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. Author manuscript; available in PMC: 2023 Oct 18.
Published in final edited form as: Biochem Soc Trans. 2022 Jun 30;50(3):1227–1243. doi: 10.1042/BST20211225

Table 1.

List of bHLH-PAS genes and their functions from animals commonly studied for biomedical research

Species bHLH–PAS
gene name
bHLH–PAS protein name Protein class Function of protein Reference (PMID)
Caenorhabditis elegans aha-1 AHA-1 II Mutation is larval lethal
Neuronal specification with ahr-1
Hypoxic transcriptional response with hif-1
Ortholog of human ARNT/ARNT2
14757639
19632181
Mus musculus Ahr AHR I Angiogenesis and hematopoiesis
Xenobiotic metabolism
Development and function of the immune system
Immune checkpoint regulation
Regulation of viral replication
Liver development and function
15590894
16443691
33941684
7732381
8806883
34446714
33408169
33239445
32690969
27783946
27158906
26005855
21068375
20676095
20676092
Homo sapiens AHR AHR I Angiogenesis and hematopoiesis
Xenobiotic metabolism
Immune response
Mutation associated with retinitis pigmentosa
21976023
21999944
23723449
29726989
Caenorhabditis elegans ahr-1 AHR-1 I Neuron development
Aggregation on lawns of bacterial food
Lifespan
33349622
24655420
16919260
15136141
14757639
Danio rerio ahr1a Ahr1a I Unclear ( possible role in transcription in endothelial cells) 30907958
28817646
Danio rerio ahr1b Ahr1b I Unclear ( possible role in transcription in endothelial cells) 30907958
28817646
Danio rerio ahr2 Ahr2 I Xenobiotic metabolism
Immune response
Fin development
Vascular morphology
Craniofacial development
30907958
22242167
28817646
32384158
29494622
Mus musculus Ahrr AHRR indeterminate Regulate AHR activity
Modulate xenobiotic metabolism
17949687
9887096
Homo sapiens AHRR AHRR indeterminate Represses AHR transcription activity
Possible tumor supressor in cancers
17890447
18172554
Danio rerio ahrra Ahrra indeterminate Modulate Ahr2 activation by dioxins
Possible role in hematopoiesis
19494032
24675095
18000031
Danio rerio ahrrb Ahrrb indeterminate Modulate Ahr2 activation by dioxins 19494032
18000031
Mus musculus Arnt ARNT (HIF1β) II Binding partner of multiple Class I bHLH–PAS transcription factors 27782878
26245371
28602820
7592839 17023418
Homo sapiens ARNT ARNT (HIF1β) II Binding partner of multiple Class I bHLH–PAS transcription factors 8662957
27782878
Danio rerio arnt Arnt (Hif1β) II Binding partner of multiple Class I bHLH–PAS transcription factors 16306231
16936225
Edwards et al. bioRxiv 2022
Mus musculus Arnt2 ARNT2 II Binding partner of multiple Class I bHLH–PAS transcription factors 8657146
10873592
31585079
Homo sapiens ARNT2 ARNT2 II Binding partner of multiple Class I bHLH–PAS transcription factors
Mutation causes post-natal microcephaly, visual and renal anomalies
24022475
27782878
Danio rerio arnt2 Arnt2 II Binding partner of multiple Class I bHLH–PAS transcription factors 19374551
19234064
16936225
Edwards et al. bioRxiv 2022
Mus musculus Arntl ARNTL (BMAL1) II Binding partner of CLOCK and NPAS2
Circadian rhythm regulation
17417633
9160755
9616112
Homo sapiens ARNTL ARNTL (BMAL1) II Binding partner of CLOCK and NPAS2
Circadian rhythm regulation
17417633
9616112
27782878
Danio rerio arntl1a Arntl1a II Circadian rhythm regulation with clock genes
Ortholog of human ARNTL
23468616
Danio rerio arntl1b Arntl1b II Circadian rhythm regulation with clock genes
Ortholog of human ARNTL
23468616
Mus musculus ARNTL2 ARNTL2 (BMAL2) II Binding partner of CLOCK and NPAS2
Circadian rhythm regulation
11207387
20153195
Homo sapiens ARNTL2 ARNTL2 (BMAL2) II Binding partner of CLOCK and NPAS2 Circadian rhythm regulation 10964693
Danio rerio arntl2 Arntl2 (Bmal2) II Circadian rhythm regulation 11517315
