Membrane transport protein

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## Contents

* 1 Overview
* 2 Facilitated diffusion
* 3 Active transport
* 3.1 Binding dependent active transport
* 4 Classification and examples
* 5 More examples
* 6 See also
* 7 External links

## Overview

[edit | edit source]

A membrane transport protein (or simply transporter) is a protein involved in the movement of ions, small molecules, or macromolecules, such as another protein across a biological membrane. Transport proteins are integral membrane proteins; that is they exist within and span the membrane across which they transport substances. The proteins may assist in the movement of substances by facilitated diffusion or active transport.

## Facilitated diffusion

[edit | edit source]

A facilitated diffusion protein speeds the movement of a chemical through a membrane in the absence of energy input; therefore, the transported chemical can only move down a concentration gradient. This can be accomplished by the formation of a high-specificity pore or channel that spans the membrane. These polar "holes" through the membrane are lined by specific amino acids residues which lower the energy barrier to the movement of polar molecules.

## Active transport

[edit | edit source]

Transport proteins are also used in active transport, which by definition does require an energy input.

Chemiosmotic transport utilizes electrochemical gradients to drive transport. As the creation and maintenance of chemiosmotic gradients require energy input from the cell, this is a form of active transport. Prokaryotes typically use hydrogen ions as the driving force for chemiosmotic transport, while eukaryotes typically use sodium ions. A symporter/coporter transports a chemical in the same direction as the electrochemical gradient, while an antiporter moves the target chemical in a direction opposite to the gradient.

The uniporter is also often included as a category of chemiosmotic transporter, although a uniporter can also be considered a facilitated diffusion protein on the basis of function.

### Binding dependent active transport

[edit | edit source]

Binding dependent active transport also moves the targeted chemical against a concentration gradient, but uses stored chemical energy, typically in the form of adenosine triphosphate, to power the transport. Generally speaking, a binding dependent transport system consists of a membrane spanning component with a high degree of specifity. The membrane spanning component changes configuration with the aid of chemical energy input (often through the use of an associated ATPase protein), thus translocating the chemical from one side of the membrane to the other.

By some definitions, proteins that catalyze the ligation of phosphate or coenzyme groups to a catabolized chemical can be considered active transport proteins in that they drive the uptake of a chemical by maintaining a steep functional concentration gradient. This pheonomenon is termed group translocation in the case of sugar phosphorylation and vectoral acylation or vectoral esterification in the case of fatty acid coenzyme A ligation.

## Classification and examples

[edit | edit source]

Classification of transmembrane transporters according to TCDB and examples of transporters with known 3D structure:

1\. Channels/Pores

*     * Voltage-gated ion channel like, including potassium channels KcsA and KvAP, and inward-rectifier potassium ion channel Kirbac [1]
* Large-conductance mechanosensitive channel, MscL [2]
* Small-conductance mechanosensitive ion channel (MscS) [3]
* CorA metal ion transporters [4]
* Ligand-gated ion channel of neurotransmitter receptors (acetylcholine receptor) [5]
* Aquaporins [6]
* Chloride channels [7]
* Outer membrane auxiliary proteins (polysaccharide transporter) [8]

2\. Electrochemical Potential-driven transporters

*     * Mitochondrial carrier proteins [9]
* Major Facilitator Superfamily (Glycerol-3-hosphate transporter, Lactose permease, and Multidrug transporter EmrD) [10]
* Resistance-nodulation-cell division (multidrug efflux transporter AcrB, see multidrug resistance)[11]
* Dicarboxylate/amino acid:cation symporter (proton glutamate symporter) [12]
* Monovalent cation/proton antiporter (Sodium/proton antiporter 1 NhaA) [13]
* Neurotransmitter sodium symporter [14]
* Ammonia transporters [15]
* Drug/Metabolite Transporter (small multidrug resistance transporter EmrE - the structures are retracted as erroneous) [16]

