R-SMAD binding]
GO:0070723 [response to cholesterol]
GO:0071141 [SMAD protein complex]
GO:0071144 [SMAD2-SMAD3 protein complex]
GO:1900224 [positive regulation of nodal signaling pathway involved in determination of lateral mesoderm left/right asymmetry]
Show all
 467 aa
52.3 kDa 		No 0
SMAD2-002
ENSP00000384449
ENST00000402690
Q15796 [Direct mapping]
Mothers against decapentaplegic homolog 2
Q53XR6 [Target identity:100%; Query identity:100%]
Mothers against decapentaplegic homolog
Show all
Predicted intracellular proteins
Plasma proteins
Transcription factors
   beta-Hairpin exposed by an alpha/beta-scaffold
Cancer-related genes
   Candidate cancer biomarkers
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Kim et al 2014)
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0000122 [negative regulation of transcription from RNA polymerase II promoter]
GO:0000790 [nuclear chromatin]
GO:0000978 [RNA polymerase II core promoter proximal region sequence-specific DNA binding]
GO:0001077 [transcriptional activator activity, RNA polymerase II core promoter proximal region sequence-specific binding]
GO:0001707 [mesoderm formation]
GO:0003677 [DNA binding]
GO:0003690 [double-stranded DNA binding]
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005160 [transforming growth factor beta receptor binding]
GO:0005515 [protein binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005654 [nucleoplasm]
GO:0005667 [transcription factor complex]
GO:0005737 [cytoplasm]
GO:0005829 [cytosol]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0006367 [transcription initiation from RNA polymerase II promoter]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
GO:0007182 [common-partner SMAD protein phosphorylation]
GO:0007183 [SMAD protein complex assembly]
GO:0007352 [zygotic specification of dorsal/ventral axis]
GO:0007369 [gastrulation]
GO:0008134 [transcription factor binding]
GO:0009952 [anterior/posterior pattern specification]
GO:0010467 [gene expression]
GO:0010718 [positive regulation of epithelial to mesenchymal transition]
GO:0017015 [regulation of transforming growth factor beta receptor signaling pathway]
GO:0019902 [phosphatase binding]
GO:0030512 [negative regulation of transforming growth factor beta receptor signaling pathway]
GO:0030513 [positive regulation of BMP signaling pathway]
GO:0030618 [transforming growth factor beta receptor, pathway-specific cytoplasmic mediator activity]
GO:0031053 [primary miRNA processing]
GO:0031625 [ubiquitin protein ligase binding]
GO:0032444 [activin responsive factor complex]
GO:0032924 [activin receptor signaling pathway]
GO:0033613 [activating transcription factor binding]
GO:0034713 [type I transforming growth factor beta receptor binding]
GO:0035019 [somatic stem cell population maintenance]
GO:0035326 [enhancer binding]
GO:0035556 [intracellular signal transduction]
GO:0038092 [nodal signaling pathway]
GO:0045165 [cell fate commitment]
GO:0045892 [negative regulation of transcription, DNA-templated]
GO:0045893 [positive regulation of transcription, DNA-templated]
GO:0045944 [positive regulation of transcription from RNA polymerase II promoter]
GO:0046332 [SMAD binding]
GO:0046872 [metal ion binding]
GO:0048340 [paraxial mesoderm morphogenesis]
GO:0051098 [regulation of binding]
GO:0060021 [palate development]
GO:0070410 [co-SMAD binding]
GO:0070411 [I-SMAD binding]
GO:0070412 [R-SMAD binding]
GO:0070723 [response to cholesterol]
GO:0071141 [SMAD protein complex]
GO:0071144 [SMAD2-SMAD3 protein complex]
GO:1900224 [positive regulation of nodal signaling pathway involved in determination of lateral mesoderm left/right asymmetry]
Show all
 467 aa
52.3 kDa 		No 0
SMAD2-003
ENSP00000467075
ENST00000591214
B7Z5N5 [Direct mapping]
Mothers against decapentaplegic homolog
Show all
Predicted membrane proteins
   Prediction method-based
   Membrane proteins predicted by MDM
   MEMSAT3 predicted membrane proteins
   MEMSAT-SVM predicted membrane proteins
   Phobius predicted membrane proteins
   SCAMPI predicted membrane proteins
   SPOCTOPUS predicted membrane proteins
   THUMBUP predicted membrane proteins
   TMHMM predicted membrane proteins
   # TM segments-based
   1TM proteins predicted by MDM
Cancer-related genes
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0003677 [DNA binding]
