Sex hormone-binding globulin
SHBG | ||||||
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UniProt | ||||||
RefSeq (mRNA) |
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RefSeq (protein) |
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Location (UCSC) | Chr 17: 7.61 – 7.63 Mb | Chr 11: 69.51 – 69.51 Mb | ||||
PubMed search | [3] | [4] |
View/Edit Human | View/Edit Mouse |
Androgen-binding protein, Sex hormone-binding globulin | |||||||
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Identifiers | |||||||
Symbol | SHBG | ||||||
Alt. symbols | ABP | ||||||
Chr. 17 p13-p12 | |||||||
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Sex hormone-binding globulin (SHBG) or sex steroid-binding globulin (SSBG) is a
Other
Function
Testosterone and estradiol circulate in the bloodstream, loosely bound mostly to
The relative binding affinity of various sex steroids for SHBG is dihydrotestosterone (DHT) > testosterone > androstenediol > estradiol > estrone.[8] DHT binds to SHBG with about 5 times the affinity of testosterone and about 20 times the affinity of estradiol.[9] Dehydroepiandrosterone (DHEA) is weakly bound to SHBG, but dehydroepiandrosterone sulfate is not bound to SHBG.[8] Androstenedione is not bound to SHBG either, and is instead bound solely to albumin.[10] Estrone sulfate and estriol are also poorly bound by SHBG.[11] Less than 1% of progesterone is bound to SHBG.[12]
SHBG levels are usually about twice as high in women than in men.[9] In women, SHBG serves to limit exposure to both androgens and estrogens.[9] Low SHBG levels in women have been associated with hyperandrogenism and endometrial cancer due to heightened exposure to androgens and estrogens, respectively.[9] During pregnancy, due to activation of SHBG production in the liver by high estrogen levels, SHBG levels increase by five-fold to ten-fold.[9] The high SHBG levels during pregnancy may serve to protect the mother from exposure to fetal androgens that escape metabolism by the placenta.[9] A case report of severe hyperandrogenism in a pregnant woman due to a rare instance of genetic SHBG deficiency illustrates this.[9][13]
Biochemistry
Biosynthesis
SHBG is produced mostly by the
Gene
The
Polymorphisms
There are variations in the genetic material for this protein that have different effects. In humans common polymorphisms include the following:
Rs6259, also called Asp327Asn location 7633209 on Chromosome 17, results in there being an extra N-glycosilation site, and so an extra sugar can be attached. This results in a longer circulation half-life for the protein, and raised levels. A health effect is a lowered risk of
Rs6258 also called Ser156Pro is at position 7631360 on the Chromosome 17.
Rs727428 position 7634474 is in several percent of humans.[19]
(TAAAA)(n) is five base pairs that repeats a variable number of times on the opposite DNA strand.[20]
Promoter activation
The mechanism of activating the promoter for SHBG in the liver involves
Protein
Sex hormone-binding globulin is homodimeric, meaning it has two identical peptide chains making up its structure. The amino acid sequence is the same as for
SHBG has two laminin G-like domains which form pockets that bind hydrophobic molecules. The steroids are bound by the LG domain at the amino end of the protein.[7] Inside the pocket of the domain is a serine residue that attracts the two different types of steroids at different points, thus changing their orientation. Androgens bind at the C3 functional groups on the A ring, and estrogens bind via a hydroxyl attached to C17 on the D ring. The two different orientations change a loop over the entrance to the pocket and the position of trp84 (in humans). Thus the whole protein signals what hormone it carries on its own surface.[7] The steroid binding LG domain is coded by exons 2 to 5.[7] A linker region joins the two LG domains together.[7]
When first produced the SHBG precursor has a leading signal peptide attached with 29 amino acids. The remaining peptide has 373 amino acids.[21] There are two sulfur bridges.
