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ATP synthase
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ATP Synthase
Molecular model of ATP synthase determined by X-ray crystallography. Stator is not shown here.
Identifiers
EC number
3.6.3.14
CAS number
9000-83-3
Databases
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IntEnz view
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BRENDA entry
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KEGG entry
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metabolic pathway
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ATP synthase is an enzyme that creates the energy storage molecule adenosine triphosphate (ATP). ATP is the most commonly used "energy currency" of cells for most organisms. It is formed from adenosine diphosphate (ADP) and inorganic phosphate (Pi). The overall reaction catalyzed by ATP synthase is:
ADP + Pi + H+out ⇌ ATP + H2O + H+in
The formation of ATP from ADP and Pi is energetically unfavorable and would normally proceed in the reverse direction. In order to drive this reaction forward, ATPase couples ATP synthesis during cellular respiration to an electrochemical gradient created by the difference in proton (H+) concentration across the mitochondrial membrane in eukaryotes or the plasma membrane in bacteria. During photosynthesis in plants, ATP is synthesized by ATPase using a proton gradient created in the thylakoid lumen through the thylakoid membrane and into the chloroplast stroma.
ATP synthase consists of two main subunits, FO and F1, which has a rotational motor mechanism allowing for ATP production.[1][2] Because of its rotating subunit ATP Synthase is a molecular machine.
Contents
[hide]
1 Nomenclature
2 Structure and function
2.1 F1 region
2.2 FO region
3 Binding model
4 Physiological role
5 Evolution
6 In different species
6.1 E. coli
6.2 Yeast
6.3 Plant
6.4 Bovine
6.5 Human
7 See also
8 References
9 Further reading
10 External links
Nomenclature[edit]
The F1 fraction derives its name from the term "Fraction 1" and FO (written as a subscript letter "o", not "zero") derives its name from being the binding fraction for oligomycin, a type of naturally-derived antibiotic that is able to inhibit the FO unit of ATP synthase.[3][4] These functional regions consist of different protein subunits — refer to tables.
Structure and function[edit]
Bovine mitochondrial ATP synthase. The FO, F1, axle, and stator regions are color coded magenta, green, orange, and cyan respectively.[5][6]
Simplified model of FOF1-ATPase alias ATP synthase of E. coli. Subunits of the enzyme are labeled accordingly.
Rotation engine of ATP synthase.
Located within the thylakoid membrane and the inner mitochondrial membrane, ATP synthase consists of two regions FO and F1. FO causes rotation of F1 and is made of c-ring and subunits a, b, d, F6. F1 is made of α , β , γ , δ {\displaystyle \alpha ,\beta ,\gamma ,\delta } subunits. F1 has a water-soluble par that can hydrolyze ATP. FO on the other hand has mainly hydrophobic regions. FO F1 creates a pathway for protons movement across the membrane.[7]
F1 region[edit]
The F1 portion of ATP synthase is hydrophilic and responsible for hydrolyzing ATP. Subunits α {\displaystyle \alpha } and β {\displaystyle \beta } make a hexamer with 6 binding sites. Three of them are catalytically inactive and they bind ADP.
Other three subunits catalyze the ATP synthesis. The other F1 subunits γ , δ , ϵ {\displaystyle \gamma ,\delta ,\epsilon } are a part of a rotational motor mechanism. γ {\displaystyle \gamma } subunit allows β {\displaystyle \beta } to go though conformational changes, i.e. closed, half open and open states allows for ATP to be bound and released once synthesized. The F1 particle is large and can be seen in the transmission electron microscope by negative staining.[8] These are particles of 9 nm diameter that pepper the inner mitochondrial membrane.
F1 - Subunits
Subunit
Human Gene
alpha ATP5A1, ATPAF2
beta ATP5B, ATPAF1, C16orf7
gamma ATP5C1
delta ATP5D
epsilon ATP5E
FO region[edit]
FO subunit F6 from the peripheral stalk region of ATP synthase.[9]
Fo is a water insoluble protein with eight subunits and transmembrane ring. The ring has a tetramer shape with protein helix loop helix that goes though conformational change when protonated and deprotonated allowing to push neighboring subunits to rotate causing the spinning of FO which then affects conformation of F1 also, resulting on switching of states of alpha and beta subunits. The FO region of ATP synthase is a proton pore that is embedded in the mitochondrial membrane. It consists of three main subunits a, b, and c, and (in humans) six additional subunits, d, e, f, g, F6, and 8 (or A6L). An atomic model for the dimeric yeast FO region was determined by cryo-EM at an overall resolution of 3.6 Å [10]
FO-Main subunits
Subunit
Human Gene
a ATP6
b ATP5F1
c ATP5G1, ATP5G2, ATP5G3
Binding model[edit]
Mechanism of ATP synthase. ADP and Pi (pink) shown being combined into ATP (red), and the rotating γ (gamma) subunit in black causing conformation.
Depiction of ATP synthase using the chemiosmotic proton gradient to power ATP synthesis through oxidative phosphorylation.
