Estrogen receptor beta

Protein-coding gene in the species Homo sapiens

ESR2

| Available structures

PDB| Ortholog search: PDBe RCSB
| List of PDB id codes

4J26, 1L2J, 1NDE, 1QKM, 1U3Q, 1U3R, 1U3S, 1U9E, 1X76, 1X78, 1X7B, 1X7J, 1YY4, 1YYE, 1ZAF, 2FSZ, 2GIU, 2I0G, 2JJ3, 2NV7, 2QTU, 2YJD, 2YLY, 2Z4B, 3OLL, 3OLS, 3OMO, 3OMP, 3OMQ, 4J24, 4ZI1

Identifiers
Aliases| ESR2, ER-BETA, ESR-BETA, ESRB, ESTRB, Erb, NR3A2, estrogen receptor 2, ODG8
External IDs| OMIM: 601663; MGI: 109392; HomoloGene: 1100; GeneCards: ESR2; OMA:ESR2 - orthologs
| Gene location (Human)

Chr.| Chromosome 14 (human)[1]

Band| 14q23.2-q23.3| Start| 64,084,232 bp[1]
End| 64,338,112 bp[1]
| Gene location (Mouse)

Chr.| Chromosome 12 (mouse)[2]

Band| 12 C3|12 33.52 cM| Start| 76,167,193 bp[2]
End| 76,224,033 bp[2]
| RNA expression pattern

Bgee| | Human| Mouse (ortholog)

|

| Top expressed in

* right adrenal gland

* right adrenal cortex

* left adrenal gland

* left adrenal cortex

* secondary oocyte

* left ovary

* right ovary

* granulocyte

* epithelium of colon

* left testis

| Top expressed in

* zygote

* secondary oocyte

* primary oocyte

* urethra

* female urethra

* male urethra

* cumulus cell

* lumbar subsegment of spinal cord

* embryo

* ovary

More reference expression data
BioGPS| |

More reference expression data
| Gene ontology

Molecular function|

* DNA-binding transcription factor activity
* nuclear receptor activity
* metal ion binding
* steroid hormone receptor activity
* steroid binding
* zinc ion binding
* protein binding
* DNA binding
* sequence-specific DNA binding
* transcription coactivator activity
* receptor antagonist activity
* lipid binding
* enzyme binding
* estrogen response element binding
* core promoter sequence-specific DNA binding
* estrogen receptor activity
* identical protein binding
* DNA-binding transcription factor activity, RNA polymerase II-specific
* RNA polymerase II cis-regulatory region sequence-specific DNA binding

Cellular component|

* mitochondrion
* nucleus
* nucleoplasm
* extracellular region

Biological process|

* regulation of transcription, DNA-templated
* transcription, DNA-templated
* negative regulation of cell growth
* transcription initiation from RNA polymerase II promoter
* positive regulation of DNA-binding transcription factor activity
* cell-cell signaling
* signal transduction
* extracellular negative regulation of signal transduction
* positive regulation of transcription, DNA-templated
* negative regulation of transcription by RNA polymerase II
* intracellular estrogen receptor signaling pathway
* positive regulation of transcription by RNA polymerase II
* cellular response to estradiol stimulus
* negative regulation of signaling receptor activity

Sources:Amigo / QuickGO
| Orthologs

Species| Human| Mouse
Entrez| |

2100

13983

Ensembl| |

ENSG00000140009

ENSMUSG00000021055

UniProt| |

Q92731

O08537

RefSeq (mRNA)| | NM_001040275
NM_001040276
NM_001214902
NM_001214903
NM_001271876

NM_001271877
NM_001291712
NM_001291723
NM_001437

NM_010157
NM_207707

RefSeq (protein)| | NP_001035365
NP_001201831
NP_001258805
NP_001258806
NP_001278641

NP_001278652
NP_001428

NP_034287
NP_997590

Location (UCSC)| Chr 14: 64.08 – 64.34 Mb| Chr 12: 76.17 – 76.22 Mb
PubMed search| [3]| [4]
Wikidata
| View/Edit Human| View/Edit Mouse

|

Estrogen receptor beta (ERβ) also known as NR3A2 (nuclear receptor subfamily 3, group A, member 2) is one of two main types of estrogen receptor—a nuclear receptor which is activated by the sex hormone estrogen.[5] In humans ERβ is encoded by the ESR2 gene.[6]

## Function

[edit]

ERβ is a member of the family of estrogen receptors and the superfamily of nuclear receptor transcription factors. The gene product contains an N-terminal DNA binding domain and C-terminal ligand binding domain and is localized to the nucleus, cytoplasm, and mitochondria. Upon binding to 17-β-estradiol, estriol or related ligands, the encoded protein forms homo-dimers or hetero-dimers with estrogen receptor α that interact with specific DNA sequences to activate transcription. Some isoforms dominantly inhibit the activity of other estrogen receptor family members. Several alternatively spliced transcript variants of this gene have been described, but the full-length nature of some of these variants has not been fully characterized.[7]

ERβ may inhibit cell proliferation and opposes the actions of ERα in reproductive tissue.[8] ERβ may also have an important role in adaptive function of the lung during pregnancy.[9]

ERβ is a potent tumor suppressor and plays a crucial role in many cancer types such as prostate cancer and ovarian cancer.[10][11]

### Mammary gland

[edit]

ERβ knockout mice show normal mammary gland development at puberty and are able to lactate normally.[12][13][14] The mammary glands of adult virgin female mice are indistinguishable from those of age-matched wild-type virgin female mice.[12] This is in contrast to ERα knockout mice, in which a complete absence of mammary gland development at puberty and thereafter is observed.[12][14] Administration of the selective ERβ agonist ERB-041 to immature ovariectomized female rats produced no observable effects in the mammary glands, further indicating that the ERβ is non-mammotrophic.[15][14][16]

Although ERβ is not required for pubertal development of the mammary glands, it may be involved in terminal differentiation in pregnancy, and may also be necessary to maintain the organization and differentiation of mammary epithelium in adulthood.[17][18] In old female ERβ knockout mice, severe cystic mammary disease that is similar in appearance to postmenopausal mastopathy develops, whereas this does not occur in aged wild-type female mice.[13] However, ERβ knockout mice are not only deficient in ERβ signaling in the mammary glands, but also have deficient progesterone exposure due to impairment of corpora lutea formation.[13][17] This complicates attribution of the preceding findings to mammary ERβ signaling.[13][17]

Selective ERβ agonism with diarylpropionitrile (DPN) has been found to counteract the proliferative effects in the mammary glands of selective ERα agonism with propylpyrazoletriol (PPT) in ovariectomized postmenopausal female rats.[19][20] Similarly, overexpression of ERβ via lentiviral infection in mature virgin female rats decreases mammary proliferation.[20] ERα signaling has proliferative effects in both normal breast and breast cancer cell lines, whereas ERβ has generally antiproliferative effects in such cell lines.[17] However, ERβ has been found to have proliferative effects in some breast cell lines.[17]

Expression of ERα and ERβ in the mammary gland have been found to vary throughout the menstrual cycle and in an ovariectomized state in female rats.[20] Whereas mammary ERα in rhesus macaques is downregulated in response to increased estradiol levels, expression of ERβ in the mammary glands is not.[21] Expression of ERα and ERβ in the mammary glands also differs throughout life in female mice.[22] Mammary ERα expression is higher and mammary ERβ expression lower in younger female mice, while mammary ERα expression is lower and mammary ERβ expression higher in older female mice as well as in parous female mice.[22] Mammary proliferation and estrogen sensitivity is higher in young female mice than in old or parous female mice, particularly during pubertal mammary gland development.[22]

## Tissue distribution

[edit]

ERβ is expressed by many tissues including the uterus,[23] blood monocytes and tissue macrophages, colonic and pulmonary epithelial cells and in prostatic epithelium and in malignant counterparts of these tissues. Also, ERβ is found throughout the brain at different concentrations in different neuron clusters.[24][25] ERβ is also highly expressed in normal breast epithelium, although its expression declines with cancer progression.[26] ERβ is expressed in all subtypes of breast cancer.[27] Controversy regarding ERβ protein expression has hindered study of ERβ, but highly sensitive monoclonal antibodies have been produced and well-validated to address these issues.[28]

## ERβ abnormalities

[edit]

ERβ function is related to various cardiovascular targets including ATP-binding cassette transporter A1 (ABCA1) and apolipoprotein A1 (ApoA-1). Polymorphism may affect ERβ function and lead to altered responses in postmenopausal women receiving hormone replacement therapy.[29] Abnormalities in gene expression associated with ERβ have also been linked to autism spectrum disorder.[30]

## Disease

[edit]

### Cardiovascular disease

[edit]