3\. Primary Active Transporters

* Light absorption-driven transporters:
* Bacteriorhodopsin-like proteins including rhodopsin (see also opsin)[17]
* Bacterial photosynthetic reaction centres and photosystems I and II [18]
* Light harvesting complexes from bacteria and chloroplasts [19]
* Oxidoreduction-driven transporters
* Transmembrane cytochrome b-like proteins [20]: coenzyme Q - cytochrome c reductase (cytochrome bc1 ); cytochrome b6f complex; formate dehydrogenase, respiratory nitrate reductase; succinate - coenzyme Q reductase (fumarate reductase); and succinate dehydrogenase. See electron transport chain.
* Cytochrome c oxidases [21] from bacteria and mitochondria
* Electrochemical potential-driven transporters
* Proton or sodium translocating F-type and V-type ATPases [22]
* P-P-bond hydrolysis-driven transporters
* P-type calcium ATPase (five different conformations) [23]
* Calcium ATPase regulators phospholamban and sarcolipin[24]
* ABC transporters: BtuCD, multidrug transporter, and molybdate uptake transporter
* General secretory pathway (Sec) translocon (preprotein translocase SecY) [25]

4\. Group Translocators

5\. Transport Electron Carriers

*     * Disulfide bond formation protein B (DsbB) [26]

8\. Accessory Factors Involved in Transport

## More examples

[edit | edit source]

* Karyopherin
* Mitochondrial membrane transport protein
* Glucose transporter
* ATP-binding cassette transporter genes
* P-glycoprotein
* CD98
* CD36
* Neurotransmitter transporters
* Glutamate transporter
* Norepinephrine transporter
* Vesicular monoamine transporter
* V-ATPase
* Ion transporters
* Na+/K+-ATPase
* Plasma membrane Ca2+ ATPase
* Proton pump
* Hydrogen potassium ATPase
* Sodium-chloride symporter

## See also

[edit | edit source]

* Carrier protein
* Ion channel
* P-loop
* Solute carrier family (classification)
* TC number (classification)
* Vesicular transport protein

## External links

[edit | edit source]

* Template:Dorlands
* Membrane+transport+proteins at the US National Library of Medicine Medical Subject Headings (MeSH)

* v
* t
* e

Proteins: carrier proteins

Fatty acid|

* FABP1
* FABP2
* FABP3
* FABP4
* FABP5
* FABP6
* FABP7

Hormone|

* peptide hormone: Follistatin
* Growth hormone binding protein
* Insulin-like growth factor binding protein
* IGFBP1
* IGFBP2
* IGFBP3
* IGFBP4
* IGFBP5
* IGFBP6
* IGFBP7
* Neurophysins
* Neurophysin I
* II

* steroid hormone: Sex hormone binding globulin/Androgen binding protein
* Transcortin
* Thyroxine-binding globulin
* Transthyretin

Metal/element|

* calcium
* Calcium-binding protein
* Calmodulin-binding proteins
* copper
* Ceruloplasmin
* iron
* Iron-binding proteins
* Transferrin receptor

Vitamin|

* Retinol binding protein
* 1
* 2
* 3
* 4
* Transcobalamin

Pigment|

* Plant Light-Harvesting Complex
* Orange Carotenoid Protein
* Phycobiliprotein
* Photosynthetic Reaction Centers
* Phytochrome
* Rhodopsin

Other|

* Acyl carrier protein
* Adaptor protein
* Cholesterylester transfer protein
* F-box protein
* GTP-binding protein
* Latent TGF-beta binding protein
* Major urinary proteins
* Membrane transport protein
* Odorant binding protein

* v
* t
* e

Membrane proteins, carrier proteins: membrane transport proteins solute carrier (TC 2A)

By group

SLC1–10

| (1):|

* high affinity glutamate and neutral amino-acid transporter
* SLC1A1
* 2
* 3
* 4
* 5
* 6
* 7

|

(2):|

* facilitative GLUT transporter
* SLC2A1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* 9
* 10
* 11
* 12
* 13
* 14

(3):|

* heavy subunits of heterodimeric amino-acid transporters
* SLC3A1
* 2

(4):|

* bicarbonate transporter
* SLC4A1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* 9
* 10
* 11

(5):|

* sodium glucose cotransporter
* SLC5A1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* 9
* 10
* 11
* 12

(6):|

* sodium\- and chloride\- dependent sodium:neurotransmitter symporters
* SLC6A1
* SLC6A2
* SLC6A3
* SLC6A4
* SLC6A5
* SLC6A6
* SLC6A7
* SLC6A8
* SLC6A9
* SLC6A10
* SLC6A11
* SLC6A12
* SLC6A13
* SLC6A14
* SLC6A15
* SLC6A16
* SLC6A17
* SLC6A18
* SLC6A19
* SLC6A20