The sugars are attached at two different
Metals
A calcium ion is needed to link the two elements of the dimer together. Also a zinc ion is used to orient an otherwise disorganised part of the peptide chain.[7]
Regulation
SHBG has both enhancing and inhibiting hormonal influences thus can be viewed as a
In an effort to explain obesity-related reductions in SHBG, recent evidence suggests sugar or monosaccharide-induced
Blood values
Reference ranges for blood tests for SHBG have been developed:[27][28]
Population | Range |
---|---|
Adult female, premenopausal | 40–120 nmol/L |
Adult female, postmenopausal | 28–112 nmol/L |
Adult male | 20–60 nmol/L |
Infant (1–23 months) | 60–252 nmol/L |
Prepubertal (2 years–8 years) | 72–220 nmol/L |
Pubertal female | 36–125 nmol/L |
Pubertal male | 16–100 nmol/L |
Clinical significance
High or low levels
SHBG levels are decreased by androgens, administration of
In the womb the human fetus has a low level of SHBG allowing increased activity of sex hormones. After birth, the SHBG level rises and remains at a high level throughout childhood. At puberty the SHBG level halves in girls and goes down to a quarter in boys.[7] The change at puberty is triggered by growth hormone, and its pulsatility differs in boys and girls.[clarification needed] In pregnant women in the third trimester of pregnancy the SHBG level escalates to five to ten times the usual level for a woman.[7][9] A hypothesis is that this protects against the effect of hormone produced by the fetus.[7]
Obese girls are more likely to have an early menarche due to lower levels of SHBG.[7] Anorexia or a lean physique in women leads to higher SHBG levels, which in turn can lead to amenorrhea.[7]
Type 2 diabetes
Reduced levels of SHBG and also certain polymorphisms of the SHBG gene are implicated in the development of insulin resistance and type 2 diabetes.[36] Such effects apparently involve direct action at the cellular level where it became apparent that cell membranes of certain tissues contain specific high-affinity SHBG receptors.[37]
Coagulation
SHBG is a useful correlate and indirect marker of estrogen-induced procoagulation and by extension
Medications
Some medications, such as certain
Compound | Structure | SHBG RBA (%) |
SHBG K (106 M−1) |
CBG RBA (%) |
CBG K (106 M−1) |
---|---|---|---|---|---|
Aminoglutethimide | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Androstanolone | Steroidal | 220 | 5500 | 1.3 | 0.83 |
Betamethasone | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Cholecalciferol | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Cimetidine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Clomifene | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Cortisol (hydrocortisone) | Steroidal | 0.13 | 1.6 | 100 | 76 |
Cortisone acetate | Steroidal | 0.10 | 1.2 | <0.1 | <0.1 |
Cyproterone acetate | Steroidal | 0.10 | 1.2 | <0.1 | <0.1 |
Danazol | Steroidal | 18 | 240 | 10 | 6.5 |
Dexamethasone | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Diazoxide | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Diethylstilbestrol | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Digitoxin | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Digoxin | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
DL-DOPA | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Dopamine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Enclomiphene |
Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Epinephrine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Estradiol | Steroidal | 49 | 680 | <0.1 | <0.1 |
Estradiol benzoate | Steroidal | 0.70 | 8.6 | <0.1 | <0.1 |
Ethinylestradiol | Steroidal | 0.80 | 9.9 | <0.1 | <0.1 |
Ethisterone | Steroidal | 55 | 780 | 0.33 | 0.21 |
Fludrocortisone | Steroidal | <0.01 | <0.2 | 0.74 | 0.47 |
Fluoxymesterone | Steroidal | 4.8 | 60 | <0.1 | <0.1 |
Flutamide | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Homovanillic acid | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Hydrocortisone hemisuccinate | Steroidal | <0.01 | <0.2 | 8.7 | 5.6 |
Indometacin | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Levonorgestrel | Steroidal | 31 | 420 | <0.1 | <0.1 |
Medroxyprogesterone | Steroidal | 0.15 | 1.9 | 13 | 8.1 |
Medroxyprogesterone acetate | Steroidal | 0.08 | 1.0 | 6.5 | 4.2 |
Melatonin | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Mesterolone | Steroidal | 180 | 3600 | <0.1 | <0.1 |