Mutations in ERβ have been shown to influence cardiomyocytes, the cells that comprise the largest part of the heart, and can lead to an increased risk of cardiovascular disease (CVD). There is a disparity in prevalence of CVD between pre- and post-menopausal women, and the difference can be attributed to estrogen levels. Many types of ERβ receptors exist in order to help regulate gene expression and subsequent health in the body, but binding of 17βE2 (a naturally occurring estrogen) specifically improves cardiac metabolism. The heart utilizes a lot of energy in the form of ATP to properly pump blood and maintain physiological requirements in order to live, and 17βE2 helps by increasing these myocardial ATP levels and respiratory function.[31]

In addition, 17βE2 can alter myocardial signaling pathways and stimulate myocyte regeneration, which can aid in inhibiting myocyte cell death. The ERβ signaling pathway plays a role in both vasodilation and arterial dilation, which contributes to an individual having a healthy heart rate and a decrease in blood pressure. This regulation can increase endothelial function and arterial perfusion, both of which are important to myocyte health. Thus, alterations in this signaling pathways due to ERβ mutation could lead to myocyte cell death from physiological stress. While ERα has a more profound role in regeneration after myocyte cell death, ERβ can still help by increasing endothelial progenitor cell activation and subsequent cardiac function.[32]

### Alzheimer's disease

[edit]

Genetic variation in ERβ is both sex and age dependent and ERβ polymorphism can lead to accelerated brain aging, cognitive impairment, and development of AD pathology. Similar to CVD, post-menopausal women have an increased risk of developing Alzheimer's disease (AD) due to a loss of estrogen, which affects proper aging of the hippocampus, neural survival and regeneration, and amyloid metabolism. ERβ mRNA is highly expressed in hippocampal formation, an area of the brain that is associated with memory. This expression contributes to increased neuronal survival and helps protect against neurodegenerative diseases such as AD. The pathology of AD is also associated with accumulation of amyloid beta peptide (Aβ). While a proper concentration of Aβ in the brain is important for healthy functioning, too much can lead to cognitive impairment. Thus, ERβ helps control Aβ levels by maintaining the protein it is derived from, β-amyloid precursor protein. ERβ helps by up-regulating insulin-degrading enzyme (IDE), which leads to β-amyloid degradation when accumulation levels begin to rise. However, in AD, lack of ERβ causes a decrease in this degradation and an increase in plaque build-up.[33]

ERβ also plays a role in regulating APOE, a risk factor for AD that redistributes lipids across cells. APOE expression in the hippocampus is specifically regulated by 17βE2, affecting learning and memory in individuals afflicted with AD. Thus, estrogen therapy via an ERβ-targeted approach can be used as a prevention method for AD either before or at the onset of menopause. Interactions between ERα and ERβ can lead to antagonistic actions in the brain, so an ERβ-targeted approach can increase therapeutic neural responses independently of ERα. Therapeutically, ERβ can be used in both men and women in order to regulate plaque formation in the brain.[34]

## Neuroprotective benefits

[edit]

### Synaptic strength and plasticity

[edit]

ERβ levels can dictate both synaptic strength and neuroplasticity through neural structure modifications. Variations in endogenous estrogen levels cause changes in dendritic architecture in the hippocampus, which affects neural signaling and plasticity. Specifically, lower estrogen levels lead to decreased dendritic spines and improper signaling, inhibiting plasticity of the brain. However, treatment of 17βE2 can reverse this affect, giving it the ability to modify hippocampal structure. As a result of the relationship between dendritic architecture and long-term potentiation (LTP), ERβ can enhance LTP and lead to an increase in synaptic strength. Furthermore, 17βE2 promotes neurogenesis in developing hippocampal neurons and neurons in the subventricular zone and dentate gyrus of the adult human brain. Specifically, ERβ increases the proliferation of progenitor cells to create new neurons and can be increased later in life through 17βE2 treatment.[35][36]

## Ligands

[edit]

### Agonists

[edit]

#### Non-selective

[edit]

* Endogenous estrogens (e.g., estradiol, estrone, estriol, estetrol)
* Natural estrogens (e.g., conjugated estrogens)
* Synthetic estrogens (e.g., ethinylestradiol, diethylstilbestrol)

#### Selective

[edit]

Agonists of ERβ selective over ERα include:

* 20-Hydroxyecdysone (ecdysterone, 20-HE, 20-E) — phytoecdysteroid[37]
* 3β-Androstanediol (3β-diol) – endogenous
* 8β-VE2
* AC-186
* Apigenin – phytoestrogen[38]
* Daidzein – phytoestrogen[38]
* DCW234
* Dehydroepiandrosterone (DHEA) – endogenous
* Diarylpropionitrile (DPN)
* ERB-79 and its active enantiomer ERB-26
* ERB-196 (WAY-202196)
* Erteberel (SERBA-1, LY-500307)
* FERb 033 – 62-fold selectivity for ERβ over ERα[39]
* Genistein – phytoestrogen; 16-fold selectivity for ERβ over ERα[38]
* Liquiritigenin (Menerba) – phytoestrogen[38]
* Penduletin – phytoestrogen[38]
* Prinaberel (ERB-041, WAY-202041)
* S-Equol ((S)-4',7-isoflavandiol) – phytoestrogen; 13-fold selectivity for ERβ over ERα[38]
* WAY-166818
* WAY-200070
* WAY-214156

### Antagonists

[edit]

#### Non-selective

[edit]

* Selective estrogen receptor modulators (e.g., tamoxifen, raloxifene)[40]
* Antiestrogens (e.g., fulvestrant, ICI-164384)

#### Selective

[edit]

Antagonists of ERβ selective over ERα include:

* PHTPP
* (R,R)-Tetrahydrochrysene ((R,R)-THC) – actually not selective over ERα, but rather an agonist instead of antagonist of ERα

### Affinities

[edit]

* v
* t
* e

Affinities of estrogen receptor ligands for the ERα and ERβ Ligand | Other names | Relative binding affinities (RBA, %)a | Absolute binding affinities (Ki, nM)a | Action