(7):|

* cationic amino-acid transporter/glycoprotein-associated
* SLC7A1
* SLC7A2
* SLC7A3
* SLC7A4
* glycoprotein-associated/light or catalytic subunits of heterodimeric amino-acid transporters
* SLC7A5
* SLC7A6
* SLC7A7
* SLC7A8
* SLC7A9
* SLC7A10
* SLC7A11
* SLC7A13
* SLC7A14

(8):|

* Na+/Ca2+ exchanger
* SLC8A1
* SLC8A2
* SLC8A3

(9):|

* Na+/H+ exchanger
* SLC9A1
* SLC9A2
* SLC9A3
* SLC9A4
* SLC9A5
* SLC9A6
* SLC9A7
* SLC9A8
* SLC9A9
* SLC9A10
* SLC9A11

(10):|

* sodium bile salt cotransport
* SLC10A1
* SLC10A2
* SLC10A3
* SLC10A4
* SLC10A5
* SLC10A6
* SLC10A7
* 10A1
* 10A2
* 10A3
* 10A7

SLC11–20

| (11):|

* proton coupled metal ion transporter
* SLC11A1
* SLC11A211A3

|

(12):|

* electroneutral cation-Cl cotransporter
* SLC12A1
* SLC12A2
* SLC12A3
* SLC12A4
* SLC12A5
* SLC12A6
* SLC12A7
* SLC12A8
* SLC12A9

(13):|

* human Na+-sulfate/carboxylate cotransporter
* SLC13A1
* SLC13A2
* SLC13A3
* SLC13A4
* SLC13A5

(14):|

* urea transporter
* SLC14A1
* SLC14A2

(15):|

* proton oligopeptide cotransporter
* SLC15A1
* SLC15A2
* SLC15A3
* SLC15A4

(16):|

* monocarboxylate transporter
* SLC16A1
* SLC16A2
* SLC16A3
* SLC16A4
* SLC16A5
* SLC16A6
* SLC16A7
* SLC16A8
* SLC16A9
* SLC16A10
* SLC16A11
* SLC16A12
* SLC16A13
* SLC16A14

(17):|

* Vesicular glutamate transporter 1
* SLC17A1
* SLC17A2
* SLC17A3
* SLC17A4
* SLC17A5
* SLC17A6
* SLC17A7
* SLC17A8
* SLC17A9

(18):|

* vesicular monoamine transporter
* SLC18A1
* SLC18A2
* SLC18A3

(19):|

* folate/thiamine transporter
* SLC19A1
* SLC19A2
* SLC19A3

(20):|

* type III Na+-phosphate cotransporter
* SLC20A1
* SLC20A2

SLC21–30

| (21):|

* Organic anion-transporting polypeptide
* SLCO1A2
* SLCO1B1
* SLCO1B3
* SLCO1B4
* SLCO1C1
*     * SLCO2A1
* SLCO2B1
*     * SLCO3A1
*     * SLCO4A1
* SLCO4C1
*     * SLCO5A1(SLCO6A1)

|

(22):|

* organic cation/anion/zwitterion transporter
* SLC22A1
* SLC22A2
* SLC22A3
* SLC22A4
* SLC22A5
* SLC22A6
* SLC22A7
* SLC22A8
* SLC22A9
* SLC22A10
* SLC22A11
* SLC22A12
* SLC22A13
* SLC22A14
* SLC22A15
* SLC22A16
* SLC22A17
* SLC22A18
* SLC22A19
* SLC22A20

(23):|

* Na+-dependent ascorbic acid transporter
* SLC23A1
* SLC23A2
* SLC23A3
* SLC23A4

(24):|

* Na+/(Ca2+-K+) exchanger
* SLC24A1
* SLC24A2
* SLC24A3
* SLC24A4
* SLC24A5
* SLC24A6