Caenorhabditis elegans cky-1 CKY-1 I Speculated to be functionally similar to NPAS4 14701734
19284974
Drosophila melanogaster clk Clock I Circadian regulation with cyc
Regulation of circadian genes per and tim
11106063
Mus musculus Clock CLOCK I Circadian rhythm regulation
Circadian regulation of genes in multiple tissues
17417633
9160755
9616112
Homo sapiens CLOCK CLOCK I Circadian rhythm regulation
Circadian regulation of genes in multiple tissues
17417633
9616112
Danio rerio clocka Clocka I Circadian rhythm regulation
Mesoderm patterning
18800057
28687631
23468616
18569451
Danio rerio clockb Clockb I Speculated function circadian rhythm regulation 23468616
Drosophila melanogaster cyc Cycle II Circadian regulation with clk
Sensitization to cocaine
11106063
10409723
10446052
Drosophila melanogaster dysf Dysfusion I Required for tracheal migration, adhesion and fusion during development
Joint formation in legs
16914738
12897136
17652079
25329825
30080872
Drosophila melanogaster gce Germ-cell expressed bHLH–PAS indeterminate Responds to juvenile hormone
Can form heterodimer with met
26161662
16516852
Caenorhabditis elegans hif-1 HIF-1 I Stimulates gene expression in response to hypoxia
Essential for heat acclimation
11427734
30696318
12686697
Mus musculus Hif1a HIF1A I Up-regulates genes in response to hypoxia
Hematopoiesis
24141110
9436976
Homo sapiens HIF1A HIF1A I Regulates genes in response to hypoxia
Vascular devleopment
9436976
9782081
Danio rerio hif1aa Hif1aa I Up-regulates genes in response to hypoxia
Hematopoesis Neural protective
29339404
31314708
Danio rerio hif1ab Hif1ab I Up-regulates genes in response to hypoxia
Hematopoesis
Maturation of intestinal goblet cells
29339404
31314708
Danio rerio hif1al Hif1al I Up-regulates genes in response to hypoxia
Regulates erythropoiesis
Ortholog of human HIF3A
33037038
23034716
26052946
Danio rerio hif1al2 Hif1al2 I Maturation of intestinal goblet cells
Speculated up-regulates genes in response to hypoxia
27222589
Mus musculus Epas1 HIF2A I Regulates gene expression in response to hypoxia
Hematopoesis
Regulates vascular endothelial growth factor (VEGF)
Spermatogenesis
15626745
20181618
15851592
Homo sapiens EPAS1 HIF2A I Regulates gene expression in response to hypoxia
Vascular development
Mutation causes familial erythrocytosis
15851592
18184961
Danio rerio epas1a Epas1a I Up-regulates genes in response to hypoxia hematopoesis
ortholog of human EPAS1 (HIF2A)
29339404
Danio rerio epas1b Epas1b I Up-regulates genes in response to hypoxia Hematopoesis
ortholog of human EPAS1 (HIF2A)
29339404
Mus musculus Hif3a HIF3A I Inhibits/represses expression of hypoxia-induced genes 25626335
9840812
11734856
Homo sapiens HIF3A HIF3A I Modulates expression of hypoxia-induced genes
Dominant negative regulator of HIF1A
11573933
16126907
19694616
9840812
Caenorhabditis elegans hlh-34 HLH-34 I Unknown, expressed in a subset of interneurons
Ortholog of human NPAS3
34604715
Drosophila melanogaster met Methoprene-tolerant indeterminate Responds to juvenile hormone
Mutants are resistant to insecticide methoprene
Can form heterodimer with Germ-cell expressed
1592245
26161662
9501163
15720391
16516852
Mus musculus Ncoa1 NCOA1 indeterminate Transciptional coactivator of steroid hormone receptors
Energy metabolism and obesity
Possible regulator of hematopoiesis
8616895
7481822
12954634
12507421
27958775
Homo sapiens NCOA1 NCOA1 indeterminate Transcriptional coactivator of steroid hormone receptors
Transcriptional coactivator of STAT5 transcription factor
9427757
7481822
9223431
12954634
9252329
Danio rerio ncoa1 Ncoa1 indeterminate Transcriptional coactivator of steroid hormone receptors
Hematopoiesis
27958775
30448381
27156127
Mus musculus Ncoa2 NCOA2 indeterminate Transcriptional coactivator of steroid hormone receptors