|
|
|
|

ERα | ERβ | ERα | ERβ
Estradiol | E2; 17β-Estradiol | 100 | 100 | 0.115 (0.04–0.24) | 0.15 (0.10–2.08) | Estrogen
Estrone | E1; 17-Ketoestradiol | 16.39 (0.7–60) | 6.5 (1.36–52) | 0.445 (0.3–1.01) | 1.75 (0.35–9.24) | Estrogen
Estriol | E3; 16α-OH-17β-E2 | 12.65 (4.03–56) | 26 (14.0–44.6) | 0.45 (0.35–1.4) | 0.7 (0.63–0.7) | Estrogen
Estetrol | E4; 15α,16α-Di-OH-17β-E2 | 4.0 | 3.0 | 4.9 | 19 | Estrogen
Alfatradiol | 17α-Estradiol | 20.5 (7–80.1) | 8.195 (2–42) | 0.2–0.52 | 0.43–1.2 | Metabolite
16-Epiestriol | 16β-Hydroxy-17β-estradiol | 7.795 (4.94–63) | 50 | ? | ? | Metabolite
17-Epiestriol | 16α-Hydroxy-17α-estradiol | 55.45 (29–103) | 79–80 | ? | ? | Metabolite
16,17-Epiestriol | 16β-Hydroxy-17α-estradiol | 1.0 | 13 | ? | ? | Metabolite
2-Hydroxyestradiol | 2-OH-E2 | 22 (7–81) | 11–35 | 2.5 | 1.3 | Metabolite
2-Methoxyestradiol | 2-MeO-E2 | 0.0027–2.0 | 1.0 | ? | ? | Metabolite
4-Hydroxyestradiol | 4-OH-E2 | 13 (8–70) | 7–56 | 1.0 | 1.9 | Metabolite
4-Methoxyestradiol | 4-MeO-E2 | 2.0 | 1.0 | ? | ? | Metabolite
2-Hydroxyestrone | 2-OH-E1 | 2.0–4.0 | 0.2–0.4 | ? | ? | Metabolite
2-Methoxyestrone | 2-MeO-E1 | <0.001–<1 | <1 | ? | ? | Metabolite
4-Hydroxyestrone | 4-OH-E1 | 1.0–2.0 | 1.0 | ? | ? | Metabolite
4-Methoxyestrone | 4-MeO-E1 | <1 | <1 | ? | ? | Metabolite
16α-Hydroxyestrone | 16α-OH-E1; 17-Ketoestriol | 2.0–6.5 | 35 | ? | ? | Metabolite
2-Hydroxyestriol | 2-OH-E3 | 2.0 | 1.0 | ? | ? | Metabolite
4-Methoxyestriol | 4-MeO-E3 | 1.0 | 1.0 | ? | ? | Metabolite
Estradiol sulfate | E2S; Estradiol 3-sulfate | <1 | <1 | ? | ? | Metabolite
Estradiol disulfate | Estradiol 3,17β-disulfate | 0.0004 | ? | ? | ? | Metabolite
Estradiol 3-glucuronide | E2-3G | 0.0079 | ? | ? | ? | Metabolite
Estradiol 17β-glucuronide | E2-17G | 0.0015 | ? | ? | ? | Metabolite
Estradiol 3-gluc. 17β-sulfate | E2-3G-17S | 0.0001 | ? | ? | ? | Metabolite
Estrone sulfate | E1S; Estrone 3-sulfate | <1 | <1 | >10 | >10 | Metabolite
Estradiol benzoate | EB; Estradiol 3-benzoate | 10 | ? | ? | ? | Estrogen
Estradiol 17β-benzoate | E2-17B | 11.3 | 32.6 | ? | ? | Estrogen
Estrone methyl ether | Estrone 3-methyl ether | 0.145 | ? | ? | ? | Estrogen
ent-Estradiol | 1-Estradiol | 1.31–12.34 | 9.44–80.07 | ? | ? | Estrogen
Equilin | 7-Dehydroestrone | 13 (4.0–28.9) | 13.0–49 | 0.79 | 0.36 | Estrogen
Equilenin | 6,8-Didehydroestrone | 2.0–15 | 7.0–20 | 0.64 | 0.62 | Estrogen
17β-Dihydroequilin | 7-Dehydro-17β-estradiol | 7.9–113 | 7.9–108 | 0.09 | 0.17 | Estrogen
17α-Dihydroequilin | 7-Dehydro-17α-estradiol | 18.6 (18–41) | 14–32 | 0.24 | 0.57 | Estrogen
17β-Dihydroequilenin | 6,8-Didehydro-17β-estradiol | 35–68 | 90–100 | 0.15 | 0.20 | Estrogen
17α-Dihydroequilenin | 6,8-Didehydro-17α-estradiol | 20 | 49 | 0.50 | 0.37 | Estrogen
Δ8-Estradiol | 8,9-Dehydro-17β-estradiol | 68 | 72 | 0.15 | 0.25 | Estrogen
Δ8-Estrone | 8,9-Dehydroestrone | 19 | 32 | 0.52 | 0.57 | Estrogen
Ethinylestradiol | EE; 17α-Ethynyl-17β-E2 | 120.9 (68.8–480) | 44.4 (2.0–144) | 0.02–0.05 | 0.29–0.81 | Estrogen
Mestranol | EE 3-methyl ether | ? | 2.5 | ? | ? | Estrogen
Moxestrol | RU-2858; 11β-Methoxy-EE | 35–43 | 5–20 | 0.5 | 2.6 | Estrogen
Methylestradiol | 17α-Methyl-17β-estradiol | 70 | 44 | ? | ? | Estrogen
Diethylstilbestrol | DES; Stilbestrol | 129.5 (89.1–468) | 219.63 (61.2–295) | 0.04 | 0.05 | Estrogen
Hexestrol | Dihydrodiethylstilbestrol | 153.6 (31–302) | 60–234 | 0.06 | 0.06 | Estrogen
Dienestrol | Dehydrostilbestrol | 37 (20.4–223) | 56–404 | 0.05 | 0.03 | Estrogen
Benzestrol (B2) | – | 114 | ? | ? | ? | Estrogen
Chlorotrianisene | TACE | 1.74 | ? | 15.30 | ? | Estrogen
Triphenylethylene | TPE | 0.074 | ? | ? | ? | Estrogen
Triphenylbromoethylene | TPBE | 2.69 | ? | ? | ? | Estrogen
Tamoxifen | ICI-46,474 | 3 (0.1–47) | 3.33 (0.28–6) | 3.4–9.69 | 2.5 | SERM
Afimoxifene | 4-Hydroxytamoxifen; 4-OHT | 100.1 (1.7–257) | 10 (0.98–339) | 2.3 (0.1–3.61) | 0.04–4.8 | SERM
Toremifene | 4-Chlorotamoxifen; 4-CT | ? | ? | 7.14–20.3 | 15.4 | SERM
Clomifene | MRL-41 | 25 (19.2–37.2) | 12 | 0.9 | 1.2 | SERM
Cyclofenil | F-6066; Sexovid | 151–152 | 243 | ? | ? | SERM
Nafoxidine | U-11,000A | 30.9–44 | 16 | 0.3 | 0.8 | SERM
Raloxifene | – | 41.2 (7.8–69) | 5.34 (0.54–16) | 0.188–0.52 | 20.2 | SERM
Arzoxifene | LY-353,381 | ? | ? | 0.179 | ? | SERM
Lasofoxifene | CP-336,156 | 10.2–166 | 19.0 | 0.229 | ? | SERM
Ormeloxifene | Centchroman | ? | ? | 0.313 | ? | SERM
Levormeloxifene | 6720-CDRI; NNC-460,020 | 1.55 | 1.88 | ? | ? | SERM
Ospemifene | Deaminohydroxytoremifene | 0.82–2.63 | 0.59–1.22 | ? | ? | SERM
Bazedoxifene | – | ? | ? | 0.053 | ? | SERM
Etacstil | GW-5638 | 4.30 | 11.5 | ? | ? | SERM
ICI-164,384 | – | 63.5 (3.70–97.7) | 166 | 0.2 | 0.08 | Antiestrogen
Fulvestrant | ICI-182,780 | 43.5 (9.4–325) | 21.65 (2.05–40.5) | 0.42 | 1.3 | Antiestrogen