(25):|

* mitochondrial carrier
* SLC25A1
* SLC25A2
* SLC25A3
* SLC25A4
* SLC25A5
* SLC25A6
* SLC25A7
* SLC25A8
* SLC25A9
* SLC25A10
* SLC25A11
* SLC25A12
* SLC25A13
* SLC25A14
* SLC25A15
* SLC25A16
* SLC25A17
* SLC25A18
* SLC25A19
* SLC25A20
* SLC25A21
* SLC25A22
* SLC25A23
* SLC25A24
* SLC25A25
* SLC25A26
* SLC25A27
* SLC25A28
* SLC25A29
* SLC25A30
* SLC25A31
* SLC25A32
* SLC25A33
* SLC25A34
* SLC25A35
* SLC25A36
* SLC25A37
* SLC25A38
* SLC25A39
* SLC25A40
* SLC25A41
* SLC25A42
* SLC25A43
* SLC25A44
* SLC25A45
* SLC25A46

(26):|

* multifunctional anion exchanger
* SLC26A1
* SLC26A2
* SLC26A3
* SLC26A4
* SLC26A5
* SLC26A6
* SLC26A7
* SLC26A8
* SLC26A9
* SLC26A10
* SLC26A11

(27):|

* fatty acid transport proteins
* SLC27A1
* SLC27A2
* SLC27A3
* SLC27A4
* SLC27A5
* SLC27A6

(28):|

* Na+-coupled nucleoside transport (SLC28A1
* SLC28A2
* SLC28A3

(29):|

* facilitative nucleoside transporter
* SLC29A1
* SLC29A2
* SLC29A3
* SLC29A4

(30):|

* zinc efflux
* SLC30A1
* SLC30A2
* SLC30A3
* SLC30A4
* SLC30A5
* SLC30A6
* SLC30A7
* SLC30A8
* SLC30A9
* SLC30A10

SLC31–40

| (31):|

* copper transporter
* SLC31A1

|

(32):|

* Vesicular glutamate transporter 1
* SLC32A1

(33):|

* Acetyl-CoA transporter
* SLC33A1

(34):|

* type II Na+-phosphate cotransporter
* SLC34A1
* SLC34A2
* SLC34A3

(35):|

* nucleoside-sugar transporter
* SLC35A1
* SLC35A2
* SLC35A3
* SLC35A4
* SLC35A5
*     * SLC35B1
* SLC35B2
* SLC35B3
* SLC35B4
*     * SLC35C1
* SLC35C2
*     * SLC35D1
* SLC35D2
* SLC35D3
*     * SLC35E1
* SLC35E2
* SLC35E3
* SLC35E4

(36):|

* proton-coupled amino-acid transporter
* SLC36A1
* SLC36A2
* SLC36A3
* SLC36A436A2

(37):|

* sugar-phosphate/phosphate exchanger
* SLC37A1
* SLC37A2
* SLC37A3
* SLC37A4

(38):|

* System A & N, sodium-coupled neutral amino-acid transporter
* SLC38A1
* SLC38A2
* SLC38A3
* SLC38A4
* SLC38A5
* SLC38A6
* SLC38A10

(39):|

* metal ion transporter
* SLC39A1
* SLC39A2
* SLC39A3
* SLC39A4
* SLC39A5
* SLC39A6
* SLC39A7
* SLC39A8
* SLC39A9
* SLC39A10
* SLC39A11
* SLC39A12
* SLC39A13
* SLC39A14

(40):|

* basolateral iron transporter
* SLC40A1

SLC41–48

| (41):|

* Magnesium transporter E
* SLC41A1
* SLC41A2
* SLC41A3

|

(42):|

* Ammonia transporter
* RhAG
* RhBG
* RhCG

(43):|

* Na+-independent, system-L like amino-acid transporter
* SLC43A1
* SLC43A2
* SLC43A3

(44):|

* Choline-like transporter
* SLC44A1
* SLC44A2
* SLC44A3
* SLC44A4
* SLC44A5

(45):|

* Putative sugar transporter
* SLC45A1
* SLC45A2
* SLC54A3
* SLC45A4

(46):|

* Folate transporter
* SLC46A1
* SLC46A2

(47):|

* multidrug and toxin extrusion
* SLC47A1
* SLC47A2

(48):|

* Heme transporter

SLCO1–4

* O1A2
* O1B1
* O1B3
* O2B1
* O431
* O4A1

Ion pumps

| Symporter, Cotransporter|

* Na+/K+,Cl−
* Na+/Pi3
* Na+/Cl−
* Na+/glucose
* Na+/I−
* Cl−/K+
* 4
* 5

|

Antiporter (exchanger)|

* Na+/H+
* Na+/Ca2+
* Na+/(Ca2+-K+) - Cl−/HCO−
3 (Band 3)
* Cl−-formate
* Cl−-oxalate

see also solute carrier disorders

* v
* t
* e

Membrane transport protein: ion channels (TC 1A)