In the 1960s through the 1970s, Paul Boyer, a UCLA Professor, developed the binding change, or flip-flop, mechanism theory, which postulated that ATP synthesis is dependent on a conformational change in ATP synthase generated by rotation of the gamma subunit. The research group of John E. Walker, then at the MRC Laboratory of Molecular Biology in Cambridge, crystallized the F1 catalytic-domain of ATP synthase. The structure, at the time the largest asymmetric protein structure known, indicated that Boyer's rotary-catalysis model was, in essence, correct. For elucidating this, Boyer and Walker shared half of the 1997 Nobel Prize in Chemistry.
The crystal structure of the F1 showed alternating alpha and beta subunits (3 of each), arranged like segments of an orange around a rotating asymmetrical gamma subunit. According to the current model of ATP synthesis (known as the alternating catalytic model), the transmembrane potential created by (H+) proton cations supplied by the electron transport chain, drives the (H+) proton cations from the intermembrane space through the membrane via the FO region of ATP synthase. A portion of the FO (the ring of c-subunits) rotates as the protons pass through the membrane. The c-ring is tightly attached to the asymmetric central stalk (consisting primarily of the gamma subunit), causing it to rotate within the alpha3beta3 of F1 causing the 3 catalytic nucleotide binding sites to go through a series of conformational changes that leads to ATP synthesis. The major F1 subunits are prevented from rotating in sympathy with the central stalk rotor by a peripheral stalk that joins the alpha3beta3 to the non-rotating portion of FO. The structure of the intact ATP synthase is currently known at low-resolution from electron cryo-microscopy (cryo-EM) studies of the complex. The cryo-EM model of ATP synthase suggests that the peripheral stalk is a flexible structure that wraps around the complex as it joins F1 to FO. Under the right conditions, the enzyme reaction can also be carried out in reverse, with ATP hydrolysis driving proton pumping across the membrane.
The binding change mechanism involves the active site of a β subunit's cycling between three states.[11] In the "loose" state, ADP and phosphate enter the active site; in the adjacent diagram, this is shown in pink. The enzyme then undergoes a change in shape and forces these molecules together, with the active site in the resulting "tight" state (shown in red) binding the newly produced ATP molecule with very high affinity. Finally, the active site cycles back to the open state (orange), releasing ATP and binding more ADP and phosphate, ready for the next cycle of ATP production.[12]
Physiological role[edit]
Like other enzymes, the activity of F1FO ATP synthase is reversible. Large-enough quantities of ATP cause it to create a transmembrane proton gradient, this is used by fermenting bacteria that do not have an electron transport chain, but rather hydrolyze ATP to make a proton gradient, which they use to drive flagella and the transport of nutrients into the cell.
In respiring bacteria under physiological conditions, ATP synthase, in general, runs in the opposite direction, creating ATP while using the proton motive force created by the electron transport chain as a source of energy. The overall process of creating energy in this fashion is termed oxidative phosphorylation. The same process takes place in the mitochondria, where ATP synthase is located in the inner mitochondrial membrane and the F1-part projects into mitochondrial matrix. The consumption of ATP by ATP-synthase pumps proton cations into the matrix.
Evolution[edit]
The evolution of ATP synthase is thought to have been modular whereby two functionally independent subunits became associated and gained new functionality.[13][14] This association appears to have occurred early in evolutionary history, because essentially the same structure and activity of ATP synthase enzymes are present in all kingdoms of life.[13] The F-ATP synthase displays high functional and mechanistic similarity to the V-ATPase.[15] However, whereas the F-ATP synthase generates ATP by utilising a proton gradient, the V-ATPase generates a proton gradient at the expense of ATP, generating pH values of as low as 1.[16]
The F1 region also shows significant similarity to hexameric DNA helicases, and the FO region shows some similarity to H+
-powered flagellar motor complexes.[15] The α3β3 hexamer of the F1 region shows significant structural similarity to hexameric DNA helicases; both form a ring with 3-fold rotational symmetry with a central pore. Both have roles dependent on the relative rotation of a macromolecule within the pore; the DNA helicases use the helical shape of DNA to drive their motion along the DNA molecule and to detect supercoiling, whereas the α3β3 hexamer uses the conformational changes through the rotation of the γ subunit to drive an enzymatic reaction.[17]
The H+
motor of the FO particle shows great functional similarity to the H+
motors that drive flagella.[15] Both feature a ring of many small alpha-helical proteins that rotate relative to nearby stationary proteins, using a H+
potential gradient as an energy source. This link is tenuous, however, as the overall structure of flagellar motors is far more complex than that of the FO particle and the ring with about 30 rotating proteins is far larger than the 10, 11, or 14 helical proteins in the FO complex.