Propylpyrazoletriol | PPT | 49 (10.0–89.1) | 0.12 | 0.40 | 92.8 | ERα agonist
16α-LE2 | 16α-Lactone-17β-estradiol | 14.6–57 | 0.089 | 0.27 | 131 | ERα agonist
16α-Iodo-E2 | 16α-Iodo-17β-estradiol | 30.2 | 2.30 | ? | ? | ERα agonist
Methylpiperidinopyrazole | MPP | 11 | 0.05 | ? | ? | ERα antagonist
Diarylpropionitrile | DPN | 0.12–0.25 | 6.6–18 | 32.4 | 1.7 | ERβ agonist
8β-VE2 | 8β-Vinyl-17β-estradiol | 0.35 | 22.0–83 | 12.9 | 0.50 | ERβ agonist
Prinaberel | ERB-041; WAY-202,041 | 0.27 | 67–72 | ? | ? | ERβ agonist
ERB-196 | WAY-202,196 | ? | 180 | ? | ? | ERβ agonist
Erteberel | SERBA-1; LY-500,307 | ? | ? | 2.68 | 0.19 | ERβ agonist
SERBA-2 | – | ? | ? | 14.5 | 1.54 | ERβ agonist
Coumestrol | – | 9.225 (0.0117–94) | 64.125 (0.41–185) | 0.14–80.0 | 0.07–27.0 | Xenoestrogen
Genistein | – | 0.445 (0.0012–16) | 33.42 (0.86–87) | 2.6–126 | 0.3–12.8 | Xenoestrogen
Equol | – | 0.2–0.287 | 0.85 (0.10–2.85) | ? | ? | Xenoestrogen
Daidzein | – | 0.07 (0.0018–9.3) | 0.7865 (0.04–17.1) | 2.0 | 85.3 | Xenoestrogen
Biochanin A | – | 0.04 (0.022–0.15) | 0.6225 (0.010–1.2) | 174 | 8.9 | Xenoestrogen
Kaempferol | – | 0.07 (0.029–0.10) | 2.2 (0.002–3.00) | ? | ? | Xenoestrogen
Naringenin | – | 0.0054 (<0.001–0.01) | 0.15 (0.11–0.33) | ? | ? | Xenoestrogen
8-Prenylnaringenin | 8-PN | 4.4 | ? | ? | ? | Xenoestrogen
Quercetin | – | <0.001–0.01 | 0.002–0.040 | ? | ? | Xenoestrogen
Ipriflavone | – | <0.01 | <0.01 | ? | ? | Xenoestrogen
Miroestrol | – | 0.39 | ? | ? | ? | Xenoestrogen
Deoxymiroestrol | – | 2.0 | ? | ? | ? | Xenoestrogen
β-Sitosterol | – | <0.001–0.0875 | <0.001–0.016 | ? | ? | Xenoestrogen
Resveratrol | – | <0.001–0.0032 | ? | ? | ? | Xenoestrogen
α-Zearalenol | – | 48 (13–52.5) | ? | ? | ? | Xenoestrogen
β-Zearalenol | – | 0.6 (0.032–13) | ? | ? | ? | Xenoestrogen
Zeranol | α-Zearalanol | 48–111 | ? | ? | ? | Xenoestrogen
Taleranol | β-Zearalanol | 16 (13–17.8) | 14 | 0.8 | 0.9 | Xenoestrogen
Zearalenone | ZEN | 7.68 (2.04–28) | 9.45 (2.43–31.5) | ? | ? | Xenoestrogen
Zearalanone | ZAN | 0.51 | ? | ? | ? | Xenoestrogen
Bisphenol A | BPA | 0.0315 (0.008–1.0) | 0.135 (0.002–4.23) | 195 | 35 | Xenoestrogen
Endosulfan | EDS | <0.001–<0.01 | <0.01 | ? | ? | Xenoestrogen
Kepone | Chlordecone | 0.0069–0.2 | ? | ? | ? | Xenoestrogen
o,p'-DDT | – | 0.0073–0.4 | ? | ? | ? | Xenoestrogen
p,p'-DDT | – | 0.03 | ? | ? | ? | Xenoestrogen
Methoxychlor | p,p'-Dimethoxy-DDT | 0.01 (<0.001–0.02) | 0.01–0.13 | ? | ? | Xenoestrogen
HPTE | Hydroxychlor; p,p'-OH-DDT | 1.2–1.7 | ? | ? | ? | Xenoestrogen
Testosterone | T; 4-Androstenolone | <0.0001–<0.01 | <0.002–0.040 | >5000 | >5000 | Androgen
Dihydrotestosterone | DHT; 5α-Androstanolone | 0.01 (<0.001–0.05) | 0.0059–0.17 | 221–>5000 | 73–1688 | Androgen
Nandrolone | 19-Nortestosterone; 19-NT | 0.01 | 0.23 | 765 | 53 | Androgen
Dehydroepiandrosterone | DHEA; Prasterone | 0.038 (<0.001–0.04) | 0.019–0.07 | 245–1053 | 163–515 | Androgen
5-Androstenediol | A5; Androstenediol | 6 | 17 | 3.6 | 0.9 | Androgen
4-Androstenediol | – | 0.5 | 0.6 | 23 | 19 | Androgen
4-Androstenedione | A4; Androstenedione | <0.01 | <0.01 | >10000 | >10000 | Androgen
3α-Androstanediol | 3α-Adiol | 0.07 | 0.3 | 260 | 48 | Androgen
3β-Androstanediol | 3β-Adiol | 3 | 7 | 6 | 2 | Androgen
Androstanedione | 5α-Androstanedione | <0.01 | <0.01 | >10000 | >10000 | Androgen
Etiocholanedione | 5β-Androstanedione | <0.01 | <0.01 | >10000 | >10000 | Androgen
Methyltestosterone | 17α-Methyltestosterone | <0.0001 | ? | ? | ? | Androgen
Ethinyl-3α-androstanediol | 17α-Ethynyl-3α-adiol | 4.0 | <0.07 | ? | ? | Estrogen
Ethinyl-3β-androstanediol | 17α-Ethynyl-3β-adiol | 50 | 5.6 | ? | ? | Estrogen
Progesterone | P4; 4-Pregnenedione | <0.001–0.6 | <0.001–0.010 | ? | ? | Progestogen
Norethisterone | NET; 17α-Ethynyl-19-NT | 0.085 (0.0015–<0.1) | 0.1 (0.01–0.3) | 152 | 1084 | Progestogen
Norethynodrel | 5(10)-Norethisterone | 0.5 (0.3–0.7) | <0.1–0.22 | 14 | 53 | Progestogen
Tibolone | 7α-Methylnorethynodrel | 0.5 (0.45–2.0) | 0.2–0.076 | ? | ? | Progestogen
Δ4-Tibolone | 7α-Methylnorethisterone | 0.069–<0.1 | 0.027–<0.1 | ? | ? | Progestogen
3α-Hydroxytibolone | – | 2.5 (1.06–5.0) | 0.6–0.8 | ? | ? | Progestogen
3β-Hydroxytibolone | – | 1.6 (0.75–1.9) | 0.070–0.1 | ? | ? | Progestogen
Footnotes: a = (1) Binding affinity values are of the format "median (range)" (# (#–#)), "range" (#–#), or "value" (#) depending on the values available. The full sets of values within the ranges can be found in the Wiki code. (2) Binding affinities were determined via displacement studies in a variety of in-vitro systems with labeled estradiol and human ERα and ERβ proteins (except the ERβ values from Kuiper et al. (1997), which are rat ERβ). Sources: See template page.

