Hämoglobinprotein (Maus)

Insight into the evolution of hemoglobins and their genes is a rich source of ideas about the function of hemoglobin and the regulation of globin gene expression. This article introduces the numerous genes and gene families encoding human globins, summarizes important events in the evolution of hemoglobin gene clusters, and discusses how these studies provide insight into the regulation of globin genes. Although the genes in and around the α-like globin gene complex are relatively stable, the β-like globin gene clusters are more dynamic, showing evidence of transposition to a new locus and frequent lineage-specific extensions and deletions.

 Das Cis-Regulatorische Module, die das Ausmaß und den Zeitpunkt der Genexpression steuern, sind eine Mischung aus konservierter und abstammungsspezifischer DNA, was möglicherweise die evolutionäre Beschränkung auf zentrale regulatorische Funktionen widerspiegelt, die bei Säugetieren weit verbreitet sind, und eine adaptive Feinabstimmung in verschiedenen Ordnungen von Säugetieren.

A wide range of animals, including vertebrates and invertebrates, use hemoglobins to transport oxygen, taking it from the lungs, gills, or other respiratory organs to peripheral tissues that need the oxygen for efficient metabolism. Therefore, it is natural to compare the structure and function of hemoglobin proteins between species, both to explore adaptation and to discover aspects of biochemistry and physiology that are conserved. Comparative studies have also been performed on the genes and gene clusters encoding the hemoglobins, revealing a rich history of gene duplication and deletion, as well as translocation. One motivation for comparative studies was to use the insights from the evolutionary analyses, to better understand the mechanisms of gene regulation. Many human hemoglobinopathies result from under-expression of globin genes, and attempts to modulate globin gene expression are a fundamental approach to finding new therapeutic options. This article summarizes general aspects of the evolution of hemoglobins and related globins, beginning with thefive basic types of globins found in humans, and then progressively focuses more specifically on hemoglobin gene clusters in vertebrates and eutherous mammals. Finally, the influence of this and related studies on the gene regulation of hemoglobin is discussed.

HEMOGLOBINS WITHIN THE GLOBIN-HEM FAMILY

Hemoglobins were originally discovered as abundant proteins in red blood cells of mammals and other vertebrates with jaws (gnathostomes) that reversibly bind and release oxygen. The major hemoglobin in adult humans, hemoglobin A, is a heterotetramer composed of two α-globin and two β-globin polypeptides, each with an associated heme group. These are encoded by the duplicated  HBA1  and  HBA2  genes or by the  HBB  gene.

  • Hemoglobins are only produced in erythroid cells, where they are the main protein. Composed of multiple subunits, hemoglobin cooperatively binds oxygen in the lungs, transports the oxygen through the blood, and releases it in peripheral tissues to support oxidative metabolism.
  • The heme molecule contains an iron atom (Fe) in the reduced state (iron or +2 oxidation state) and five of the six coordination sites of Fe are occupied, four by the porphyrin ring and one by the „proximal“ histidine (His) of the surrounding globin polypeptide . The sixth coordination site is reversibly bound by oxygen. Oxygen is loaded, transported, and unloaded onto hemoglobin without chemical (covalent) change to either the oxygen or the heme group, and the iron in the heme group remains in the reduced state.
  • This is in contrast to the known role of many heme proteins, myoglobin (encoded by the  MB  gene) is a related, monomeric heme-linked globin protein found predominantly in skeletal and cardiac muscle. The Fe in heme is five-coordinate. It has long been described as an oxygen storage protein and facilitates the diffusion of oxygen to the mitochondria (  Wittenberg and Wittenberg 1987 ).
  • Additional heme-containing globins were discovered by searching the wealth of information contained in the sequence of the human genome and transcripts made from it. Cytoglobin encoded by the  CYGB  gene  is found in many tissues (  Burmester et al. 2002  ;  Trent and Hargrove 2002  ), in sharp contrast to the strictly tissue-specific expression patterns of hemoglobin and myoglobin genes. The most distantly related globin found in the human genome is neuroglobin, which   is encoded  by NGB ( Burmester et al. 2000  ).
  • Its mRNA is abundant in brain tissue but is also present in many other tissues. It is related to invertebrate nerve globins , indicating that an ancestral gene was present more than 800 million years ago, before the divergence of vertebrates and invertebrates. In contrast to the five-coordinate heme complex in hemoglobins and MB, heme forms a six-coordinate complex with both NGB and CYGB, with two His residues, designated proximal and distal, coordinated to Fe. Ligands such as oxygen and nitric oxide compete with the distal His for binding, but despite this, NGB and CYBG still have high affinity for the ligands.
  • These two hexacoordinate heme-globins have been implicated in nitric oxide metabolism, with CYGB showing nitric oxide dioxygenase activity that converts nitric oxide to nitrate (  Oleksiewicz et al. 2011  ), and NGB showing nitrite reductase activity to form Nitric oxide shows (  Tiso et al. 2011  ). ). The latter activity has also been found for myoglobin (  Hendgen-Cotta et al. 2008  ) and deoxy-hemoglobin ( Gladwin and Kim-Shapiro 2008 ). Physiologically, nitrite reductase activity could provide a means of producing nitric oxide under hypoxic conditions, the signaling of which could regulate mitochondrial respiration and protect tissues (nerves by NGB, heart muscle by MB) from damage under ischemic conditions (  Dietz 2011  ).
  • A role for CYGB in oxygen-requiring reactions such as hydroxylation has not been ruled out. These proposed enzymatic roles in nitric oxide and other metabolism may derive from functions performed by ancestral hemoglobins in primordial life.