The modular evolution theory for the origin of ATP synthase suggests that two subunits with independent function, a DNA helicase with ATPase activity and a H+
motor, were able to bind, and the rotation of the motor drove the ATPase activity of the helicase in reverse.[13][17] This complex then evolved greater efficiency and eventually developed into today's intricate ATP synthases. Alternatively, the DNA helicase/H+
motor complex may have had H+
pump activity with the ATPase activity of the helicase driving the H+
motor in reverse.[13] This may have evolved to carry out the reverse reaction and act as an ATP synthase.[14][18][19]
In different species[edit]
E. coli[edit]
E. coli ATP synthase is the simplest known form of ATP synthase, with 8 different subunit types.[20]
Yeast[edit]
Yeast ATP synthase is one of the best-studied eukaryotic ATP synthases; and five F1, eight FO subunits, and seven associated proteins have been identified.[7] Most of these proteins have homologues in other eukaryotes.[21][22][23]
Plant[edit]
In plants, ATP synthase is also present in chloroplasts (CF1FO-ATP synthase). The enzyme is integrated into thylakoid membrane; the CF1-part sticks into stroma, where dark reactions of photosynthesis (also called the light-independent reactions or the Calvin cycle) and ATP synthesis take place. The overall structure and the catalytic mechanism of the chloroplast ATP synthase are almost the same as those of the mitochondrial enzyme. However, in chloroplasts, the proton motive force is generated not by respiratory electron transport chain but by primary photosynthetic proteins.
Bovine[edit]
The ATP synthase isolated from bovine heart mitochondria (Bos taurus) is, in terms of biochemistry and structure, the best-characterized ATP synthase. Beef heart is used as a source for the enzyme because of the high concentration of mitochondria in cardiac muscle.[24][25][26]
Human[edit]
The following is a list of humans genes that encode components of ATP synthases:
ATP5A1, ATP5AL1
ATP5B, ATP5BL1
ATP5C2, ATP5D, ATP5E, ATP5F1, ATP5G1, ATP5G2, ATP5G3, ATP5H, ATP5HP1, ATP5I, ATP5J, ATP5J2, ATP5L, ATP5L2, ATP5O, ATP5S
ATP6, ATP6AP1, ATP6AP2
ATPSBL1, ATPSBL2
MT-ATP6, MT-ATP8
See also[edit]
ATP10 protein required for the assembly of the FO sector of the mitochondrial ATPase complex.
Chloroplast
Electron transfer chain
Flavoprotein
Mitochondrion
Oxidative phosphorylation
P-ATPase
Proton pump
Rotating locomotion in living systems
Transmembrane ATPase
V-ATPase
References[edit]
Jump up ^ Okuno D, Iino R, Noji H (April 2011). "Rotation and structure of FOF1-ATP synthase". Journal of Biochemistry. 149 (6): 655–64. doi:10.1093/jb/mvr049. PMID 21524994.
Jump up ^ Junge W, Nelson N (June 2015). "ATP synthase". Annual Review of Biochemistry. 84: 631–57. doi:10.1146/annurev-biochem-060614-034124. PMID 25839341.
Jump up ^ Kagawa Y, Racker E (May 1966). "Partial resolution of the enzymes catalyzing oxidative phosphorylation. 8. Properties of a factor conferring oligomycin sensitivity on mitochondrial adenosine triphosphatase". The Journal of Biological Chemistry. 241 (10): 2461–6. PMID 4223640.
Jump up ^ Mccarty RE (November 1992). "A plant biochemist's view of H+
-ATPases and ATP synthases". The Journal of Experimental Biology. 172 (Pt 1): 431–441. PMID 9874753.
Jump up ^ PDB: 5ARA; Zhou A, Rohou A, Schep DG, Bason JV, Montgomery MG, Walker JE, Grigorieff N, Rubinstein JL (October 2015). "Structure and conformational states of the bovine mitochondrial ATP synthase by cryo-EM". eLife. 4: e10180. doi:10.7554/eLife.10180. PMC 4718723  . PMID 26439008.
Jump up ^ Goodsell D (December 2005). "ATP Synthase". Molecule of the Month. RCSB PDB. doi:10.2210/rcsb_pdb/mom_2005_12.
^ Jump up to: a b Velours J, Paumard P, Soubannier V, Spannagel C, Vaillier J, Arselin G, Graves PV (May 2000). "Organisation of the yeast ATP synthase F(0):a study based on cysteine mutants, thiol modification and cross-linking reagents". Biochimica et Biophysica Acta. 1458 (2–3): 443–56. doi:10.1016/S0005-2728(00)00093-1. PMID 10838057.
Jump up ^ Fernandez Moran H, Oda T, Blair PV, Green DE (July 1964). "A macromolecular repeating unit of mitochondrial structure and function. Correlated electron microscopic and biochemical studies of isolated mitochondria and submitochondrial particles of beef heart muscle". The Journal of Cell Biology. 22 (1): 63–100. doi:10.1083/jcb.22.1.63. PMC 2106494  . PMID 14195622.
Jump up ^ PDB: 1VZS; Carbajo RJ, Silvester JA, Runswick MJ, Walker JE, Neuhaus D (2004). "Solution structure of subunit F(6) from the peripheral stalk region of ATP synthase from bovine heart mitochondria". Journal of Molecular Biology. 342 (2): 593–603. doi:10.1016/j.jmb.2004.07.013. PMID 15327958.
Jump up ^ Guo H, Bueler SA, Rubinstein JL (Nov 2017). "Atomic model for the dimeric FO region of mitochondrial ATP synthase". Science. 358 (6365): 936–40. doi:10.1126/science.aao4815. PMID 29074581.