## Interactions

[edit]

Estrogen receptor beta has been shown to interact with:

* CCND1,[41]
* ESR1[42][43]
* MAD2L1,[43]
* NCOA3,[44][45]
* NCOA6,[46][47]
* RBM39,[48] and
* SRC.[49][50]

## References

[edit]

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## Further reading

[edit]

* Pettersson K, Gustafsson JA (2001). "Role of estrogen receptor beta in estrogen action". Annual Review of Physiology. 63: 165–192\. doi:10.1146/annurev.physiol.63.1.165. PMID 11181953.165-192&rft.date=2001&rft_id=info:doi/10.1146/annurev.physiol.63.1.165&rft_id=info:pmid/11181953&rft.aulast=Pettersson&rft.aufirst=K&rft.au=Gustafsson,+JA&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Warner M, Saji S, Gustafsson JA (July 2000). "The normal and malignant mammary gland: a fresh look with ER beta onboard". Journal of Mammary Gland Biology and Neoplasia. 5 (3): 289–294\. doi:10.1023/A:1009598828267. PMID 14973391. S2CID 34129981.289-294&rft.date=2000-07&rft_id=https://api.semanticscholar.org/CorpusID:34129981#id-name=S2CID&rft_id=info:pmid/14973391&rft_id=info:doi/10.1023/A:1009598828267&rft.aulast=Warner&rft.aufirst=M&rft.au=Saji,+S&rft.au=Gustafsson,+JA&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Saxon LK, Turner CH (February 2005). "Estrogen receptor beta: the antimechanostat?". Bone. 36 (2): 185–192\. doi:10.1016/j.bone.2004.08.003. PMID 15780944.185-192&rft.date=2005-02&rft_id=info:doi/10.1016/j.bone.2004.08.003&rft_id=info:pmid/15780944&rft.aulast=Saxon&rft.aufirst=LK&rft.au=Turner,+CH&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Halachmi S, Marden E, Martin G, MacKay H, Abbondanza C, Brown M (June 1994). "Estrogen receptor-associated proteins: possible mediators of hormone-induced transcription". Science. 264 (5164): 1455–1458\. Bibcode:1994Sci...264.1455H. doi:10.1126/science.8197458. PMID 8197458.1455-1458&rft.date=1994-06&rft_id=info:pmid/8197458&rft_id=info:doi/10.1126/science.8197458&rft_id=info:bibcode/1994Sci...264.1455H&rft.aulast=Halachmi&rft.aufirst=S&rft.au=Marden,+E&rft.au=Martin,+G&rft.au=MacKay,+H&rft.au=Abbondanza,+C&rft.au=Brown,+M&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Schwabe JW, Chapman L, Finch JT, Rhodes D (November 1993). "The crystal structure of the estrogen receptor DNA-binding domain bound to DNA: how receptors discriminate between their response elements". Cell. 75 (3): 567–578\. doi:10.1016/0092-8674(93)90390-C. PMID 8221895. S2CID 20795587.567-578&rft.date=1993-11&rft_id=https://api.semanticscholar.org/CorpusID:20795587#id-name=S2CID&rft_id=info:pmid/8221895&rft_id=info:doi/10.1016/0092-8674(93)90390-C&rft.aulast=Schwabe&rft.aufirst=JW&rft.au=Chapman,+L&rft.au=Finch,+JT&rft.au=Rhodes,+D&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Chen H, Lin RJ, Schiltz RL, Chakravarti D, Nash A, Nagy L, et al. (August 1997). "Nuclear receptor coactivator ACTR is a novel histone acetyltransferase and forms a multimeric activation complex with P/CAF and CBP/p300". Cell. 90 (3): 569–580\. doi:10.1016/S0092-8674(00)80516-4. PMID 9267036. S2CID 15284825.569-580&rft.date=1997-08&rft_id=https://api.semanticscholar.org/CorpusID:15284825#id-name=S2CID&rft_id=info:pmid/9267036&rft_id=info:doi/10.1016/S0092-8674(00)80516-4&rft.aulast=Chen&rft.aufirst=H&rft.au=Lin,+RJ&rft.au=Schiltz,+RL&rft.au=Chakravarti,+D&rft.au=Nash,+A&rft.au=Nagy,+L&rft.au=Privalsky,+ML&rft.au=Nakatani,+Y&rft.au=Evans,+RM&rft_id=https://doi.org/10.1016%2FS0092-8674%2800%2980516-4&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Pace P, Taylor J, Suntharalingam S, Coombes RC, Ali S (October 1997). "Human estrogen receptor beta binds DNA in a manner similar to and dimerizes with estrogen receptor alpha". The Journal of Biological Chemistry. 272 (41): 25832–25838\. doi:10.1074/jbc.272.41.25832. PMID 9325313.25832-25838&rft.date=1997-10&rft_id=info:doi/10.1074/jbc.272.41.25832&rft_id=info:pmid/9325313&rft.aulast=Pace&rft.aufirst=P&rft.au=Taylor,+J&rft.au=Suntharalingam,+S&rft.au=Coombes,+RC&rft.au=Ali,+S&rft_id=https://doi.org/10.1074%2Fjbc.272.41.25832&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Brandenberger AW, Tee MK, Lee JY, Chao V, Jaffe RB (October 1997). "Tissue distribution of estrogen receptors alpha (ER-alpha) and beta (ER-beta) mRNA in the midgestational human fetus". The Journal of Clinical Endocrinology and Metabolism. 82 (10): 3509–3512\. doi:10.1210/jcem.82.10.4400. PMID 9329394.3509-3512&rft.date=1997-10&rft_id=info:doi/10.1210/jcem.82.10.4400&rft_id=info:pmid/9329394&rft.aulast=Brandenberger&rft.aufirst=AW&rft.au=Tee,+MK&rft.au=Lee,+JY&rft.au=Chao,+V&rft.au=Jaffe,+RB&rft_id=https://doi.org/10.1210%2Fjcem.82.10.4400&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Enmark E, Pelto-Huikko M, Grandien K, Lagercrantz S, Lagercrantz J, Fried G, et al. (December 1997). "Human estrogen receptor beta-gene structure, chromosomal localization, and expression pattern". The Journal of Clinical Endocrinology and Metabolism. 82 (12): 4258–4265\. doi:10.1210/jcem.82.12.4470. PMID 9398750.4258-4265&rft.date=1997-12&rft_id=info:doi/10.1210/jcem.82.12.4470&rft_id=info:pmid/9398750&rft.aulast=Enmark&rft.aufirst=E&rft.au=Pelto-Huikko,+M&rft.au=Grandien,+K&rft.au=Lagercrantz,+S&rft.au=Lagercrantz,+J&rft.au=Fried,+G&rft.au=Nordenskjöld,+M&rft.au=Gustafsson,+JA&rft_id=https://doi.org/10.1210%2Fjcem.82.12.4470&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Vladusic EA, Hornby AE, Guerra-Vladusic FK, Lupu R (January 1998). "Expression of estrogen receptor beta messenger RNA variant in breast cancer". Cancer Research. 58 (2): 210–214\. PMID 9443393.210-214&rft.date=1998-01&rft_id=info:pmid/9443393&rft.aulast=Vladusic&rft.aufirst=EA&rft.au=Hornby,+AE&rft.au=Guerra-Vladusic,+FK&rft.au=Lupu,+R&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Ogawa S, Inoue S, Watanabe T, Hiroi H, Orimo A, Hosoi T, et al. (February 1998). "The complete primary structure of human estrogen receptor beta (hER beta) and its heterodimerization with ER alpha in vivo and in vitro". Biochemical and Biophysical Research Communications. 243 (1): 122–126\. doi:10.1006/bbrc.1997.7893. PMID 9473491.122-126&rft.date=1998-02&rft_id=info:doi/10.1006/bbrc.1997.7893&rft_id=info:pmid/9473491&rft.aulast=Ogawa&rft.aufirst=S&rft.au=Inoue,+S&rft.au=Watanabe,+T&rft.au=Hiroi,+H&rft.au=Orimo,+A&rft.au=Hosoi,+T&rft.au=Ouchi,+Y&rft.au=Muramatsu,+M&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Alves SE, Lopez V, McEwen BS, Weiland NG (March 1998). "Differential colocalization of estrogen receptor beta (ERbeta) with oxytocin and vasopressin in the paraventricular and supraoptic nuclei of the female rat brain: an immunocytochemical study". Proceedings of the National Academy of Sciences of the United States of America. 95 (6): 3281–3286\. Bibcode:1998PNAS...95.3281A. doi:10.1073/pnas.95.6.3281. PMC 19733. PMID 9501254.3281-3286&rft.date=1998-03&rft_id=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC19733#id-name=PMC&rft_id=info:pmid/9501254&rft_id=info:doi/10.1073/pnas.95.6.3281&rft_id=info:bibcode/1998PNAS...95.3281A&rft.aulast=Alves&rft.aufirst=SE&rft.au=Lopez,+V&rft.au=McEwen,+BS&rft.au=Weiland,+NG&rft_id=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC19733&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Brandenberger AW, Tee MK, Jaffe RB (March 1998). "Estrogen receptor alpha (ER-alpha) and beta (ER-beta) mRNAs in normal ovary, ovarian serous cystadenocarcinoma and ovarian cancer cell lines: down-regulation of ER-beta in neoplastic tissues". The Journal of Clinical Endocrinology and Metabolism. 83 (3): 1025–1028\. doi:10.1210/jcem.83.3.4788. PMID 9506768.1025-1028&rft.date=1998-03&rft_id=info:doi/10.1210/jcem.83.3.4788&rft_id=info:pmid/9506768&rft.aulast=Brandenberger&rft.aufirst=AW&rft.au=Tee,+MK&rft.au=Jaffe,+RB&rft_id=https://doi.org/10.1210%2Fjcem.83.3.4788&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Moore JT, McKee DD, Slentz-Kesler K, Moore LB, Jones SA, Horne EL, et al. (June 1998). "Cloning and characterization of human estrogen receptor beta isoforms". Biochemical and Biophysical Research Communications. 247 (1): 75–78\. doi:10.1006/bbrc.1998.8738. PMID 9636657.75-78&rft.date=1998-06&rft_id=info:doi/10.1006/bbrc.1998.8738&rft_id=info:pmid/9636657&rft.aulast=Moore&rft.aufirst=JT&rft.au=McKee,+DD&rft.au=Slentz-Kesler,+K&rft.au=Moore,+LB&rft.au=Jones,+SA&rft.au=Horne,+EL&rft.au=Su,+JL&rft.au=Kliewer,+SA&rft.au=Lehmann,+JM&rft.au=Willson,+TM&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Ogawa S, Inoue S, Watanabe T, Orimo A, Hosoi T, Ouchi Y, Muramatsu M (August 1998). "Molecular cloning and characterization of human estrogen receptor betacx: a potential inhibitor ofestrogen action in human". Nucleic Acids Research. 26 (15): 3505–3512\. doi:10.1093/nar/26.15.3505. PMC 147730. PMID 9671811.3505-3512&rft.date=1998-08&rft_id=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC147730#id-name=PMC&rft_id=info:pmid/9671811&rft_id=info:doi/10.1093/nar/26.15.3505&rft.aulast=Ogawa&rft.aufirst=S&rft.au=Inoue,+S&rft.au=Watanabe,+T&rft.au=Orimo,+A&rft.au=Hosoi,+T&rft.au=Ouchi,+Y&rft.au=Muramatsu,+M&rft_id=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC147730&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Lu B, Leygue E, Dotzlaw H, Murphy LJ, Murphy LC, Watson PH (March 1998). "Estrogen receptor-beta mRNA variants in human and murine tissues". Molecular and Cellular Endocrinology. 138 (1–2): 199–203\. doi:10.1016/S0303-7207(98)00050-1. PMID 9685228. S2CID 54243493.1–2&rft.pages=199-203&rft.date=1998-03&rft_id=https://api.semanticscholar.org/CorpusID:54243493#id-name=S2CID&rft_id=info:pmid/9685228&rft_id=info:doi/10.1016/S0303-7207(98)00050-1&rft.aulast=Lu&rft.aufirst=B&rft.au=Leygue,+E&rft.au=Dotzlaw,+H&rft.au=Murphy,+LJ&rft.au=Murphy,+LC&rft.au=Watson,+PH&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Seol W, Hanstein B, Brown M, Moore DD (October 1998). "Inhibition of estrogen receptor action by the orphan receptor SHP (short heterodimer partner)". Molecular Endocrinology. 12 (10): 1551–1557\. doi:10.1210/mend.12.10.0184. PMID 9773978.1551-1557&rft.date=1998-10&rft_id=info:doi/10.1210/mend.12.10.0184&rft_id=info:pmid/9773978&rft.aulast=Seol&rft.aufirst=W&rft.au=Hanstein,+B&rft.au=Brown,+M&rft.au=Moore,+DD&rft_id=https://doi.org/10.1210%2Fmend.12.10.0184&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Hanstein B, Liu H, Yancisin MC, Brown M (January 1999). "Functional analysis of a novel estrogen receptor-beta isoform". Molecular Endocrinology. 13 (1): 129–137\. doi:10.1210/mend.13.1.0234. PMID 9892018.129-137&rft.date=1999-01&rft_id=info:doi/10.1210/mend.13.1.0234&rft_id=info:pmid/9892018&rft.aulast=Hanstein&rft.aufirst=B&rft.au=Liu,+H&rft.au=Yancisin,+MC&rft.au=Brown,+M&rft_id=https://doi.org/10.1210%2Fmend.13.1.0234&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">
* Vidal O, Kindblom LG, Ohlsson C (June 1999). "Expression and localization of estrogen receptor-beta in murine and human bone". Journal of Bone and Mineral Research. 14 (6): 923–929\. doi:10.1359/jbmr.1999.14.6.923. PMID 10352100. S2CID 85604096.923-929&rft.date=1999-06&rft_id=https://api.semanticscholar.org/CorpusID:85604096#id-name=S2CID&rft_id=info:pmid/10352100&rft_id=info:doi/10.1359/jbmr.1999.14.6.923&rft.aulast=Vidal&rft.aufirst=O&rft.au=Kindblom,+LG&rft.au=Ohlsson,+C&rfr_id=info:sid/en.wikipedia.org:Estrogen+receptor+beta" class="Z3988">

## External links

[edit]

* Estrogen+Receptor+beta at the U.S. National Library of Medicine Medical Subject Headings (MeSH)
* Overview of all the structural information available in the PDB for UniProt: Q92731 (Estrogen receptor beta) at the PDBe-KB.