The five types of globin genes listed in Figure 1 are located on five different chromosomes:  HBA1  and  HBA2  at chromosome position 16p13.3,  HBB  at 11p15.4,  MB  at 22q12.3,  CYGB  at 17q25.1 and  NGB  at 14q24.3. 

MB  ,  CYGB  and  NGB  are present as single copy genes while  HBB  and  HBA  s are present in clusters with multiple related genes. All genes consist of at least three exons separated by two introns. Although the introns differ dramatically in size, they are in homologous locations. CYGB  and  NGB  _genes each have an extra exon. It has been suggested that preservation of intron position in vertebrate globin genes facilitates exon shuffling during protein evolution. However, intron positions in globin genes differ considerably outside of vertebrates, suggesting that the conservation of intron position may simply reflect an ancestral state that has not changed during vertebrate evolution.

Hemoglobin protein (Mouse)

30R-3102 Fitzgerald 100 mg 159.6 EUR

Hemoglobin protein (Mouse)

30-1135 Fitzgerald 5 mg 630 EUR

Mouse Hemoglobin (HB) Protein

20-abx168595 Abbexa
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  • 226.80 EUR
  • 727.20 EUR
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  • 100 ug
  • 10 ug
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  • 200 ug
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Mouse Hemoglobin Beta (HBb) Protein

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  • 777.60 EUR
  • 326.40 EUR
  • 2331.60 EUR
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  • 100 ug
  • 10 ug
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  • 50 ug

Hemoglobin protein

30R-3084 Fitzgerald 100 mg 159.6 EUR

Hemoglobin protein

30-1134 Fitzgerald 1 gram 214.8 EUR

Hemoglobin Protein

abx061580-100ug Abbexa 100 ug 760.8 EUR

Mouse Hemoglobin Alpha 1 (HBa1) Protein

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  • 811.20 EUR
  • 343.20 EUR
  • 2498.40 EUR
  • 961.20 EUR
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  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
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Hemoglobin Ao protein

30-AH32 Fitzgerald 1 mg 181.2 EUR

Human Hemoglobin Protein

abx060919-2g Abbexa 2 g 526.8 EUR

Hemoglobin protein (Rabbit)

30R-3103 Fitzgerald 100 mg 159.6 EUR

Mouse Glycated Hemoglobin A1c (HbA1c) Protein

20-abx168596 Abbexa
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  • 309.60 EUR
  • 1963.20 EUR
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  • 10 ug
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Glycated Hemoglobin Protein

20-abx260544 Abbexa
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  • 276.00 EUR
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  • 0.5 mg
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Glycated Hemoglobin Protein

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Glycated Hemoglobin Protein

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Dog Hemoglobin (HB) Protein

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  • 276.00 EUR
  • 1479.60 EUR
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  • 10 ug
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Pig Hemoglobin (HB) Protein