Mestranol | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Methoxytryptophol |
Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Methyldopa | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Methylserotonin | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Methyltestosterone | Steroidal | 39 | 530 | <0.1 | <0.1 |
Metiamide | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Metribolone | Steroidal | 1.7 | 21 | 0.36 | 0.23 |
Metyrapone | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Mexrenone | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Nafoxidine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Nandrolone | Steroidal | 5.8 | 72 | 0.10 | 0.63 |
Norepinephrine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Norethisterone | Steroidal | 11 | 140 | 0.28 | 0.18 |
Noretynodrel | Steroidal | 1.3 | 16 | 0.16 | 0.10 |
Normetanephrine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Phenytoin | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Potassium canrenoate | Steroidal | 0.18 | 2.2 | 0.83 | 0.53 |
Prednisolone | Steroidal | 0.04 | 0.49 | 59 | 41 |
Prednisone | Steroidal | 0.17 | 2.1 | 5.0 | 3.2 |
Progesterone | Steroidal | 0.71 | 8.8 | 36 | 24 |
Promegestone | Steroidal | 0.007 | 0.09 | 0.40 | 0.25 |
Prorenone | Steroidal | 8.2 | 100 | <0.1 | <0.1 |
Reserpine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Rifampin |
Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Serotonin | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Spironolactone | Steroidal | 0.03 | 0.37 | <0.1 | <0.1 |
Tamoxifen | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Testolactone | Steroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Testosterone | Steroidal | 100 | 1600 | 8.3 | 5.3 |
Testosterone enanthate | Steroidal | 0.007 | 0.086 | <0.1 | <0.1 |
7α-Thioprogesterone | Steroidal | 0.06 | 0.74 | 36 | 24 |
7α-Thiospironolactone | Steroidal | 0.59 | 7.3 | <0.1 | <0.1 |
Thyroxine |
Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Triiodothyronine | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Trimethyltrienolone | Steroidal | 0.90 | 11 | 0.11 | 0.07 |
Vanillylmandelic acid | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
Zuclomifene | Nonsteroidal | <0.01 | <0.2 | <0.1 | <0.1 |
The reference CBG .
|
Progestogen | SHBG (%) | CBG (%) |
---|---|---|
17α-Allyl-19-nortestosterone | <1 | ? |
Allylestrenol | <1 | ? |
Chlormadinone acetate | <1 | <1 |
Cyproterone acetate | <1 | <1 |
Desogestrel | <1 | <1 |
Dienogest | <1 | <1 |
Drospirenone | <1 | <1 |
Etonogestrel | 15 | <1 |
Gestodene | 40 | <1 |
Levonorgestrel | 50 | <1 |
Medroxyprogesterone acetate | <1 | <1 |
Megestrol acetate | <1 | <1 |
Nomegestrol acetate | <1 | <1 |
Norelgestromin | <1 | ? |
Norethisterone | 16 | <1 |
Noretynodrel | <1 | <1 |
Norgestimate | <1 | <1 |
Progesterone | <1 | 36 |
Promegestone | <1 | <1 |
Segesterone acetate | <1 | ? |
Δ4-Tibolone | 1 | <1 |
Values are CBGcortisol .
was |
Compound | SHBG (%) |
---|---|
5α-Androstane-3β,17β-diol |
17 |
5β-Androstane-3α,17β-diol |
5 |
Dihydrotestosterone | 100 |
Ethylestrenol | <1 |
Fluoxymesterone | <1 |
Mesterolone | 440 |
Metandienone | 2 |
Metenolone | 3 |
Methyltestosterone | 5 |
Metribolone | <1 |
Nandrolone | 1 |
Oxymetholone | <1 |
Stanozolol | 1 |
Testosterone | 19 |
Values are RBAsligand (100%) for SHBG was dihydrotestosterone .
(%). The reference |
Compound | SHBG (%) |
---|---|
3β-Androstanediol | 100 |
Androstenediol | 77 |
Bolandiol | 24 |
Dihydroethisterone | 100 |
Dihydroethyltestosterone | 18–21 |
Dihydromethylandrostenediol | 77 |
Dihydronandrolone | 44 |
Dihydrotestosterone | 100 |
Dihydrotrestolone | 47 |
4,17α-Dimethyltestosterone | 97 |
Drostanolone | 39 |
Ethisterone | 92 |
Fluoxymesterone | 3 |
11-Ketodihydrotestosterone | 0 |
Medroxyprogesterone acetate | 16 |
Megestrol acetate | 0 |
Mestanolone | 84 |
Methasterone | 58 |
Methyl-1-testosterone | 69 |
Methylandrostenediol |
40 |
Methyltestosterone | 64 |
Mibolerone | 6 |
Nandrolone | 16 |
Nandrolone decanoate | 0 |
Nandrolone phenylpropionate | 0 |
Norethandrolone | 3 |
Norethisterone | 21 |
Normethandrone | 7 |
Oxandrolone | 0 |
Oxymetholone | 3 |
Progesterone | 13 |
Stanozolol | 36 |
1-Testosterone | 98 |
Testosterone | 82 |
Testosterone benzoate | 8 |
Testosterone cypionate | 6 |
Testosterone enanthate | 9 |
Δ4-Tibolone | 8 |
Trestolone | 12 |
Trestolone enanthate | 12 |
Vinyltestosterone | 36 |
Values are RBAsligand (100%) for SHBG was dihydrotestosterone .