This article incorporates text from the United States National Library of Medicine, which is in the public domain.

* v
* t
* e

PDB gallery

* 1hj1: RAT ESTROGEN RECEPTOR BETA LIGAND-BINDING DOMAIN IN COMPLEX WITH PURE ANTIESTROGEN ICI164,384

* 1l2j: Human Estrogen Receptor beta Ligand-binding Domain in Complex with (R,R)-5,11-cis-diethyl-5,6,11,12-tetrahydrochrysene-2,8-diol

* 1nde: Estrogen Receptor beta with Selective Triazine Modulator

* 1qkm: HUMAN ESTROGEN RECEPTOR BETA LIGAND-BINDING DOMAIN IN COMPLEX WITH PARTIAL AGONIST GENISTEIN

* 1qkn: RAT ESTROGEN RECEPTOR BETA LIGAND-BINDING DOMAIN IN COMPLEX WITH ANTAGONIST RALOXIFENE

* 1u3q: Crystal Structure of Estrogen Receptor beta complexed with CL-272

* 1u3r: Crystal Structure of Estrogen Receptor beta complexed with WAY-338

* 1u3s: Crystal Structure of Estrogen Receptor beta complexed with WAY-797

* 1u9e: CRYSTAL STRUCTURE OF ESTROGEN RECEPTOR BETA COMPLEXED WITH WAY-397

* 1x76: CRYSTAL STRUCTURE OF ESTROGEN RECEPTOR BETA COMPLEXED WITH WAY-697

* 1x78: CRYSTAL STRUCTURE OF ESTROGEN RECEPTOR BETA COMPLEXED WITH WAY-244

* 1x7b: CRYSTAL STRUCTURE OF ESTROGEN RECEPTOR BETA COMPLEXED WITH ERB-041

* 1x7j: CRYSTAL STRUCTURE OF ESTROGEN RECEPTOR BETA COMPLEXED WITH GENISTEIN

* 1yy4: Crystal structure of estrogen receptor beta complexed with 1-chloro-6-(4-hydroxy-phenyl)-naphthalen-2-ol

* 1yye: Crystal structure of estrogen receptor beta complexed with way-202196

* 1zaf: Crystal structure of estrogen receptor beta complexed with 3-Bromo-6-hydroxy-2-(4-hydroxy-phenyl)-inden-1-one

* 2fsz: A second binding site for hydroxytamoxifen within the coactivator-binding groove of estrogen receptor beta

* 2giu: Human estrogen receptor beta ligand-binding domain in complex with compound 45

* 2i0g: Benzopyrans are Selective Estrogen Receptor beta Agonists (SERBAs) with Novel Activity in Models of Benign Prostatic Hyperplasia

* 2j7x: STRUCTURE OF ESTRADIOL-BOUND ESTROGEN RECEPTOR BETA LBD IN COMPLEX WITH LXXLL MOTIF FROM NCOA5

* 2j7y: STRUCTURE OF 17-EPIESTRIOL-BOUND ESTROGEN RECEPTOR BETA LBD IN COMPLEX WITH LXXLL MOTIF FROM NCOA5

* v
* t
* e

Transcription factors and intracellular receptors

(1) Basic domains

| (1.1) Basic leucine zipper (bZIP)|

* Activating transcription factor
* AATF
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* AP-1
* c-Fos
* FOSB
* FOSL1
* FOSL2
* JDP2
* c-Jun
* JUNB
* JunD
* BACH
* 1
* 2
* BATF
* BLZF1
* C/EBP
* α
* β
* γ
* δ
* ε
* ζ
* CREB
* 1
* 3
* L1
* CREM
* DBP
* DDIT3
* GABPA
* GCN4
* HLF
* MAF
* B
* sMaf
* F
* G
* K
* NFE
* 2
* L1
* L2
* L3
* NFIL3
* NRL
* NRF
* 1
* 2
* 3
* XBP1

|

(1.2) Basic helix-loop-helix (bHLH)|

| Group A|

* AS-C
* ASCL1
* ASCL2
* ATOH1
* HAND
* 1
* 2
* MESP2
* Myogenic regulatory factors
* MyoD
* Myogenin
* MYF5
* MYF6
* NeuroD
* 1
* 2
* Neurogenins
* 1
* 2
* 3
* OLIG
* 1
* 2
* Paraxis
* TCF15
* Scleraxis
* SLC
* LYL1
* TAL
* 1
* 2
* Twist

|

Group B|

* FIGLA
* Myc
* c-Myc
* l-Myc
* n-Myc
* MXD4
* TCF4

Group C
bHLH-PAS|

* AhR
* AHRR
* ARNT
* ARNTL
* ARNTL2
* CLOCK
* HIF
* 1A
* EPAS1
* 3A
* NPAS
* 1
* 2
* 3
* PER
* 1
* 2
* 3
* Period
* SIM
* 1
* 2

Group D|

* BHLH
* 2
* 3
* 9
* Pho4
* ID
* 1
* 2
* 3
* 4

Group E|

* HES
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* HEY
* 1
* 2
* L

Group F
bHLH-COE|

* EBF1

(1.3) bHLH-ZIP|

* AP-4
* MAX
* MXD1
* MXD3
* MITF
* MNT
* MLX
* MLXIPL
* MXI1
* Myc
* SREBP
* 1
* 2
* USF1

(1.4) NF-1|

* NFI
* A
* B
* C
* X
* SMAD
* R-SMAD
* 1
* 2
* 3
* 5
* 9
* I-SMAD
* 6
* 7
* 4)

(1.5) RF-X|

* RFX
* 1
* 2
* 3
* 4
* 5
* 6
* ANK

(1.6) Basic helix-span-helix (bHSH)|

* AP-2
* α
* β
* γ
* δ
* ε

(2) Zinc finger DNA-binding domains

| (2.1) Nuclear receptor (Cys4)|

| subfamily 1|

* Thyroid hormone
* α
* β
* CAR
* FXR
* LXR
* α
* β
* PPAR
* α
* β/δ
* γ
* PXR
* RAR
* α
* β
* γ
* ROR
* α
* β
* γ
* Rev-ErbA
* α
* β
* VDR

|

subfamily 2|

* COUP-TF
* (I
* II
* Ear-2
* HNF4
* α
* γ
* PNR
* RXR
* α
* β
* γ
* Testicular receptor
* 2
* 4
* TLX

subfamily 3|

* Steroid hormone
* Androgen
* Estrogen
* α
* β
* Glucocorticoid
* Mineralocorticoid
* Progesterone
* Estrogen related
* α
* β
* γ

subfamily 4|

* NUR
* NGFIB
* NOR1
* NURR1

subfamily 5|

* LRH-1
* SF1

subfamily 6|

* GCNF

subfamily 0|

* DAX1
* SHP

(2.2) Other Cys4|

* GATA
* 1
* 2
* 3
* 4
* 5
* 6
* MTA
* 1
* 2
* 3
* TRPS1

(2.3) Cys2His2|

* General transcription factors
* TFIIA
* TFIIB
* TFIID
* TFIIE
* 1
* 2
* TFIIF
* 1
* 2
* TFIIH
* 1
* 2
* 4
* 2I
* 3A
* 3C1
* 3C2

* ATBF1
* BCL
* 6
* 11A
* 11B
* CTCF
* E4F1
* EGR
* 1
* 2
* 3
* 4
* ERV3
* GFI1
* GLI family
* 1
* 2
* 3
* REST
* S1
* S2
* YY1
* HIC
* 1
* 2
* HIVEP
* 1
* 2
* 3
* IKZF
* 1
* 2
* 3
* ILF
* 2
* 3
* Sp/KLF family
* KLF
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* 9
* 10
* 11
* 12
* 13
* 14
* 15
* 17
* SP
* 1
* 2
* 4
* 7
* 8
* MTF1
* MYT1
* OSR1
* PRDM9
* SALL
* 1
* 2
* 3
* 4
* TSHZ3
* WT1
* Zbtb7
* 7A
* 7B
* ZBTB
* 11
* 16
* 17
* 20
* 21
* 32
* 33
* 40
* zinc finger
* 3
* 7
* 9
* 10
* 19
* 22
* 24
* 33B
* 34
* 35
* 41
* 43
* 44
* 51
* 74
* 143
* 146
* 148
* 165
* 202
* 217
* 219
* 238
* 239
* 259
* 267
* 268
* 281
* 300
* 318
* 330
* 346
* 350
* 365
* 366
* 384
* 423
* 451
* 452
* 471
* 593
* 638
* 644
* 649
* 655
* 804A