20-abx167813 Abbexa
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  • 260.40 EUR
  • 1161.60 EUR
  • 526.80 EUR
  • 343.20 EUR
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  • 10 ug
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Rat Hemoglobin (HB) Protein

20-abx167814 Abbexa
  • 393.60 EUR
  • 243.60 EUR
  • 994.80 EUR
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  • 100 ug
  • 10 ug
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Cow Hemoglobin (HB) Protein

20-abx168572 Abbexa
  • 510.00 EUR
  • 276.00 EUR
  • 1378.80 EUR
  • 594.00 EUR
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  • 100 ug
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Human Hemoglobin (HB) Protein

20-abx651581 Abbexa
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  • 226.80 EUR
  • 594.00 EUR
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  • 100 ug
  • 10 ug
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Rabbit Hemoglobin (HB) Protein

20-abx168603 Abbexa
  • 326.40 EUR
  • 226.80 EUR
  • 727.20 EUR
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  • 100 ug
  • 10 ug
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Monkey Hemoglobin (HB) Protein

20-abx168612 Abbexa
  • 510.00 EUR
  • 276.00 EUR
  • 1378.80 EUR
  • 594.00 EUR
  • 376.80 EUR
  • 100 ug
  • 10 ug
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  • 200 ug
  • 50 ug

Hemoglobin A1c protein

30C-CP1048U Fitzgerald 100 ug 198 EUR

Hemoglobin A1c protein

30R-AH012X Fitzgerald 1 mg 812.4 EUR

Hemoglobin A1c protein

30-AH31 Fitzgerald 1 mg 632.4 EUR

Hemoglobin protein (Guinea Pig)

30R-3101 Fitzgerald 100 mg 159.6 EUR

Hemoglobin Ao protein (Glycated)

30-AH33 Fitzgerald 1 mg 489.6 EUR

Hemoglobin Bovine Protein

abx169705-100g Abbexa 100 g 444 EUR

Purified Mouse Hemoglobin

HEMG17-N-1 Alpha Diagnostics 1 mg 270 EUR

Hemoglobin (Mouse) ELISA Kit

E4739-100 Biovision each 744 EUR

Mouse Alpha Hemoglobin Stabilizing Protein (AHSP) ELISA Kit

20-abx258774 Abbexa
  • 8684.40 EUR
  • 4626.00 EUR
  • 1074.00 EUR
  • 10 × 96 tests
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Mouse Alpha Hemoglobin Stabilizing Protein (AHSP) ELISA Kit

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Mouse Alpha hemoglobin stabilizing protein(ERAF) ELISA kit

E03A1852-192T BlueGene 192 tests 1524 EUR

Mouse Alpha hemoglobin stabilizing protein(ERAF) ELISA kit

E03A1852-48 BlueGene 1 plate of 48 wells 624 EUR

Mouse Alpha hemoglobin stabilizing protein(ERAF) ELISA kit

E03A1852-96 BlueGene 1 plate of 96 wells 822 EUR

Human Hemoglobin Mu (HBm) Protein

20-abx067043 Abbexa
  • 878.40 EUR
  • 343.20 EUR
  • 2749.20 EUR
  • 1045.20 EUR
  • 627.60 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Cow Hemoglobin Beta (HBb) Protein

20-abx166293 Abbexa
  • 878.40 EUR
  • 343.20 EUR
  • 2766.00 EUR
  • 1062.00 EUR
  • 627.60 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Mouse Hemoglobin (HB) CLIA Kit

abx197126-96tests Abbexa 96 tests 990 EUR

Mouse Hemoglobin (HB) CLIA Kit

20-abx492738 Abbexa
  • 8853.60 EUR
  • 4719.60 EUR
  • 1093.20 EUR
  • 10 × 96 tests
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Horse Hemoglobin Beta (HBb) Protein

20-abx168036 Abbexa
  • 895.20 EUR
  • 360.00 EUR
  • 2815.20 EUR
  • 1078.80 EUR
  • 644.40 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Sheep Hemoglobin Beta (HBb) Protein

20-abx168611 Abbexa
  • 627.60 EUR
  • 292.80 EUR
  • 1796.40 EUR
  • 727.20 EUR
  • 460.80 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Sheep Hemoglobin Beta (HBb) Protein

20-abx067040 Abbexa
  • 627.60 EUR
  • 292.80 EUR
  • 1796.40 EUR
  • 727.20 EUR
  • 460.80 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Human Hemoglobin Beta (HBb) Protein