Jump up ^ Gresser MJ, Myers JA, Boyer PD (October 1982). "Catalytic site cooperativity of beef heart mitochondrial F1 adenosine triphosphatase. Correlations of initial velocity, bound intermediate, and oxygen exchange measurements with an alternating three-site model". The Journal of Biological Chemistry. 257 (20): 12030–8. PMID 6214554.
Jump up ^ Nakamoto RK, Baylis Scanlon JA, Al-Shawi MK (August 2008). "The rotary mechanism of the ATP synthase". Archives of Biochemistry and Biophysics. 476 (1): 43–50. doi:10.1016/j.abb.2008.05.004. PMC 2581510  . PMID 18515057.
^ Jump up to: a b c d Doering C, Ermentrout B, Oster G (December 1995). "Rotary DNA motors". Biophysical Journal. 69 (6): 2256–67. Bibcode:1995BpJ....69.2256D. doi:10.1016/S0006-3495(95)80096-2. PMC 1236464  . PMID 8599633.
^ Jump up to: a b Crofts A. "Lecture 10:ATP synthase". Life Sciences at the University of Illinois at Urbana-Champaign.
^ Jump up to: a b c "ATP Synthase". InterPro Database.
Jump up ^ Kim MS, Jang J, Ab Rahman NB, Pethe K, Berry EA, Huang LS (June 2015). "Isolation and Characterization of a Hybrid Respiratory Supercomplex Consisting of Mycobacterium tuberculosis Cytochrome bcc and Mycobacterium smegmatis Cytochrome aa3". The Journal of Biological Chemistry. 290 (23): 14350–60. doi:10.1074/jbc.m114.624312. PMC 4505504  . PMID 25861988.
^ Jump up to: a b Martinez LO, Jacquet S, Esteve JP, Rolland C, Cabezón E, Champagne E, Pineau T, Georgeaud V, Walker JE, Tercé F, Collet X, Perret B, Barbaras R (January 2003). "Ectopic beta-chain of ATP synthase is an apolipoprotein A-I receptor in hepatic HDL endocytosis". Nature. 421 (6918): 75–9. Bibcode:2003Natur.421...75M. doi:10.1038/nature01250. PMID 12511957.
Jump up ^ Cross RL, Taiz L (January 1990). "Gene duplication as a means for altering H+/ATP ratios during the evolution of FOF1 ATPases and synthases". FEBS Letters. 259 (2): 227–9. doi:10.1016/0014-5793(90)80014-a. PMID 2136729.
Jump up ^ Cross RL, Müller V (October 2004). "The evolution of A-, F-, and V-type ATP synthases and ATPases: reversals in function and changes in the H+/ATP coupling ratio". FEBS Letters. 576 (1–2): 1–4. doi:10.1016/j.febslet.2004.08.065. PMID 15473999.
Jump up ^ Ahmad Z, Okafor F, Laughlin TF. "Role of Charged Residues in the Catalytic Sites of Escherichia coli ATP Synthase". Journal of Amino Acids. 2011: 785741. doi:10.4061/2011/785741. PMID 22312470.
Jump up ^ Devenish RJ, Prescott M, Roucou X, Nagley P (May 2000). "Insights into ATP synthase assembly and function through the molecular genetic manipulation of subunits of the yeast mitochondrial enzyme complex". Biochimica et Biophysica Acta. 1458 (2–3): 428–42. doi:10.1016/S0005-2728(00)00092-X. PMID 10838056.
Jump up ^ Kabaleeswaran V, Puri N, Walker JE, Leslie AG, Mueller DM (November 2006). "Novel features of the rotary catalytic mechanism revealed in the structure of yeast F1 ATPase". The EMBO Journal. 25 (22): 5433–42. doi:10.1038/sj.emboj.7601410. PMID 17082766.
Jump up ^ Stock D, Leslie AG, Walker JE (November 1999). "Molecular architecture of the rotary motor in ATP synthase". Science. 286 (5445): 1700–5. doi:10.1126/science.286.5445.1700. PMID 10576729.
Jump up ^ Abrahams JP, Leslie AG, Lutter R, Walker JE (August 1994). "Structure at 2.8 A resolution of F1-ATPase from bovine heart mitochondria". Nature. 370 (6491): 621–8. Bibcode:1994Natur.370..621A. doi:10.1038/370621a0. PMID 8065448.
Jump up ^ Gibbons C, Montgomery MG, Leslie AG, Walker JE (November 2000). "The structure of the central stalk in bovine F(1)-ATPase at 2.4 A resolution". Nature Structural Biology. 7 (11): 1055–61. doi:10.1038/80981. PMID 11062563.
Jump up ^ Menz RI, Walker JE, Leslie AG (August 2001). "Structure of bovine mitochondrial F(1)-ATPase with nucleotide bound to all three catalytic sites: implications for the mechanism of rotary catalysis". Cell. 106 (3): 331–41. doi:10.1016/s0092-8674(01)00452-4. PMID 11509182.