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005515 [protein binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005667 [transcription factor complex]
GO:0005737 [cytoplasm]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
GO:0016021 [integral component of membrane]
Show all
 431 aa
48.7 kDa 		No 1
SMAD2-004
ENSP00000466193
ENST00000586040
Q15796 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
Predicted intracellular proteins
Plasma proteins
Transcription factors
   beta-Hairpin exposed by an alpha/beta-scaffold
Cancer-related genes
   Candidate cancer biomarkers
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0000122 [negative regulation of transcription from RNA polymerase II promoter]
GO:0000790 [nuclear chromatin]
GO:0000978 [RNA polymerase II core promoter proximal region sequence-specific DNA binding]
GO:0001077 [transcriptional activator activity, RNA polymerase II core promoter proximal region sequence-specific binding]
GO:0001707 [mesoderm formation]
GO:0003677 [DNA binding]
GO:0003690 [double-stranded DNA binding]
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005160 [transforming growth factor beta receptor binding]
GO:0005515 [protein binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005654 [nucleoplasm]
GO:0005667 [transcription factor complex]
GO:0005737 [cytoplasm]
GO:0005829 [cytosol]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0006367 [transcription initiation from RNA polymerase II promoter]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
GO:0007182 [common-partner SMAD protein phosphorylation]
GO:0007183 [SMAD protein complex assembly]
GO:0007352 [zygotic specification of dorsal/ventral axis]
GO:0007369 [gastrulation]
GO:0008134 [transcription factor binding]
GO:0009952 [anterior/posterior pattern specification]
GO:0010467 [gene expression]
GO:0010718 [positive regulation of epithelial to mesenchymal transition]
GO:0017015 [regulation of transforming growth factor beta receptor signaling pathway]
GO:0019902 [phosphatase binding]
GO:0030512 [negative regulation of transforming growth factor beta receptor signaling pathway]
GO:0030513 [positive regulation of BMP signaling pathway]
GO:0030618 [transforming growth factor beta receptor, pathway-specific cytoplasmic mediator activity]
GO:0031053 [primary miRNA processing]
GO:0031625 [ubiquitin protein ligase binding]
GO:0032444 [activin responsive factor complex]
GO:0032924 [activin receptor signaling pathway]
GO:0033613 [activating transcription factor binding]
GO:0034713 [type I transforming growth factor beta receptor binding]
GO:0035019 [somatic stem cell population maintenance]
GO:0035326 [enhancer binding]
GO:0035556 [intracellular signal transduction]
GO:0038092 [nodal signaling pathway]
GO:0045165 [cell fate commitment]
GO:0045892 [negative regulation of transcription, DNA-templated]
GO:0045893 [positive regulation of transcription, DNA-templated]
GO:0045944 [positive regulation of transcription from RNA polymerase II promoter]
GO:0046332 [SMAD binding]
GO:0046872 [metal ion binding]
GO:0048340 [paraxial mesoderm morphogenesis]
GO:0051098 [regulation of binding]
GO:0060021 [palate development]
GO:0070410 [co-SMAD binding]
GO:0070411 [I-SMAD binding]
GO:0070412 [R-SMAD binding]
GO:0070723 [response to cholesterol]
GO:0071141 [SMAD protein complex]
GO:0071144 [SMAD2-SMAD3 protein complex]
GO:1900224 [positive regulation of nodal signaling pathway involved in determination of lateral mesoderm left/right asymmetry]
Show all
 437 aa
49 kDa 		No 0
SMAD2-005
ENSP00000465355
ENST00000586514
K7EJX0 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
   SCAMPI predicted membrane proteins
Predicted intracellular proteins
Cancer-related genes
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005667 [transcription factor complex]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
Show all
 217 aa
24.5 kDa 		No 0
SMAD2-006
ENSP00000465971
ENST00000589877
K7EL92 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
Predicted intracellular proteins
Cancer-related genes
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005667 [transcription factor complex]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
Show all
 109 aa
11.9 kDa 		No 0
SMAD2-007
ENSP00000468202
ENST00000587421
K7ERC7 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
   SCAMPI predicted membrane proteins
Predicted intracellular proteins
Cancer-related genes