20-abx166677 Abbexa
  • 794.40 EUR
  • 326.40 EUR
  • 2448.00 EUR
  • 944.40 EUR
  • 577.20 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Mouse Hemoglobin (HB) ELISA Kit

abx254159-96tests Abbexa 96 tests 764.4 EUR

Mouse Hemoglobin (HB) ELISA Kit

20-abx154130 Abbexa
  • 6373.20 EUR
  • 3404.40 EUR
  • 801.60 EUR
  • 10 × 96 tests
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Mouse hemoglobin (Hb) ELISA Kit

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Mouse hemoglobin (Hb) ELISA Kit

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Mouse Hemoglobin H ELISA kit

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Mouse Hemoglobin H ELISA kit

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Mouse Hemoglobin H ELISA kit

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Mouse Hemoglobin A ELISA kit

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Mouse Hemoglobin A ELISA kit

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Mouse Hemoglobin A ELISA kit

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Mouse hemoglobin (Hb) ELISA kit

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Mouse hemoglobin (Hb) ELISA kit

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Mouse hemoglobin (Hb) ELISA kit

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Mouse Hemoglobin, Mu ELISA kit

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Mouse Hemoglobin, Mu ELISA kit

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Mouse Hemoglobin, Mu ELISA kit

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Mouse Hemoglobin(HB)ELISA kit

GA-E0840MS-48T GenAsia Biotech 48T 403.2 EUR

Mouse Hemoglobin(HB)ELISA kit

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Mouse Hemoglobin (HB) ELISA Kit

SEB409Mu-10x96wellstestplate Cloud-Clone 10x96-wells test plate 3305.89 EUR

Mouse Hemoglobin (HB) ELISA Kit

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Mouse Hemoglobin (HB) ELISA Kit

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Mouse Hemoglobin (HB) ELISA Kit

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Mouse Hemoglobin (HB) ELISA Kit

4-SEB409Mu Cloud-Clone
  • 3366.00 EUR
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  • 488.40 EUR
  • 10 plates of 96 wells
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  • 1 plate of 96 wells

Mouse Hemoglobin ELISA Kit (HB)

RK02880 Abclonal 96 Tests 625.2 EUR

Mouse Hemoglobin(HB)ELISA kit

QY-E20731 Qayee Biotechnology 96T 433.2 EUR

Mouse Hemoglobin (HB)ELISA kit

YLA1089MO-48T Shanghai YL Biotech 48T 465 EUR

Mouse Hemoglobin (HB)ELISA kit

YLA1089MO-96T Shanghai YL Biotech 96T 600 EUR

Hemoglobin Zeta Recombinant Protein

91-211 ProSci 0.05 mg 852.9 EUR

Human Hemoglobin Delta (HBd) Protein

20-abx166874 Abbexa
  • 811.20 EUR
  • 343.20 EUR
  • 2498.40 EUR
  • 961.20 EUR
  • 577.20 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

alpha Hemoglobin Stabilizing Protein

20-abx260463 Abbexa
  • 276.00 EUR
  • 2798.40 EUR
  • 393.60 EUR
  • 10 ug
  • 1 mg
  • 50 ug

Hemoglobin alpha Recombinant Protein

92-524 ProSci 0.05 mg 739.5 EUR

Hemoglobin Subunit Zeta (HBZ) Protein

20-abx260582 Abbexa
  • 3433.20 EUR
  • 393.60 EUR
  • 276.00 EUR
  • 1 mg
  • 25 ug
  • 5 ug

Hemoglobin subunit alpha 2 Protein

20-abx262533 Abbexa
  • 4101.60 EUR
  • 393.60 EUR
  • 276.00 EUR
  • 1 mg
  • 20 ug
  • 5 ug

Chicken Hemoglobin Beta (HBb) Protein

20-abx067041 Abbexa
  • 560.40 EUR
  • 276.00 EUR
  • 1528.80 EUR
  • 644.40 EUR
  • 393.60 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Cow Hemoglobin Alpha 1 (HBa1) Protein

20-abx166292 Abbexa
  • 878.40 EUR
  • 343.20 EUR
  • 2766.00 EUR
  • 1062.00 EUR
  • 627.60 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