Further reading[edit]
Nick Lane: The Vital Question - Energy, Evolution, and the Origins of Complex Life, Ww Norton, 2015-07-20, ISBN 978-0393088816 (Link points to Figure 10 showing model of ATP synthase)
External links[edit]
Boris A. Feniouk: "ATP synthase — a splendid molecular machine"
Well illustrated ATP synthase lecture by Antony Crofts of the University of Illinois at Urbana-Champaign.
Proton and Sodium translocating F-type, V-type and A-type ATPases in OPM database
The Nobel Prize in Chemistry 1997 to Paul D. Boyer and John E. Walker for the enzymatic mechanism of synthesis of ATP; and to Jens C. Skou, for discovery of an ion-transporting enzyme, Na+
, K+
-ATPase.
Harvard Multimedia Production Site — Videos — ATP synthesis animation
David Goodsell: "ATP Synthase- Molecule of the Month"
[show]
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
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MT-CYB
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MT-CO1
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ATP synthase
MT-ATP6
MT-ATP8
tRNA
MT-TA
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MT-TL1
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MT-TS1
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MT-TY
see also mitochondrial diseases
[show]
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Hydrolases: acid anhydride hydrolases (EC 3.6)
3.6.1
Pyrophosphatase
Inorganic
Thiamine
Apyrase
Thiamine-triphosphatase
3.6.2
Adenylylsulfatase
Phosphoadenylylsulfatase
3.6.3-4: ATPase
3.6.3
Cu++ (3.6.3.4)
Menkes/ATP7A
Wilson/ATP7B
Ca+ (3.6.3.8)
SERCA
ATP2A1
ATP2A2
ATP2A3
Plasma membrane
ATP2B1
ATP2B2
ATP2B3
ATP2B4
SPCA
ATP2C1
ATP2C2
Na+/K+ (3.6.3.9)
ATP1A1
ATP1A2
ATP1A3
ATP1A4
ATP1B1
ATP1B2
ATP1B3
ATP1B4
H+/K+ (3.6.3.10)
ATP4A
Other P-type ATPase
ATP8B1
ATP10A
ATP11B
ATP12A
ATP13A2
ATP13A3
3.6.4
Dynein
Kinesin
Myosin
Katanin
3.6.5: GTPase
3.6.5.1: Heterotrimeric G protein
Gαs
Gαi
GNAI1
GNAI2
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Gαq/11
GNAQ
GNA11
Gα12/13
GNA12
GNA13
Transducin
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Gustducin
GNAT3
3.6.5.2: Small GTPase > Ras superfamily
Rho family of GTPases: Cdc42
CDC42
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RhoUV
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2
3
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1
2
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C
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1
2
3
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other: Ras
HRAS
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NRAS
Rab
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RAB27
Arf
ARF6
SAR1B
ARL13B
ARL6
Ran
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RGK
3.6.5.3: Protein-synthesizing GTPase
Prokaryotic
IF-2
EF-Tu
EF-G
Eukaryotic
3.6.5.5-6: Polymerization motors
Dynamin
Tubulin
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Molecular and Cellular Biology portal
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History of laptops
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Alan Kay with his 1972 "Dynabook" prototype (photo: 5 November 2008 in Mountain View, California)
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The history of laptops describes the efforts in the 1970 s and 1980 s to build small, portable personal computers that combine the components, inputs, outputs and capabilities of a desktop computer in a small chassis. Before laptop/notebook computers were technically feasible, similar ideas had been proposed, most notably Alan Kay's Dynabook concept, developed at Xerox PARC in the early 1970 s. One of the first reasonably portable computers was the Xerox NoteTaker, again developed at Xerox PARC, in 1976. However, only 10 prototypes were built.
Contents
[hide]
1 Portal R2E CCMC
2 Osborne 1
3 Bondwell 2
4 Compaq Portable
5 Epson HX-20
6 Grid Compass
7 Dulmont Magnum/Kookaburra
8 Ampere
9 Tandy Model 100
10 Sharp and Gavilan
11 Kyotronic 85
12 Commodore SX-64
13 Kaypro 2000
14 IBM PC Convertible
15 Toshiba T1100, T1000, and T1200
16 US Air Force
17 Hewlett-Packard Vectra Portable CS
18 Cambridge Z88
19 Compaq SLT/286
20 NEC UltraLite
21 Apple
21.1 Macintosh Portable
21.2 Powerbook
22 IBM RS/6000 N40
23 Windows 95 operating system
24 Intel Pentium processor
25 Improved technology
26 Netbooks
27 Smartbooks
28 See also
29 References
30 External links
31 Further reading
Portal R2E CCMC[edit]
R2E CCMC Portal laptop
The portable micro computer the “Portal" of the French company R2E Micral CCMC officially appeared in September 1980 at the Sicob show in Paris. The Portal was a portable microcomputer designed and marketed by the studies and developments department of the French firm R2E Micral in 1980 at the request of the company CCMC specializing in payroll and accounting. It was based on an Intel 8085 processor, 8-bit, clocked at 2 MHz. It was equipped with a central 64K byte Ram, a keyboard with 58 alpha numeric keys and 11 numeric keys (separate blocks), a 32-character screen, a floppy disk: capacity = 140 000 characters, of a thermal printer: speed = 28 characters / second, an asynchronous channel, a synchronous channel, a 220V power supply. Designed for an operating temperature of 15 ° C to 35 ° C, it weighed 12 kg and its dimensions were 45cm x 45cm x 15cm. It provided total mobility. Its operating system was Prologue.