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005667 [transcription factor complex]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
Show all
 137 aa
15.2 kDa 		No 0
SMAD2-008
ENSP00000466254
ENST00000587269
K7EJX0 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
   SCAMPI predicted membrane proteins
Predicted intracellular proteins
Cancer-related genes
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005667 [transcription factor complex]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
Show all
 217 aa
24.5 kDa 		No 0
SMAD2-009
ENSP00000468732
ENST00000585978
K7ESI8 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
Predicted intracellular proteins
Cancer-related genes
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005667 [transcription factor complex]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
Show all
 108 aa
11.7 kDa 		No 0
SMAD2-201
ENSP00000349282
ENST00000356825
Q15796 [Direct mapping]
Mothers against decapentaplegic homolog 2
Show all
Predicted intracellular proteins
Plasma proteins
Transcription factors
   beta-Hairpin exposed by an alpha/beta-scaffold
Cancer-related genes
   Candidate cancer biomarkers
   Mutated cancer genes
   Mutational cancer driver genes
Protein evidence (Kim et al 2014)
Protein evidence (Ezkurdia et al 2014)
Show all
GO:0000122 [negative regulation of transcription from RNA polymerase II promoter]
GO:0000790 [nuclear chromatin]
GO:0000978 [RNA polymerase II core promoter proximal region sequence-specific DNA binding]
GO:0001077 [transcriptional activator activity, RNA polymerase II core promoter proximal region sequence-specific binding]
GO:0001707 [mesoderm formation]
GO:0003677 [DNA binding]
GO:0003690 [double-stranded DNA binding]
GO:0003700 [transcription factor activity, sequence-specific DNA binding]
GO:0005160 [transforming growth factor beta receptor binding]
GO:0005515 [protein binding]
GO:0005622 [intracellular]
GO:0005634 [nucleus]
GO:0005654 [nucleoplasm]
GO:0005667 [transcription factor complex]
GO:0005737 [cytoplasm]
GO:0005829 [cytosol]
GO:0006351 [transcription, DNA-templated]
GO:0006355 [regulation of transcription, DNA-templated]
GO:0006367 [transcription initiation from RNA polymerase II promoter]
GO:0007179 [transforming growth factor beta receptor signaling pathway]
GO:0007182 [common-partner SMAD protein phosphorylation]
GO:0007183 [SMAD protein complex assembly]
GO:0007352 [zygotic specification of dorsal/ventral axis]
GO:0007369 [gastrulation]
GO:0008134 [transcription factor binding]
GO:0009952 [anterior/posterior pattern specification]
GO:0010467 [gene expression]
GO:0010718 [positive regulation of epithelial to mesenchymal transition]
GO:0017015 [regulation of transforming growth factor beta receptor signaling pathway]
GO:0019902 [phosphatase binding]
GO:0030512 [negative regulation of transforming growth factor beta receptor signaling pathway]
GO:0030513 [positive regulation of BMP signaling pathway]
GO:0030618 [transforming growth factor beta receptor, pathway-specific cytoplasmic mediator activity]
GO:0031053 [primary miRNA processing]
GO:0031625 [ubiquitin protein ligase binding]
GO:0032444 [activin responsive factor complex]
GO:0032924 [activin receptor signaling pathway]
GO:0033613 [activating transcription factor binding]
GO:0034713 [type I transforming growth factor beta receptor binding]
GO:0035019 [somatic stem cell population maintenance]
GO:0035326 [enhancer binding]
GO:0035556 [intracellular signal transduction]
GO:0038092 [nodal signaling pathway]
GO:0045165 [cell fate commitment]
GO:0045892 [negative regulation of transcription, DNA-templated]
GO:0045893 [positive regulation of transcription, DNA-templated]
GO:0045944 [positive regulation of transcription from RNA polymerase II promoter]
GO:0046332 [SMAD binding]
GO:0046872 [metal ion binding]
GO:0048340 [paraxial mesoderm morphogenesis]
GO:0051098 [regulation of binding]
GO:0060021 [palate development]
GO:0070410 [co-SMAD binding]
GO:0070411 [I-SMAD binding]
GO:0070412 [R-SMAD binding]
GO:0070723 [response to cholesterol]
GO:0071141 [SMAD protein complex]
GO:0071144 [SMAD2-SMAD3 protein complex]
GO:1900224 [positive regulation of nodal signaling pathway involved in determination of lateral mesoderm left/right asymmetry]
Show all
 437 aa
49 kDa 		No 0

Contact

The Project

The Human Protein Atlas

The Human Protein Atlas project is funded
by the Knut & Alice Wallenberg foundation.