CLIA kit for Mouse HB (Hemoglobin)

E-CL-M0345 Elabscience Biotech 1 plate of 96 wells 700.8 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, E.coli-1mg

QP6146-ec-1mg EnQuireBio 1mg 2763.6 EUR

ELISA kit for Mouse HB (Hemoglobin)

E-EL-M0573 Elabscience Biotech 1 plate of 96 wells 452.4 EUR

ELISA kit for Mouse HB (Hemoglobin)

ELK1354 ELK Biotech 1 plate of 96 wells 446.4 EUR

Mouse Hemoglobin subunit alpha (Hba)

1-CSB-YP010147MO Cusabio
  • 604.80 EUR
  • 318.00 EUR
  • 2198.40 EUR
  • 915.60 EUR
  • 1459.20 EUR
  • 400.80 EUR
  • 100ug
  • 10ug
  • 1MG
  • 200ug
  • 500ug
  • 50ug

Mouse Hemoglobin subunit alpha (Hba)

1-CSB-EP010147MO Cusabio
  • 606.00 EUR
  • 318.00 EUR
  • 2192.40 EUR
  • 919.20 EUR
  • 1461.60 EUR
  • 402.00 EUR
  • 100ug
  • 10ug
  • 1MG
  • 200ug
  • 500ug
  • 50ug

Recombinant Mouse Hemoglobin subunit alpha Protein, His, E.coli-100ug

QP6146-ec-100ug EnQuireBio 100ug 685.2 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, E.coli-10ug

QP6146-ec-10ug EnQuireBio 10ug 326.4 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, E.coli-200ug

QP6146-ec-200ug EnQuireBio 200ug 1077.6 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, E.coli-500ug

QP6146-ec-500ug EnQuireBio 500ug 1816.8 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, E.coli-50ug

QP6146-ec-50ug EnQuireBio 50ug 434.4 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, Yeast-10ug

QP6146-ye-10ug EnQuireBio 10ug 326.4 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, Yeast-1mg

QP6146-ye-1mg EnQuireBio 1mg 2763.6 EUR

Recombinant Mouse Hemoglobin subunit alpha Protein, His, Yeast-50ug

QP6146-ye-50ug EnQuireBio 50ug 424.8 EUR

Mouse Hemoglobin Polyclonal Antibody

A57573 EpiGentek
  • 684.66 EUR
  • Ask for price
  • Ask for price
  • 423.50 EUR
  • 100 µg
  • 20 ul
  • 50 ul
  • 100 ul

Human Hemoglobin Alpha 1 (HBa1) Protein

20-abx067039 Abbexa
  • 777.60 EUR
  • 326.40 EUR
  • 2397.60 EUR
  • 927.60 EUR
  • 560.40 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Human Hemoglobin Gamma 1 (HBg1) Protein

20-abx653709 Abbexa
  • 693.60 EUR
  • 309.60 EUR
  • 2064.00 EUR
  • 828.00 EUR
  • 510.00 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Human Hemoglobin Gamma 2 (HBg2) Protein

20-abx653710 Abbexa
  • 693.60 EUR
  • 309.60 EUR
  • 2064.00 EUR
  • 828.00 EUR
  • 510.00 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Horse Hemoglobin Alpha 1 (HBa1) Protein

20-abx166390 Abbexa
  • 878.40 EUR
  • 343.20 EUR
  • 2715.60 EUR
  • 1045.20 EUR
  • 627.60 EUR
  • 100 ug
  • 10 ug
  • 1 mg
  • 200 ug
  • 50 ug

Hemoglobin

E61I012 EnoGene 100ug 411.6 EUR

Hemoglobin

GL7038-10 Glentham Life Sciences 10 37.7 EUR

Hemoglobin theta-1 Recombinant Protein

92-519 ProSci 0.05 mg 739.5 EUR

Species in an early divergent branch of vertebrates, the cyclostomes (represented by hagfish and lampreys), also use a heme-containing globin for oxygen transport, but surprisingly it is more closely related to CYGB than to the gnathostome hemoglobins. This suggests that the oxygen transport function of heme-containing globins arose through independent, convergent evolution in the two major vertebrate branches. In cyclostomes, oxygen transport appears to be derived by co-option of the widely expressed  CYGB  gene.

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