Osborne 1[edit]
An opened Osborne 1 computer, ready for use. The keyboard sits on the inside of the lid.
The Osborne 1 is considered the first true mobile computer by most historians. Adam Osborne founded Osborne Computer and formed the Osborne 1 in 1981. The Osborne 1 had a five-inch screen, incorporating a modem port, two 5 1/4 floppy drives, a big collection of bundled software applications, and a battery pack. The computer company was a failure and did not last for very long. Although it was large and heavy compared to today's laptops, with a tiny 5" CRT monitor, it had a near-revolutionary impact on business, as professionals were able to take their computer and data with them for the first time. This and other "luggables" were inspired by what was probably the first portable computer, the Xerox NoteTaker. The Osborne was about the size of a portable sewing machine, and could be carried on commercial aircraft. The Osborne 1 weighed close to 11 kg and was priced at $1795.
Bondwell 2[edit]
Although it wasn't released until 1985, well after the decline of CP/M as a major operating system, the Bondwell 2 is one of only a handful of CP/M laptops. It used a Z-80 CPU running at 4 MHz, had 64 KBs of RAM, and a 3.5" floppy disk drive built in, which was unusual for CP/M laptops. The flip-up LCD display's resolution was 640x200 pixels. Bondwell 2 also included MicroPro's complete line of CP/M software, including WordStar. The Bondwell 2 was capable of displaying bitmapped graphics. The price of the Bondwell 2 was listed at $995.
Compaq Portable[edit]
The Compaq Portable was the first portable computer created in 1982 which one of the first 100% IBM PC compatible systems. The first shipment was in March 1983 and was priced at $2,995. The Compaq Portable folded up into a luggable case the size of a portable sewing machine. The third model of this development, Compaq Portable II, featured high resolution graphics on its tube display. It was the first portable computer ready to be used on the shop floor, and for CAD and diagram display. It established Compaq as a major brand on the market.
Epson HX-20[edit]
Epson HX-20 (1981)
Epson L3s (~1990?)
Another significant machine announced in 1981, although first sold widely in 1983, was the Epson HX-20. A simple handheld computer, it featured a full-transit 68-key keyboard, rechargeable nickel-cadmium batteries, a small (120×32-pixel) dot-matrix LCD with 4 lines of text, 20 characters per line text mode, a 24 column dot matrix printer, a Microsoft BASIC interpreter, and 16 KB of RAM (expandable to 32 KB). Through at least the late 1980s Epson continued to release laptops such as the L3s.
Grid Compass[edit]
Main article: Grid Compass
The first laptop in the modern form was the Grid Compass 1101, designed by Bill Moggridge in 1979–80, and released in 1983. Enclosed in a magnesium case, it introduced the now familiar clamshell design, in which the flat display folded shut against the keyboard. The computer could be run from batteries, and was equipped with a 320×200-pixel electroluminescent display and 384 kilobyte bubble memory. It was not IBM-compatible, and its high price (US$8,000–10,000) limited it to specialized applications. However, it was used heavily by the U.S. military, and by NASA on the Space Shuttle during the 1980s. The GRiD's manufacturer subsequently earned significant returns on its patent rights as its innovations became commonplace. GRiD Systems Corp. was later bought by the Tandy (now RadioShack) Corporation.
Dulmont Magnum/Kookaburra[edit]
Main article: Dulmont Magnum
Another contender for the first true laptop was the Dulmont Magnum, designed Barry Wilkinson and Terry Crews Engineering Manager at Dulmison in 1982 and released in Australia in 1983 It included an 80×8 display in a lid that closed against the keyboard. It was based on the MS-DOS operating system and applications stored in ROM (A:) and also supported removable modules in expansion slots (B: and C:) that could be custom programmed EPROM or standard word processing and spreadsheet applications. However, the Magnum had no nonvolatile memory, but could suspend and retain memory in RAM, including a RAM Disk (D:). A separate expansion box provided dual 5.25-inch floppy or 10 MB hard disk storage. Dulmont was eventually taken over by Time Office Computers, who marketed the Magnum internationally in 16 and 25 line LCD versions, and also introduced the brandname Kookaburra to emphasize its Australian origins.
Ampere[edit]
The Ampere,[1] a laptop with a sleek clamshell design by Ryu Oosake, was made in 1983. It offered a MC68008 microprocessor dedicated to running an APL interpreter residing in ROM. It was released as the Ampere WS-1.[2]
Tandy Model 100[edit]
The TRS-80 Model 100 was an early portable computer introduced in 1983. It was one of the first notebook-style computers, featuring a keyboard and LCD, battery-powered, in a package roughly the size and shape of a notepad or a large book.
