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General Information
Symbol
Dmel\Hsp22
Species
D. melanogaster
Name
Heat shock protein 22
Annotation Symbol
CG4460
Feature Type
FlyBase ID
FBgn0001223
Gene Model Status
Stock Availability
Gene Snapshot
Heat shock protein 22 (Hsp22) encodes a molecular chaperone that is encoded in the nucleus and targeted to the mitochondrial matrix. Hsp22 expression is dramatically increased during aging, and both Hsp22 mutation and over-expression are reported to modulate adult life span and stress resistance. [Date last reviewed: 2018-09-13]
Also Known As

DmHsp22

Key Links
Genomic Location
Cytogenetic map
Sequence location
3L:9,372,931..9,374,964 [+]
Recombination map

3-29

RefSeq locus
NT_037436 REGION:9372931..9374964
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (7 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from sequence model
Biological Process (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the small heat shock protein (HSP20) family. (P02515)
Summaries
Gene Group (FlyBase)
SMALL HEAT SHOCK PROTEINS -
The small Heat Shock Protein (sHSP) family is characterized by the presence of an α-crystallin domain. sHSPs play a crucial role in protein folding. Most sHSPs have the ability to prevent protein aggregation and to maintain substrate proteins in a refoldable state. (Adapted from FBrf0228040 and FBrf0231212).
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
Hsp22 - Hsp-G3
There are seven closely related heat-shock genes at 67B (Ayme and Tissieres, 1985; Pauli, Arrigo, Vasquez, Tonka, and Tissieres, 1989, Genome 31: 671-76). In addition to the four small heat-shock genes previously identified (Hsp22, Hsp23, Hsp26, and Hsp27), three more genes (Hsp-G1, Hsp-G2, and Hsp-G3, formerly called Gene1, Gene2, and Gene3) have been found clustered within 15 kb of DNA at the same 67B cytological location. All seven genes are heat-shock inducible in almost all cells at the stages tested (Ayme and Tissieres, 1985). The genes are also transcribed during certain developmental stages in the absence of heat shock (Sirotkin and Davidson, 1982, Dev. Biol. 89: 196-210). Pauli et al (1989) report that the maximum accumulation of developmental rRNA in a majority of these small heat-shock genes occurs in the white pupae stage; in Hsp-G2, however, a small transcipt is found in embryos, first and second instar larvae, and young pupae; and a larger transcript in the pupal and adult stages of males (Pauli and Tonka, 1987, J. Mol. Biol. 198: 235-40; Pauli, Tonka, and Ayme-Southgate, 1988, J. Mol. Biol. 200: 47-53). In absence of stress, the expression of Hsp26 has been observed in spermatocytes, nurse cells, epithelium, imaginal discs, proventriculus, and neurocytes (Glaser, Wolfner, and Lis, 1986, EMBO 5: 747-54). Transcripts of Hsp26 and Hsp27 accumulate in adult ovaries, apparently originating in nurse cells (Zimmerman, Petri, and Meselson, 1983, Cell 32: 1161-70).
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Hsp22 for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model

gene_with_dicistronic_mRNA ; SO:0000722

Gene model reviewed during 5.46

Dicistronic transcript isoform(s) appear to be relatively rare based on RNA-Seq and/or EST data; may be stage- or tissue-specific.

Gene model reviewed during 5.54

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0100558
1900
174
FBtr0100559
968
174
Additional Transcript Data and Comments
Reported size (kB)

1.03 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0100010
19.8
174
4.95
FBpp0100011
19.8
174
4.95
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

174 aa isoforms: Hsp22-PA, Hsp22-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

174 (aa); 19.705 (kD predicted)

22.666 (kD predicted)

22 (kD observed)

Comments

Size determined by in vitro translation.