Ca2+: Calcium channel

| Ligand-gated|

* Inositol trisphosphate receptor
* 1
* 2
* 3
* Ryanodine receptor
* 1
* 2
* 3

|

Voltage-gated|

* L-type/Cavα
* 1.1
* 1.2
* 1.3
* 1.4
* N-type/Cavα2.2
* P-type/Cavα
* 2.1
* Q-type/Cavα2.1
* R-type/Cavα2.3
* T-type/Cavα
* 3.1
* 3.2
* 3.3

* α2δ-subunits
* 1
* 2
* β-subunits
* β1
* β2
* β3
* β4
* γ-subunits
* γ1
* γ2
* γ3
* γ4

* Cation channels of sperm
* 1
* 2
* 3
* 4
* Two-pore channel
* 1
* 2

Na+: Sodium channel

| Constitutively active|

* Epithelial sodium channel
* α
* β
* γ
* δ

|

Proton-gated|

* Amiloride-sensitive cation channel
* 1
* 2
* 3
* 4

Voltage-gated|

* Navα
* 1.1
* 1.2
* 1.3
* 1.4
* 1.5
* 1.6
* 1.7
* 1.8
* 1.9
* 7A
* Navβ
* 1
* 2
* 3
* 4

K+: Potassium channel

| Calcium-activated|

* BK channel
* α1
* β1
* β2
* β3
* β4
* SK channel
* SK1
* SK2
* SK3
* IK channel
* IK1
* KCa
* 1.1
* 2.1
* 2.2
* 2.3
* 3.1
* 4.1
* 4.2
* 5.1

|

Inward-rectifier|

* KATP
* Kir
* 1.1
* 2.1
* 2.2
* 2.3
* 2.4
* 2.6
* GIRK/Kir
* 3.1
* 3.2
* 3.3
* 3.4
* Kir
* 4.1
* 4.2
* 5.1
* 6.1
* 6.2
* 7.1

Tandem pore domain|

* K2P
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* 9
* 10
* 12
* 13
* 15
* 16
* 17
* 18

Voltage-gated|

* Kvα1-6
* 1.1
* 1.2
* 1.3
* 1.4
* 1.5
* 1.6
* 1.7
* 1.8
*     * 2.1
* 2.2
*     * 3.1
* 3.2
* 3.3
* 3.4
*     * 4.1
* 4.2
* 4.3
*     * 5.1
*     * 6.1
* 6.2
* 6.3
* 6.4

* Kvα7-12
* 7.1
* 7.2
* 7.3
* 7.4
* 7.5
*     * 8.1
* 8.2
*     * 9.1
* 9.2
* 9.3
*     * 10.1
* 10.2
*     * 11.1/hERG
* 11.2
* 11.3
*     * 12.1
* 12.2
* 12.3

* Kvβ
* 1
* 2
* 3
* KCNIP
* 1
* 2
* 3
* 4
* minK/ISK
* minK/ISK-like
* MiRP
* 1
* 2
* 3
* Shaker gene

Miscellaneous

| Cl−: Chloride channel|

* Calcium-activated chloride channels
* Anoctamin
* ANO1
* Bestrophin
* 1
* 2
* Chloride Channel Accessory
* 1
* 2
* 3
* 4
* CFTR
* CLCN
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* KA
* KB
* CLIC
* 1
* 2
* 3
* 4
* 5
* 6
* L1
* CLNS
* 1A
* 1B

|

H+: Proton channel|

* HVCN1

M+: CNG cation channel|

* α
* 1
* 2
* 3
* 4
* β
* 1
* 2
* 3
* HCN
* FC
* 1
* 2
* 3
* 4

M+: TRP cation channel|

* TRPA (1)
* TRPC
* 1
* 2
* 3
* 4
* 4AP
* 5
* 6
* 7
* TRPM
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* TRPML
* 1
* 2
* 3
* TRPN
* TRPP
* 1
* 2
* TRPV
* 1
* 2
* 3
* 4
* 5
* 6