(%). The reference |
Compound | SHBG (%) | CBG (%)
| ||||||
---|---|---|---|---|---|---|---|---|
Aldosterone | <0.2 | 6.0 | ||||||
Corticosterone | <0.2 | 107 | ||||||
Cortisol | <0.2 | 100 | ||||||
Dexamethasone | <0.2 | <0.1 | ||||||
Dihydrotestosterone | 100 | 0.8 | ||||||
Estradiol | 8.7 | <0.1 | ||||||
Metribolone | 0.2 | <0.1 | ||||||
Moxestrol | <0.2 | <0.1 | ||||||
Progesterone | <0.2 | 25 | ||||||
Promegestone | <0.2 | 0.9 | ||||||
Testosterone | 26 | 3 | ||||||
Values are CBGcortisol .
was |
Compound | RBA to (%)SHBG |
Bound to SHBG (%) |
Bound to albumin (%) |
---|---|---|---|
17β-Estradiol | 50 | 37 | 61 |
Estrone | 12 | 16 | 80 |
Estriol | 0.3 | 1 | 91 |
Estrone sulfate | 0 | 0 | 99 |
17β-Dihydroequilin | 30 | ? | ? |
Equilin | 8 | 26 | 13 |
17β-Dihydroequilin sulfate |
0 | ? | ? |
Equilin sulfate |
0 | ? | ? |
Δ8-Estrone | ? | ? | ? |
The reference RBAtestosterone .
(%) values was |
Endogenous steroids
Measurement
When checking serum estradiol or testosterone, a total level that includes free and bound fractions can be assayed, or the free portion may be measured alone. Sex hormone-binding globulin can be measured separately from the total fraction of testosterone.
A free androgen index expresses the ratio of testosterone to SHBG and can be used to summarize the activity of free testosterone.
Affinity and binding
Steroid | SHBG affinity | Plasma protein binding in men | Plasma protein binding in women (follicular phase) | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
RBA (%) | K (106 M−1) | Total (nM) | Unbound (%) | SHBG (%) | CBG (%) | Albumin (%) | Total (nM) | Unbound (%) | SHBG (%) | CBG (%) | Albumin (%) | |
Aldosterone | 0.017 | 0.21 | 0.35 | 37.1 | 0.10 | 21.2 | 41.6 | 0.24 | 36.8 | 0.23 | 21.9 | 41.2 |
3α-Androstanediol | 82 | 1300 | 0.41 | 0.85 | 13.7 | <0.1 | 85.5 | 0.068 | 0.71 | 27.9 | <0.1 | 71.4 |
Androstenediol | 97 | 1500 | 4.3 | 3.24 | 60.4 | <0.1 | 36.3 | 2.4 | 1.73 | 78.8 | <0.1 | 19.4 |
Androstenedione | 2.3 | 29 | 4.1 | 7.85 | 2.82 | 1.37 | 88.0 | 5.4 | 7.54 | 6.63 | 1.37 | 84.5 |
Androsterone | 1.1 | 14 | 2.0 | 4.22 | 0.73 | 0.52 | 94.5 | 1.5 | 4.18 | 1.77 | 0.54 | 93.5 |
Corticosterone | 0.18 | 2.2 | 12 | 3.39 | 0.09 | 77.5 | 19.0 | 7.0 | 3.28 | 0.22 | 78.1 | 18.4 |
Cortisol | 0.13 | 1.6 | 400 | 3.91 | 0.08 | 89.5 | 6.57 | 400 | 3.77 | 0.18 | 89.7 | 6.33 |
Cortisone | 0.22 | 2.7 | 72 | 16.2 | 0.54 | 38.0 | 45.3 | 54 | 15.8 | 1.30 | 38.6 | 44.3 |
Dehydroepiandrosterone | 5.3 | 66 | 24 | 4.13 | 3.38 | <0.1 | 92.4 | 17 | 3.93 | 7.88 | <0.1 | 88.1 |
11-Deoxycorticosterone | 1.9 | 24 | 0.20 | 2.69 | 0.80 | 36.4 | 60.1 | 0.12 | 2.62 | 1.91 | 36.9 | 58.6 |
11-Deoxycortisol | 1.3 | 16 | 1.4 | 3.37 | 0.67 | 77.1 | 18.9 | 0.60 | 3.24 | 1.57 | 77.1 | 18.1 |