External Data
Crossreferences
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Hsp22 using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Hsp22 in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 12 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 3 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Hsp22
Transgenic constructs containing regulatory region of Hsp22
Deletions and Duplications ( 3 )
Phenotypes
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (11)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
4 of 15
Yes
No
4 of 15
Yes
No
4 of 15
Yes
No
5  
3 of 15
No
No
2  
3 of 15
No
No
3 of 15
No
No
3 of 15
No
Yes
4  
2 of 15
No
Yes
2 of 15
No
No
2 of 15
No
Yes
1  
2 of 15
No
Yes
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (9)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
5 of 15
Yes
No
5 of 15
Yes
No
4 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
Yes
2 of 15
No
No
2 of 15
No
Yes
2 of 15
No
No
Rattus norvegicus (Norway rat) (9)
5 of 13
Yes
No
5 of 13
Yes
No
4 of 13
No
No
3 of 13
No
No
3 of 13
No
No
3 of 13
No
Yes
2 of 13
No
No
2 of 13
No
Yes
2 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (14)
6 of 12
Yes
Yes
4 of 12
No
No
4 of 12
No
No
4 of 12
No
No
3 of 12
No
No
3 of 12
No
No
3 of 12
No
Yes
2 of 12
No
No
2 of 12
No
No
2 of 12
No
No
1 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (13)
5 of 15
Yes
No
5 of 15
Yes
No
5 of 15
Yes
Yes
4 of 15
No
No
4 of 15
No
No
4 of 15
No
Yes
3 of 15
No
No
3 of 15
No
No
3 of 15
No
Yes
3 of 15
No
No
2 of 15
No
No
2 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (19)
7 of 15
Yes
No
7 of 15
Yes
No
7 of 15
Yes
Yes
6 of 15
No
Yes
6 of 15
No
No
6 of 15
No
Yes
6 of 15
No
Yes
5 of 15
No
No
5 of 15
No
No
5 of 15
No
Yes
4 of 15
No
No
4 of 15
No
No
4 of 15
No
No
4 of 15
No
Yes
4 of 15
No
No
4 of 15
No
Yes
3 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (15)
3 of 9
Yes
No
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
1 of 9
No
No
1 of 9
No
No
1 of 9
No
No
1 of 9
No
No
1 of 9
No
No
1 of 9
No
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
2 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (2)
3 of 12
Yes
Yes
2 of 12
No
Yes
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG09190GOL )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( None identified )
No non-Drosophilid orthologies identified
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( None identified )
No non-Dipteran orthologies identified
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
No non-Insect Arthropod orthologies identified
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0USC )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (10)
5 of 10
5 of 10
5 of 10
5 of 10
5 of 10
4 of 10
4 of 10
4 of 10
3 of 10
3 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 7 )
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v8.0 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Functional Complementation Data
    Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
    Interactions
    Summary of Physical Interactions
    esyN Network Diagram
    Show neighbor-neighbor interactions:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3L
    Recombination map

    3-29

    Cytogenetic map
    Sequence location
    3L:9,372,931..9,374,964 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    67B2-67B2
    Limits computationally determined from genome sequence between P{PZ}l(3)0162901629&P{PZ}mRpL1210534 and P{EP}Hsp26EP3336&P{EP}Hsp26EP3315
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    67B-67B
    (determined by in situ hybridisation)
    A probe containing both Hsp22 and Hsp26 hybridises to cytological locations 67B1--67B2 and 67B4--67B5.
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (7)
    Genomic Clones (18)
     

    Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

    cDNA Clones (16)
     

    Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.

    cDNA clones, fully sequenced
    BDGP DGC clones
    Other clones
    Drosophila Genomics Resource Center cDNA clones

    For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

    cDNA Clones, End Sequenced (ESTs)
    BDGP DGC clones
    RNAi and Array Information
    Linkouts
    DRSC - Results frm RNAi screens
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    Antibody Information
    Laboratory Generated Antibodies
     
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Additional comments

    Dicistronic annotation CG32041 split out into separate annotations for each open reading frame, CG4456 and CG4460, in release 4.2 of the genome annotation. CG4456 corresponds to Hsp67Bb and CG4460 corresponds to Hsp22.

    One or more of the processed transcripts for this gene contain(s) two non-overlapping open reading frames (ORFs). The non-overlapping ORFs are represented by Hsp67Bb and Hsp22.

    Other Comments

    Expression is rapidly induced in the testis after heat stress. Activation of Hsp22 in testes and heads necessitates the presence of a functional Hsf.

    Increase in RNA levels is observed with aging.

    Chromosome staining reveals that Trl and heat shock transcription factors (HSF) colocalises at Hsp22.

    In vivo UV cross-linking and nuclear run-on assays shows that RNA polymerase II density on the Hsp22 gene is rapidly increased by heat shock.

    In unshocked cells Hsp83 is moderately transcribed while transcription from the other heat shock genes is undetectable. Engaged but paused RNA molecules are found at the various Hsp70 and Hsp26 genes but not at the other heat shock genes. Increased transcription of the heat shock genes is observed within 1-2 mins of heat shock and maximal rates were reached within 2-5 minutes. Rates of transcription vary over a 20-fold range.

    Exposure of cells to pulses of elevated temperature initiates the heat-shock response. A restricted subset of genes, the Hsp genes, is activated and the majority of transcription and translation is shut down. 3H-uridine incorporation ceases at its usual positions and commences at new puff sites. Preexisting polysomes disaggregate and within a few minutes a new population of polysomes appears containing newly transcribed mRNA; this RNA hybridizes to some of the heat-shock puffs. Similar response inducible by other stressful treatments. The response may be elicited at all stages of the life cycle and in cultured cells.

    Hsp22 regulatory regions involved in heat shock expression and ecdysterone-induced expression have been identified.