Energy metabolism and obesity
Fertility
9238002
8670870
12507241
12138202
Homo sapiens NCOA2 NCOA2 indeterminate Transcriptional coactivator of steroid hormone receptors 9238002
8670870
Danio rerio ncoa2 Ncoa2 indeterminate Possible regulator of hematopoiesis
Transcriptional coactivator of steroid hormone receptors
18295965
18248177
Mus musculus Ncoa3 NCOA3 indeterminate Transcriptional coactivator of steroid hormone receptors
Important for hearing
Regulates adipogenesis hematopoiesis
Inflammation
33326993
9765300
Homo sapiens NCOA3 NCOA3 indeterminate Transcriptional coactivator of steroid hormone receptors
Mutation associated with progressive hearing loss
33326993
9765300
Danio rerio ncoa3 Ncoa3 indeterminate Development and function of otoliths Proposed trancriptional coactivator 33326993
Mus musculus Npas1 NPAS1 I Neural development
Behavior
15347806
9012850
28499489
Homo sapiens NPAS1 NPAS1 I Neuronal devleopment
Regulates genes involved in psychiatric diseases
9012850
28499489
Danio rerio npas1 Npas1 I Expressed in adult telencephalon
Speculated to be involved in neural development
25556858
Mus musculus Npas2 NPAS2 I Circadian rhythm regulation 11441147
26895328
11163178
17417633
Homo sapiens NPAS2 NPAS2 I Circadian rhythm regulation
Mutation associated with nonobstructive azoospermia
11441147
11163178
25956372
Danio rerio npas2 Npas2 I Not characterized none
Mus musculus Npas3 NPAS3 I Neural development
behavior
Lung development and function
15347806
28499489
19581591
Homo sapiens NPAS3 NPAS3 I Neural development
Mutations associated with schizophrenia
12746393
21709683
22228753
15924306
Danio rerio npas3 Npas3 I Insufficent data
Speculated to be involved in neuronal development
-
Mus musculus Npas4 NPAS4 I Synaptic plasticity
Memory
24201284
22194569
18815592
Homo sapiens NPAS4 NPAS4 I Variants associated with intellectual disability 33758288
24465693
Danio rerio npas4a Npas4a I Forebrain development
Angiogenesis of intersegmental vessicles
25538572
28082451
Danio rerio npas4b Npas4b I Angiogenesis of intersegmental vessicles 28082451
Danio rerio npas4l Npas4l I Master regulator of hemato-vascular specification
Formation of lens cataract
27411634
7588049
30861003
Drosophila melanogaster sim Single-minded I Midline central nervous system development
Axonal extension
3345560
3345559
2242162
1760843
16439478
Mus musculus Sim1 SIM1 I Neuronal development
Hypothalamus development and function
Obesity
9784500
16291793
12947113
10587584
Homo sapiens SIM1 SIM1 I Obesity
Neuroendocrine development
10587584
16924270
33434169
Danio rerio sim1a Sim1a I Hypothalamus development and function
Axonal guidance
19234064
23222439
16691572
Danio rerio sim1b Sim1b I Hypothalamus development and function 19234064
16691572
Mus musculus Sim2 SIM1 I Development and function of neuroendocrine cells
Axon guidance
Overexpression causes Down syndrome phenotypes
Growth and integrity of skeleton
14988428
12024028
16291793
10400987
10915774
Homo sapiens SIM2 SIM2 I Promotes glioblastoma cell invasion
Part of Down syndrome critical region on chromosome 21
7647800
7568099
20448453
15946822
Danio rerio sim2 Sim2 I Not characterized none
Drosophila melanogaster sima Similar I Hypoxic response
Trachea development
Ortholog of human HIF1A
12215541
19285057
Drosophila melanogaster ss Spineless I Distal limb formation
Mutants have reduced sensory bristles
Ortholog of human AHR
10433921
17084833
9573046
Drosophila melanogaster tgo Tango II Neuronal devleopment
Glial migration
Trachea development
Limb formation
Coregulator of class I genes
Ortholog of human ARNT/ARNT2
8557198
10433921
9409674
Drosophila melanogaster trh Trachealess I Development of salivary duct
Development of trachea distal limb Development
Most similar to human NPAS1, NPAS3
17187773
8557198