Dihydrotestosterone | 220 | 5500 | 1.7 | 0.88 | 59.7 | 0.22 | 39.2 | 0.65 | 0.47 | 78.4 | 0.12 | 21.0 |
Estradiol | 49 | 680 | 0.084 | 2.32 | 19.6 | <0.1 | 78.0 | 0.29 | 1.81 | 37.3 | <0.1 | 60.8 |
Estriol | 0.35 | 4.3 | 0.037 | 8.15 | 0.44 | <0.2 | 91.3 | 0.10 | 8.10 | 1.06 | <0.2 | 90.7 |
Estrone | 12 | 150 | 0.081 | 3.96 | 7.37 | <0.1 | 88.6 | 0.23 | 3.58 | 16.3 | <0.1 | 80.1 |
Etiocholanolone | 0.11 | 1.4 | 1.3 | 8.15 | 0.14 | 0.44 | 91.3 | 1.2 | 8.13 | 0.35 | 0.46 | 91.1 |
Pregnenolone | 1.1 | 14 | 2.4 | 2.87 | 0.50 | 0.16 | 96.5 | 2.2 | 2.85 | 1.21 | 0.16 | 95.8 |
17α-Hydroxypregnenolone | 0.19 | 2.3 | 5.4 | 4.27 | 0.12 | <0.1 | 95.5 | 3.5 | 4.26 | 0.30 | <0.1 | 95.4 |
Progesterone | 0.71 | 8.8 | 0.57 | 2.39 | 0.26 | 17.2 | 80.1 | 0.65 | 2.36 | 0.63 | 17.7 | 79.3 |
17α-Hydroxyprogesterone | 0.8 | 9.9 | 5.4 | 2.50 | 0.31 | 41.3 | 55.9 | 1.8 | 2.44 | 0.73 | 42.1 | 54.7 |
Testosterone | 100 | 1600 | 23 | 2.23 | 44.3 | 3.56 | 49.9 | 1.3 | 1.36 | 66.0 | 2.26 | 30.4 |
In men, the concentrations of SHBG, CBG, and albumin were 28 nM, 0.7 μM, and 0.56 mM, respectively. In women, the concentrations of SHBG, CBG, and albumin were 37 nM, 0.7 μM, and 0.56 mM, respectively. |
Synonyms
SHBG has been known under a variety of different names including:[52][53][54]
- Sex hormone-binding globulin (SHBG)
- Sex steroid-binding globulin (SSBG, SBG)
- Sex steroid-binding protein (SBP, SSBP)
- Androgen-binding protein (ABP)
- Estradiol-binding-protein (EBP)
- Testosterone–estradiol binding globulin (TeBG, TEBG)
References
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- ^ a b c GRCm38: Ensembl release 89: ENSMUSG00000005202 – Ensembl, May 2017
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- ^ Unit Code 91215 Archived 2011-07-20 at the Wayback Machine at Mayo Clinic Medical Laboratories. Retrieved April 2011
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Further reading
- Rosner W, Hryb DJ, Khan MS, Nakhla AM, Romas NA (1999). "Sex hormone-binding globulin mediates steroid hormone signal transduction at the plasma membrane". The Journal of Steroid Biochemistry and Molecular Biology. 69 (1–6): 481–5. S2CID 6499033.
- Power SG, Bocchinfuso WP, Pallesen M, Warmels-Rodenhiser S, Van Baelen H, Hammond GL (October 1992). "Molecular analyses of a human sex hormone-binding globulin variant: evidence for an additional carbohydrate chain". The Journal of Clinical Endocrinology and Metabolism. 75 (4): 1066–70. PMID 1400872.
- Gershagen S, Lundwall A, Fernlund P (November 1989). "Characterization of the human sex hormone binding globulin (SHBG) gene and demonstration of two transcripts in both liver and testis". Nucleic Acids Research. 17 (22): 9245–58. PMID 2587256.