H2O (+ solutes): Porin|

* Aquaporin
* 0
* 1
* 2
* 3
* 4
* 5
* 6
* 7
* 8
* 9
* Voltage-dependent anion channel
* 1
* 2
* 3
* General bacterial porin family

Cytoplasm: Gap junction|

* Connexin
* A
* GJA1
* GJA3
* GJA4
* GJA5
* GJA8
* GJA9
* GJA10
* B
* GJB1
* GJB2
* GJB3
* GJB4
* GJB5
* GJB6
* GJB7
* C
* GJC1
* GJC2
* GJC3
* D
* GJD2
* GJD3
* GJD4
* Innexin

By gating mechanism

| Ion channel class|

* Ligand-gated
* Light-gated
* Voltage-gated
* Stretch-activated

|

see also disorders

* v
* t
* e

Membrane transport protein: ion pumps, ATPases / ATP synthase (TC 3A2-3A3)

F-, V-, and A-type ATPase (3.A.2)|

| H+ (F-type)|

* H+ transporting, mitochondrial: ATP5A1
* ATP5B
* ATP5C1
* ATP5C2
* ATP5D
* ATP5E
* ATP5F1
* ATP5G1
* ATP5G2
* ATP5G3
* ATP5H
* ATP5I
* ATP5J
* ATP5J2
* ATP5L
* ATP5L2
* ATP5O
* ATP5S

|

H+ (V-type)|

* H+ transporting, lysosomal: ATP6AP1
* ATP6AP2
* ATP6V1A
* ATP6V1B1
* ATP6V1B2
* ATP6V1C1
* ATP6V1C2
* ATP6V1D
* ATP6V1E1
* ATP6V1E2
* ATP6V1F
* ATP6V1G1
* ATP6V1G2
* ATP6V1G3
* ATP6V1H
* ATP6V0A1
* ATP6V0A2
* ATP6V0A4
* ATP6V0B
* ATP6V0C
* ATP6V0D1
* ATP6V0D2
* ATP6V0E
* ATP6V0E1

* TCIRG1

A-ATPase|

found in Archea

P-type ATPase (3.A.3)|

* 3.A.3.1.1: Na+/K+ transporting: ATP1A1
* ATP1A2
* ATP1A3
* ATP1A4
* ATP1B1
* ATP1B2
* ATP1B3
* ATP1B4
* ATP1G1

* 3.A.3.1.2: H+/K+
* H+/K+ exchanging: ATP4A
* ATP4B

* 3.A.3.1.4: H+/K+ transporting, nongastric: ATP12A

* 3.A.3.2: Ca2+ (SERCA, PMCA, SPCA) / Ca2+ transporting: ATP2A1
* ATP2A2
* ATP2A3
* ATP2B1
* ATP2B2
* ATP2B3
* ATP2B4
* ATP2C1

* 3.A.3.5: Cu2+ transporting: ATP7A
* ATP7B

* 3.A.3.8.8: flippase: ATP8A2

* Other/ungrouped:

* Na+/K+ – H+

* Mg2+ transporting: ATP3
* Class I, type 8: ATP8A1
* ATP8B1
* ATP8B2
* ATP8B3
* ATP8B4
* Class II, type 9: ATP9A
* ATP9B
* Class V, type 10: ATP10A
* ATP10B
* ATP10D
* Class VI, type 11: ATP11A
* ATP11B
* ATP11C
* type 13: ATP13A1
* ATP13A2
* ATP13A3
* ATP13A4
* ATP13A5

see also ATPase disorders

* v
* t
* e

Membrane transport protein: neurotransmitter transporters (TC 2.A.1.2)

Vesicular|

* Vesicular monoamine
* VMAT1
* VMAT2
* Vesicular acetylcholine
* Vesicular glutamate
* Vesicular GABA

Other|

* Monoamine
* DAT
* NET
* SERT
* PMAT
* Excitatory amino-acid transporter
* GABA transporter

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