It was made by Kyocera, and originally sold in Japan as the Kyotronic 85. Although a slow seller for Kyocera, the rights to the machine were purchased by Tandy Corporation, and the computer was sold through Radio Shack stores in the United States and Canada as well as affiliated dealers in other countries, becoming one of the company's most popular models, with over 6,000,000 units sold worldwide. The Olivetti M-10 and the NEC PC-8201 and PC-8300 were also built on the same Kyocera platform.
Sharp and Gavilan[edit]
Two other noteworthy early laptops were the Sharp PC-5000 and the Gavilan SC, announced in 1983 but first sold in 1984. The Gavilan was notably the first computer to be marketed as a "laptop". It was also equipped with a pioneering touchpad-like pointing device, installed on a panel above the keyboard. Like the GRiD Compass, the Gavilan and the Sharp were housed in clamshell cases, but they were partly IBM-compatible, although primarily running their own system software. Both had LCDs, and could connect to optional external printers. The Dulmont Magnum, launched internationally in 1984, was an Australian portable similar in layout to the Gavilan, which used the Intel 80186 processor.[3]
Kyotronic 85[edit]
The year 1983 also saw the launch of what was probably the biggest-selling early laptop, the Kyocera Kyotronic 85. Owing much to the design of the previous Epson HX-20, and although at first a slow seller in Japan, it was quickly licensed by Tandy Corporation, Olivetti, and NEC, who recognised its potential and marketed it respectively as the TRS-80 Model 100 line (or Tandy 100), Olivetti M-10, and NEC PC-8201.[4] The machines ran on standard AA batteries. The Tandy's built-in programs, including a BASIC interpreter, a text editor, and a terminal program, were supplied by Microsoft, and was written in part by Bill Gates himself. The computer was not a clamshell, but provided a tiltable 8 line × 40-character LCD screen above a full-travel keyboard. With its internal modem, it was a highly portable communications terminal. Due to its portability, good battery life (and ease of replacement), reliability (it had no moving parts), and low price (as little as US$300), the model was highly regarded, becoming a favorite among journalists. It weighed less than 2 kg with dimensions of 30×21.5×4.5 centimeters (12×8½×1¾ in). Initial specifications included 8 kilobytes of RAM (expandable to 24 KB) and a 3 MHz processor. The machine was in fact about the size of a paper notebook, but the term had yet to come into use and it was generally described as a "portable" computer.
Commodore SX-64[edit]
The Commodore SX-64, also known as the Executive 64, or VIC-64 in Europe, was a portable, briefcase or suitcase-size "luggable" version of the popular Commodore 64 home computer, and was the first full-color portable computer.[5]
The SX-64 featured a built-in five-inch composite monitor and a built-in 1541 floppy drive. It weighed 20 pounds. The machine was carried by its sturdy handle, which doubled as an adjustable stand. It was announced in January 1983 and released a year later, at $995 USD.[6]
Kaypro 2000[edit]
Possibly the first commercial IBM-compatible laptop was the Kaypro 2000, introduced in 1985. With its brushed aluminum clamshell case, it was remarkably similar in design to modern laptops. It featured a 25 line by 80 character LCD, a detachable keyboard, and a pop-up 90 mm (3.5-inch) floppy drive.
IBM PC Convertible[edit]
Also among the first commercial IBM-compatible laptops was the IBM PC Convertible, introduced in 1986. It had a CGA-compatible LCD and 2 floppy drives. It weighed 13 pounds (5.9 kg).
Toshiba T1100, T1000, and T1200[edit]
Toshiba launched the Toshiba T1100 in 1985, and has subsequently described it as "the world's first mass-market laptop computer".[7] It did not have a hard drive, and ran entirely from floppy disks. The CPU was a 4.77 MHz Intel 80C88, a variation of the popular Intel 8088, and the display was a monochrome, text-only 640x200 LCD. It was followed in 1987 by the T1000 and T1200. Although limited floppy-based DOS machines, with the operating system stored in ROM, the Toshiba models were small and light enough to be carried in a backpack, and could be run from lead-acid batteries. They also introduced the now-standard "resume" feature to DOS-based machines: the computer could be paused between sessions without having to be restarted each time.
US Air Force[edit]
The first laptops successful on a large scale came in large part due to a Request For Proposal (RFP) by the U.S. Air Force in 1987. This contract would eventually lead to the purchase of over 200,000 laptops. Competition to supply this contract was fiercely contested and the major PC companies of the time; IBM Corporation, To shins, Compaq, NEC, and Zenith Data Systems (ZDS), rushed to develop laptops in an attempt to win this deal. ZDS, which had earlier won a landmark deal with the IRS for its Z-171, was awarded this contract for its SupersPort series. The SupersPort series was originally launched with an Intel 8086 processor, dual floppy disk drives, a backlit, blue and white STN LCD screen, and a NiCd battery pack. Later models featured an Intel 80286 processor and a 20 MB hard disk drive. On the strength of this deal, ZDS became the world's largest laptop supplier in 1987 and 1988. ZDS partnered with Tottori Sanyo in the design and manufacturing of these laptops. This relationship is notable because it was the first deal between a major brand and an Asian original equipment manufacturer.