(2.4) Cys6|

* HIVEP1

(2.5) Alternating composition|

* AIRE
* DIDO1
* GRLF1
* ING
* 1
* 2
* 4
* JARID
* 1A
* 1B
* 1C
* 1D
* 2
* JMJD1B

(2.6) WRKY|

* WRKY

(3) Helix-turn-helix domains

| (3.1) Homeodomain|

| Antennapedia
ANTP class|

| protoHOX
Hox-like|

* ParaHox
* Gsx
* 1
* 2
* Xlox
* PDX1
* Cdx
* 1
* 2
* 4
* extended Hox: Evx1
* Evx2
* MEOX1
* MEOX2
* Homeobox
* A1
* A2
* A3
* A4
* A5
* A7
* A9
* A10
* A11
* A13
* B1
* B2
* B3
* B4
* B5
* B6
* B7
* B8
* B9
* B13
* C4
* C5
* C6
* C8
* C9
* C10
* C11
* C12
* C13
* D1
* D3
* D4
* D8
* D9
* D10
* D11
* D12
* D13
* GBX1
* GBX2
* MNX1

|

metaHOX
NK-like|

* BARHL1
* BARHL2
* BARX1
* BARX2
* BSX
* DBX
* 1
* 2
* DLX
* 1
* 2
* 3
* 4
* 5
* 6
* EMX
* 1
* 2
* EN
* 1
* 2
* HHEX
* HLX
* LBX1
* LBX2
* MSX
* 1
* 2
* NANOG
* NKX
* 2-1
* 2-2
* 2-3
* 2-5
* 3-1
* 3-2
* HMX1
* HMX2
* HMX3
* 6-1
* 6-2
* NOTO
* TLX1
* TLX2
* TLX3
* VAX1
* VAX2

other|

* ARX
* CRX
* CUTL1
* FHL
* 1
* 2
* 3
* HESX1
* HOPX
* LMX
* 1A
* 1B
* NOBOX
* TALE
* IRX
* 1
* 2
* 3
* 4
* 5
* 6
* MKX
* MEIS
* 1
* 2
* PBX
* 1
* 2
* 3
* PKNOX
* 1
* 2
* SIX
* 1
* 2
* 3
* 4
* 5
* PHF
* 1
* 3
* 6
* 8
* 10
* 16
* 17
* 20
* 21A
* POU domain
* PIT-1
* BRN-3: A
* B
* C
* Octamer transcription factor: 1
* 2
* 3/4
* 6
* 7
* 11
* SATB2
* ZEB
* 1
* 2

(3.2) Paired box|

* PAX
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* 9
* PRRX
* 1
* 2
* PROP1
* PHOX
* 2A
* 2B
* RAX
* SHOX
* SHOX2
* VSX1
* VSX2
* Bicoid
* GSC
* BICD2
* OTX
* 1
* 2
* PITX
* 1
* 2
* 3

(3.3) Fork head / winged helix|

* E2F
* 1
* 2
* 3
* 4
* 5
* FOX proteins
* A1
* A2
* A3
* B1
* B2
* C1
* C2
* D1
* D2
* D3
* D4
* D4L1
* D4L3
* D4L4
* D4L5
* D4L6
* E1
* E3
* F1
* F2
* G1
* H1
* I1
* I2
* I3
* J1
* J2
* J3
* K1
* K2
* L1
* L2
* M1
* N1
* N2
* N3
* N4
* O1
* O3
* O4
* O6
* P1
* P2
* P3
* P4
* Q1
* R1
* R2
* S1

(3.4) Heat shock factors|

* HSF
* 1
* 2
* 4

(3.5) Tryptophan clusters|

* ELF
* 2
* 4
* 5
* EHF
* ELK
* 1
* 3
* 4
* ERF
* ETS
* 1
* 2
* ERG
* SPIB
* ETV
* 1
* 4
* 5
* 6
* FLI1
* Interferon regulatory factors
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* MYB
* MYBL2

(3.6) TEA domain|

* transcriptional enhancer factor
* 1
* 2
* 3
* 4

(4) β-Scaffold factors with minor groove contacts

| (4.1) Rel homology region|

* NF-κB
* NFKB1
* NFKB2
* REL
* RELA
* RELB
* NFAT
* C1
* C2
* C3
* C4
* 5

|

(4.2) STAT|

* STAT
* 1
* 2
* 3
* 4
* 5
* 6

(4.3) p53-like|

* p53 p63 p73 family
* p53
* TP63
* p73
* TBX
* 1
* 2
* 3
* 5
* 19
* 21
* 22
* TBR1
* TBR2
* TFT
* MYRF

(4.4) MADS box|

* Mef2
* A
* B
* C
* D
* SRF

(4.6) TATA-binding proteins|

* TBP
* TBPL1

(4.7) High-mobility group|

* BBX
* HMGB
* 1
* 2
* 3
* 4
* HMGN
* 1
* 2
* 3
* 4
* HNF
* 1A
* 1B
* SOX
* 1
* 2
* 3
* 4
* 5
* 6
* 8
* 9
* 10
* 11
* 12
* 13
* 14
* 15
* 17
* 18
* 21
* SRY
* SSRP1
* TCF/LEF
* TCF
* 1
* 3
* 4
* LEF1
* TOX
* 1
* 2
* 3
* 4

(4.9) Grainyhead|

* TFCP2

(4.10) Cold-shock domain|

* CSDA
* YBX1

(4.11) Runt|

* CBF
* CBFA2T2
* CBFA2T3
* RUNX1
* RUNX2
* RUNX3
* RUNX1T1

(0) Other transcription factors

| (0.2) HMGI(Y)|

* HMGA
* 1
* 2
* HBP1

|

(0.3) Pocket domain|

* Rb
* RBL1
* RBL2

(0.5) AP-2/EREBP-related factors|

* Apetala 2
* EREBP
* B3

(0.6) Miscellaneous|

* ARID
* 1A
* 1B
* 2
* 3A
* 3B
* 4A
* CAP
* IFI
* 16
* 35
* MLL
* 2
* 3
* T1
* MNDA
* NFY
* A
* B
* C
* Rho/Sigma

see also transcription factor/coregulator deficiencies

* v
* t
* e

Estrogen receptor modulators

ERTooltip Estrogen receptor|

| Agonists|

* Steroidal: 2-Hydroxyestradiol
* 2-Hydroxyestrone
* 3-Methyl-19-methyleneandrosta-3,5-dien-17β-ol
* 3α-Androstanediol
* 3α,5α-Dihydrolevonorgestrel
* 3β,5α-Dihydrolevonorgestrel
* 3α-Hydroxytibolone
* 3β-Hydroxytibolone
* 3β-Androstanediol
* 4-Androstenediol
* 4-Androstenedione
* 4-Fluoroestradiol
* 4-Hydroxyestradiol
* 4-Hydroxyestrone
* 4-Methoxyestradiol
* 4-Methoxyestrone
* 5-Androstenediol
* 7-Oxo-DHEA
* 7α-Hydroxy-DHEA
* 7α-Methylestradiol
* 7β-Hydroxyepiandrosterone
* 8,9-Dehydroestradiol
* 8,9-Dehydroestrone
* 8β-VE2
* 10β,17β-Dihydroxyestra-1,4-dien-3-one (DHED)
* 11β-Chloromethylestradiol
* 11β-Methoxyestradiol
* 15α-Hydroxyestradiol
* 16-Ketoestradiol
* 16-Ketoestrone
* 16α-Fluoroestradiol
* 16α-Hydroxy-DHEA
* 16α-Hydroxyestrone
* 16α-Iodoestradiol
* 16α-LE2
* 16β-Hydroxyestrone
* 16β,17α-Epiestriol (16β-hydroxy-17α-estradiol)
* 17α-Estradiol (alfatradiol)
* 17α-Dihydroequilenin
* 17α-Dihydroequilin
* 17α-Epiestriol (16α-hydroxy-17α-estradiol)
* 17α-Ethynyl-3α-androstanediol
* 17α-Ethynyl-3β-androstanediol
* 17β-Dihydroequilenin
* 17β-Dihydroequilin
* 17β-Methyl-17α-dihydroequilenin
* Abiraterone
* Abiraterone acetate
* Alestramustine
* Almestrone
* Anabolic steroids (e.g., testosterone and esters, methyltestosterone, metandienone (methandrostenolone), nandrolone and esters, many others; via estrogenic metabolites)
* Atrimustine
* Bolandiol
* Bolandiol dipropionate
* Butolame
* Clomestrone
* Cloxestradiol
* Cloxestradiol acetate
* Conjugated estriol
* Conjugated estrogens
* Cyclodiol
* Cyclotriol
* DHEA
* DHEA-S
* ent-Estradiol
* Epiestriol (16β-epiestriol, 16β-hydroxy-17β-estradiol)
* Epimestrol
* Equilenin
* Equilin
* ERA-63 (ORG-37663)
* Esterified estrogens
* Estetrol
* Estradiol
* Estradiol esters
* Lipoidal estradiol
* Polyestradiol phosphate
* Estramustine
* Estramustine phosphate
* Estrapronicate
* Estrazinol
* Estriol
* Estriol esters
* Polyestriol phosphate
* Estrofurate
* Estrogenic substances
* Estromustine
* Estrone
* Estrone esters
* Estrone methyl ether
* Estropipate
* Etamestrol (eptamestrol)
* Ethinylandrostenediol
* Ethandrostate
* Ethinylestradiol
* Ethinylestradiol 3-benzoate
* Ethinylestradiol sulfonate
* Ethinylestriol
* Ethylestradiol
* Etynodiol
* Etynodiol diacetate
* Hexolame
* Hippulin
* Hydroxyestrone diacetate
* Lynestrenol
* Lynestrenol phenylpropionate
* Mestranol
* Methylestradiol
* Moxestrol
* Mytatrienediol
* Nilestriol
* Norethisterone
* Noretynodrel
* Orestrate
* Pentolame
* Prodiame
* Prolame
* Promestriene
* RU-16117
* Quinestradol
* Quinestrol
* Tibolone