    Translation of Hsp70 mRNAs and to a lesser extent the mRNAs for the small heat shock proteins is almost independent of eIF-4E.

    Mutations at br reduce the transcription rate or stability of the small heat shock protein mRNAs.

    Heat shock induces the accumulation of a dicistronic mRNA which contains both the Hsp22 and Hsp67Bb open reading frames. Overall orientation not stated: Hsp27- Hsp23- Hsp67Ba+ Hsp26+ Hsp22- Hsp67Bb- Hsp67Bc+

    The binding sites for the protein factors required for activation of transcription of Hsp genes are multiple short upstream sequence elements called HSEs or heat shock consensus elements.

    Heat-shock inducible in almost all cells at the stages tested.

    Activation of transcription of Hsp genes apparently involves the sequential binding of two or more protein factors in vicinity of TATA box.

    Polymerase II dissociates from most chromosome regions and accumulates at the new heat shock puff sites upon heat shock.

    Hsp22 is transcribed during certain developmental stages in the absence of heat shock.

    The effects of heat shock may be abrogated to some degree by pretreatment with a pulse of a slightly lower temperature.

    Mitochondrial and histone-gene activities persist transcription and translation.

    In polytene cells, during heat shock response, existing puffs regress and a novel group quickly appear at 33B, 63C, 64F, 67B, 70A, 87A, 87C, 93D, 95D.

    The heat shock response follows a pulse of 36oC to 40oC; treatments above 40oC inhibit all activity and lead to death; treatments of 30oC-35oC induce heat-shock-protein synthesis without repressing normal protein synthesis.

    In polytene cells, during heat shock response, existing puffs regress and a novel group quickly appear at cytological locations 33B, 63C, 64F, 67B, 70A, 87A, 87C, 93D, 95D.

    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 47 )
    Sequence Crossreferences
    NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
    GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
    GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
    RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
    UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
    Other crossreferences
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    Flygut - An atlas of the Drosophila adult midgut
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    KEGG Genes - Molecular building blocks of life in the genomic space.
    modMine - A data warehouse for the modENCODE project
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DPiM - Drosophila Protein interaction map
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
    FlyMine - An integrated database for Drosophila genomics
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (17)
    Reported As
    Symbol Synonym
    Dmel20.8
    Hsp22
    (Santana et al., 2020, Bouska et al., 2019, Engel et al., 2019, Krittika and Yadav, 2019, Maitra et al., 2019, Meltzer et al., 2019, Michalak et al., 2019, Su et al., 2019, Boardman et al., 2018, Dabbaghizadeh et al., 2018, Duncan et al., 2018, Walls et al., 2018, Zhang et al., 2018, Dabbaghizadeh et al., 2017, Kuintzle et al., 2017, Donovan and Marr, 2016, Gene Disruption Project members, 2016-, Kiss et al., 2016, Lopez et al., 2016, Maistrenko et al., 2016, Morrow et al., 2016, Zhang et al., 2016, Castillo et al., 2015, Kim et al., 2015, Kopp et al., 2015, Moskalev et al., 2015, Štětina et al., 2015, Tower, 2015, Van Bortle et al., 2015, Benbahouche et al., 2014, Huang et al., 2014, Marr et al., 2014, Taylor et al., 2014, Tower et al., 2014, Vartiainen et al., 2014, Colinet et al., 2013, Stefanatos et al., 2013, Eleftherianos and Castillo, 2012, Fredriksson et al., 2012, Hirano et al., 2012, Thomas et al., 2012, Wilczynski et al., 2012, Friedman et al., 2011, Morettini et al., 2011, Navarro et al., 2011, Takahashi et al., 2011, Colinet et al., 2010, Colinet et al., 2010, Fernández-Ayala et al., 2010, Groebe et al., 2010, Kim et al., 2010, Kim et al., 2010, Kong et al., 2010, Lorbeck et al., 2010, Takahashi et al., 2010, Tian et al., 2010, Wadhwa et al., 2010, Gruenewald et al., 2009, Moskalev et al., 2009, Bettencourt et al., 2008, Duncan, 2008, Gilchrist et al., 2008, Harvey et al., 2008, Sudi et al., 2008, Xi et al., 2008, Neal et al., 2006, Walser et al., 2006, Yepiskoposyan et al., 2006, Duncan, 2005, Geiger-Thornsberry and Mackay, 2004, Morrow et al., 2004, Gim et al., 2001)
    anon-WO0118547.50
    anon-WO0140519.138
    small hsp locus 67B
    Name Synonyms
    Secondary FlyBase IDs
    • FBgn0044743
    • FBgn0052041
    • FBgn0061943
    • FBtr0076448
    • FBpp0076176
    • FBtr0076449
    • FBpp0076177
    • FBtr0076450
    • FBpp0076178
    • FBtr0076451
    • FBpp0076179
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (236)