- Hammond GL, Underhill DA, Rykse HM, Smith CL (November 1989). "The human sex hormone-binding globulin gene contains exons for androgen-binding protein and two other testicular messenger RNAs". Molecular Endocrinology. 3 (11): 1869–76. PMID 2608061.
- Que BG, Petra PH (July 1987). "Characterization of a cDNA coding for sex steroid-binding protein of human plasma". FEBS Letters. 219 (2): 405–9. S2CID 21654093.
- Gershagen S, Fernlund P, Lundwall A (August 1987). "A cDNA coding for human sex hormone binding globulin. Homology to vitamin K-dependent protein S". FEBS Letters. 220 (1): 129–35. S2CID 13166151.
- Walsh KA, Titani K, Takio K, Kumar S, Hayes R, Petra PH (November 1986). "Amino acid sequence of the sex steroid binding protein of human blood plasma". Biochemistry. 25 (23): 7584–90. PMID 3542030.
- Hammond GL, Robinson PA, Sugino H, Ward DN, Finne J (April 1986). "Physicochemical characteristics of human sex hormone binding globulin: evidence for two identical subunits". Journal of Steroid Biochemistry. 24 (4): 815–24. PMID 3702459.
- Hardy DO, Cariño C, Catterall JF, Larrea F (April 1995). "Molecular characterization of a genetic variant of the steroid hormone-binding globulin gene in heterozygous subjects". The Journal of Clinical Endocrinology and Metabolism. 80 (4): 1253–6. PMID 7714097.
- Cargill M, Altshuler D, Ireland J, Sklar P, Ardlie K, Patil N, Shaw N, Lane CR, Lim EP, Kalyanaraman N, Nemesh J, Ziaugra L, Friedland L, Rolfe A, Warrington J, Lipshutz R, Daley GQ, Lander ES (July 1999). "Characterization of single-nucleotide polymorphisms in coding regions of human genes". Nature Genetics. 22 (3): 231–8. S2CID 195213008.
- Grishkovskaya I, Avvakumov GV, Sklenar G, Dales D, Hammond GL, Muller YA (February 2000). "Crystal structure of human sex hormone-binding globulin: steroid transport by a laminin G-like domain". The EMBO Journal. 19 (4): 504–12. PMID 10675319.
- Hogeveen KN, Talikka M, Hammond GL (September 2001). "Human sex hormone-binding globulin promoter activity is influenced by a (TAAAA)n repeat element within an Alu sequence". The Journal of Biological Chemistry. 276 (39): 36383–90. PMID 11473114.
- Hryb DJ, Nakhla AM, Kahn SM, St George J, Levy NC, Romas NA, Rosner W (July 2002). "Sex hormone-binding globulin in the human prostate is locally synthesized and may act as an autocrine/paracrine effector". The Journal of Biological Chemistry. 277 (29): 26618–22. PMID 12015315.
- Raineri M, Catalano MG, Hammond GL, Avvakumov GV, Frairia R, Fortunati N (March 2002). "O-Glycosylation of human sex hormone-binding globulin is essential for inhibition of estradiol-induced MCF-7 breast cancer cell proliferation". Molecular and Cellular Endocrinology. 189 (1–2): 135–43. S2CID 24123789.
- Grishkovskaya I, Avvakumov GV, Hammond GL, Muller YA (May 2002). "Resolution of a disordered region at the entrance of the human sex hormone-binding globulin steroid-binding site". Journal of Molecular Biology. 318 (3): 621–6. PMID 12054810.
- Thompson DJ, Healey CS, Baynes C, Kalmyrzaev B, Ahmed S, Dowsett M, Folkerd E, Luben RN, Cox D, Ballinger D, Pharoah PD, Ponder BA, Dunning AM, Easton DF (December 2008). "Identification of common variants in the SHBG gene affecting sex hormone-binding globulin levels and breast cancer risk in postmenopausal women". Cancer Epidemiology, Biomarkers & Prevention. 17 (12): 3490–8. PMID 19064566.
- Trkiehl (2011). "SHBG - SNPedia". Retrieved 13 July 2014.
External links
- Overview of all the structural information available in the PDB for UniProt: P04278 (Sex hormone-binding globulin) at the PDBe-KB.
- Androgen-Binding+Protein at the U.S. National Library of Medicine Medical Subject Headings (MeSH)