* Nonsteroidal: (R,R)-THC
* (S,S)-THC
* 2,8-DHHHC
* β-LGND1
* β-LGND2 (GTx-878)
* AC-186
* Allenestrol
* Allenolic acid
* Benzestrol
* Bifluranol
* Bisdehydrodoisynolic acid
* Butestrol
* Carbestrol
* D-15414
* DCW234
* Diarylpropionitrile
* Dienestrol
* Dienestrol diacetate
* Diethylstilbestrol
* Diethylstilbestrol esters
* Dimestrol (dianisylhexene)
* Dimethylstilbestrol
* Doisynoestrol (fenocycline)
* Doisynolic acid
* Efavirenz
* Elacestrant
* ERB-196 (WAY-202196)
* Erteberel (SERBA-1, LY-500307)
* Estrobin (DBE)
* Fenestrel
* FERb 033
* Fosfestrol (diethylstilbestrol diphosphate)
* Furostilbestrol (diethylstilbestrol difuroate)
* GTx-758
* Hexestrol
* Hexestrol esters
* ICI-85966 (Stilbostat)
* M2613
* meso-Butestrol
* meso-Hexestrol
* Mestilbol
* Methallenestril
* Methestrol
* Methestrol dipropionate
* Paroxypropione
* Pentafluranol
* Phenestrol
* Prinaberel (ERB-041, WAY-202041)
* Propylpyrazoletriol
* Quadrosilan
* SC-3296
* SC-4289
* SERBA-2
* SKF-82,958
* Terfluranol
* Triphenylbromoethylene
* Triphenylchloroethylene
* Triphenyliodoethylene
* Triphenylmethylethylene (triphenylpropene)
* WAY-166818
* WAY-169916
* WAY-200070
* WAY-204688 (SIM-688)
* WAY-214156

* Unknown/unsorted: ERB-26
* ERA-45
* ERB-79
* ZK-283197

* Xenoestrogens: Anise-related (e.g., anethole, anol, dianethole, dianol, photoanethole)
* Chalconoids (e.g., isoliquiritigenin, phloretin, phlorizin (phloridzin), wedelolactone)
* Coumestans (e.g., coumestrol, psoralidin)
* Flavonoids (incl. 7,8-DHF, 8-prenylnaringenin, apigenin, baicalein, baicalin, biochanin A, calycosin, catechin, daidzein, daidzin, ECG, EGCG, epicatechin, equol, formononetin, glabrene, glabridin, genistein, genistin, glycitein, kaempferol, liquiritigenin, mirificin, myricetin, naringenin, penduletin, pinocembrin, prunetin, puerarin, quercetin, tectoridin, tectorigenin)
* Lavender oil
* Lignans (e.g., enterodiol, enterolactone, nyasol (cis-hinokiresinol))
* Metalloestrogens (e.g., cadmium)
* Pesticides (e.g., alternariol, dieldrin, endosulfan, fenarimol, HPTE, methiocarb, methoxychlor, triclocarban, triclosan)
* Phytosteroids (e.g., digitoxin (digitalis), diosgenin, guggulsterone)
* Phytosterols (e.g., β-sitosterol, campesterol, stigmasterol)
* Resorcylic acid lactones (e.g., zearalanone, α-zearalenol, β-zearalenol, zearalenone, zeranol (α-zearalanol), taleranol (teranol, β-zearalanol))
* Steroid-like (e.g., deoxymiroestrol, miroestrol)
* Stilbenoids (e.g., resveratrol, rhaponticin)
* Synthetic xenoestrogens (e.g., alkylphenols, bisphenols (e.g., BPA, BPF, BPS), DDT, parabens, PBBs, PHBA, phthalates, PCBs)
* Others (e.g., agnuside, rotundifuran)

|

Mixed
(SERMsTooltip Selective estrogen receptor modulators)|

* 2-Phenylbenzofuran
* 2-Phenylbenzothiophene
* 4'-Hydroxynorendoxifen
* 27-Hydroxycholesterol
* Acefluranol
* Acolbifene
* Afimoxifene
* Anordiol
* Anordrin
* Arzoxifene
* Bazedoxifene
* Brilanestrant
* Broparestrol
* Camizestrant
* Chlorotrianisene
* Clomifene
* Clomifenoxide
* CN-55945-27
* Cyclofenil
* D-15413
* Desmethylchlorotrianisene
* Droloxifene
* Enclomifene
* Endoxifen
* Etacstil (GW-5638, DPC-974)
* Ethamoxytriphetol (MER-25)
* Femarelle
* Fispemifene
* GW-7604
* ICI-55548
* Idoxifene
* Lasofoxifene
* Levormeloxifene
* LN-1643
* LN-2299
* LY-117018
* Menerba
* Miproxifene
* Miproxifene phosphate
* MRL-37
* Nafoxidine
* Nitromifene
* NNC 45-0095
* NNC 45-0320
* NNC 45-0781
* NNC 45-1506
* Ormeloxifene
* Ospemifene
* Panomifene
* Pipendoxifene
* Promensil
* Raloxifene
* Rimostil (P-081)
* Spironolactone
* SS1010
* Tamoxifen
* TAS-108 (SR-16234)
* Toremifene
* Trioxifene
* TZE-5323
* U-11555A
* U-11634
* Y-134
* Zindoxifene
* Zuclomifene

Antagonists|

* (R,R)-THC
* 7β-Hydroxy-DHEA
* Chloroindazole
* Cytestrol acetate
* EM-800 (SCH-57050)
* Epitiostanol
* ERA-90
* ERB-88
* Fulvestrant (ICI-182780)
* Glyceollins (I, II, III, IV)
* ICI-164384
* MDL-101906
* Mepitiostane
* Methylepitiostanol
* Methylpiperidinopyrazole
* MIBE
* Oxabicycloheptene sulfonate
* Phenytoin
* PHTPP
* Prochloraz
* RU-39411
* RU-58668
* SS1020
* TAS-108 (SR-16234)
* ZB716
* ZK-164015
* ZK-191703

* Coregulator-binding modulators: ERX-11

* Noncompetitive inhibitors: Trilostane

GPERTooltip G protein-coupled estrogen receptor|

| Agonists|

* 2-Methoxyestradiol
* 7β-Hydroxyepiandrosterone
* Afimoxifene (4-hydroxytamoxifen)
* Aldosterone
* Atrazine
* Bisphenol A
* Daidzein
* DDT (p,p'-DDT, o',p'-DDE)
* Diarylpropionitrile
* Equol
* Estradiol
* Ethinylestradiol
* Fulvestrant (ICI-182780)
* G-1
* Genistein
* GPER-L1
* GPER-L2
* Hydroxytyrosol
* Kepone
* Nicotinic acid
* Nicotinamide
* Nonylphenol
* Oleuropein
* PCBs (2,2',5'-PCB-4-OH)
* Propylpyrazoletriol
* Quercetin
* Raloxifene
* Resveratrol
* STX
* Tamoxifen
* Tectoridin

|

Antagonists|

* CCL18
* Estriol
* G-15
* G-36
* MIBE

Unknown|

* Diethylstilbestrol
* Zearalenone

See also
Receptor/signaling modulators
Estrogens and antiestrogens
Androgen receptor modulators
Progesterone receptor modulators
List of estrogens
*[v]: View this template
*[t]: Discuss this template
*[e]: Edit this template
*[ER]: Estrogen receptor
*[SERMs]: Selective estrogen receptor modulators
*[GPER]: G protein-coupled estrogen receptor