Hewlett-Packard Vectra Portable CS[edit]
In 1987, HP released a portable version of their Vectra CS computer.[8] It had the classic laptop configuration (keyboard and monitor closes up clam-shell style in order to carry), however, it was very heavy and fairly large. It had a full-size keyboard (with separate numeric keypad) and a large amber LCD screen. While it was offered with dual 3.5-inch floppy disk drives, the most common configuration was a 20 MB hard drive and a single floppy drive. It was one of the first machines with a 1.44 MB density 3.5-inch disk drive.
Cambridge Z88[edit]
Another notable computer was the Cambridge Z88, designed by Clive Sinclair, introduced in 1988. About the size of an A4 sheet of paper as well, it ran on standard batteries, and contained basic spreadsheet, word processing, and communications programs. It anticipated the future miniaturization of the portable computer, and as a ROM-based machine with a small display, can – like the TRS-80 Model 100 – also be seen as a forerunner of the personal digital assistant.
Compaq SLT/286[edit]
By the end of the 1980s, laptop computers were becoming popular among business people. The COMPAQ SLT/286 debuted in October 1988, being the first battery-powered laptop to support an internal hard disk drive and a VGA compatible LCD screen. It weighed 14 lbs.[9]
NEC UltraLite[edit]
Main article: NEC UltraLite
The NEC UltraLite, released in mid-1989, was perhaps the first "notebook" computer, weighing just 2 kg (4.4 lbs). In lieu of a floppy or hard drive, it contained a 2 megabyte RAM drive, but this reduced its utility as well as its size. Although portable computers with clamshell LCD screens already existed before it, the Ultralite was the first computer in a notebook form-factor. It was significantly smaller than all portable computers that came before it. People can actually carry-it like a notebook and fold its clamshell LCD like a book cover over the rest of its body.
Apple[edit]
Macintosh Portable[edit]
The Macintosh Portable, Apple's first attempt at a battery-powered computer
The first Apple Computer machine designed to be used on the go was the 1989 Macintosh Portable (although an LCD screen had been an option for the transportable Apple IIc in 1984). Unlike the Compaq LTE laptop released earlier in the year the Macintosh Portable was actually a "luggable" not a laptop, but the Mac Portable was praised for its clear active matrix display and long battery life, but was a poor seller due to its bulk. In the absence of a true Apple laptop, several compatible machines such as the Outbound Laptop were available for Mac users; however, for copyright reasons, the user had to supply a set of Mac ROMs, which usually meant having to buy a new or used Macintosh as well.
Powerbook[edit]
The Apple PowerBook series, introduced in October 1991, pioneered changes that are now de facto standards on laptops, such as room for a palm rest, and the inclusion of a pointing device (a trackball). The following year, IBM released its ThinkPad 700C, featuring a similar design (though with a distinctive red TrackPoint pointing device).
Later PowerBooks featured optional color displays (PowerBook 165c, 1993), and first true touchpad (PowerBook 500 series, 1994), first 16-bit stereo audio, and first built-in Ethernet network adapter (PowerBook 500, 1994).
IBM RS/6000 N40[edit]
In 1994, IBM released the RS/6000 N40 laptop based on a PowerPC microprocessor running the AIX operating system, a variant of UNIX. It was manufactured by Tadpole Technology (now Tadpole Computer), who also manufactured laptops based on SPARC and Alpha microprocessors, the SPARCbook and ALPHAbook lines, respectively.
Windows 95 operating system[edit]
The summer of 1995 was a significant turning point in the history of notebook computing. In August of that year Microsoft introduced Windows 95. It was the first time that Microsoft had implemented the advanced power management specification with control in the operating system. Prior to this point each brand used custom BIOS, drivers and in some cases, ASICs, to optimize the battery life of its machines. This move by Microsoft was controversial in the eyes of notebook designers because it greatly reduced their ability to innovate; however, it did serve its role in simplifying and stabilizing certain aspects of notebook design.
Intel Pentium processor[edit]
Windows 95 also ushered in the importance of the CD-ROM drive in mobile computing, and initiated the shift to the Intel Pentium processor as the base platform for notebooks. The Gateway Solo was the first notebook introduced with a Pentium processor and a CD-ROM. Also featuring a removable hard disk drive and floppy drive, the Solo was the first three-spindle (optical, floppy, and hard disk drive) notebook computer, and was extremely successful within the consumer segment of the market. In roughly the same time period the Dell Latitude, Toshiba Satellite, and IBM ThinkPad were reaching great success with Pentium-based two-spindle (hard disk and floppy disk drive) systems directed toward the corporate market.
Improved technology[edit]
A 1997 Micron laptop
Early laptop displays were so primitive that PC Magazine in 1986 published an article discussing them with the headline "Is It On Yet?". It said of the accompanying montage of nine portable computers, "Pictured at the right are two screens and seven elongated smudges". The article stated that "LCD screens still look to many observers like Etch-a-Sketch toys, or gray chalk on a dirty blackboard", and predicted that until displays improved, "laptops will continue to be a niche rather than a mainstream direction".[10] As technology improved during the 1990s, the usefulness and popularity of laptops increased. Correspondingly prices went down. Several developments specific to laptops were quickly implemented, improving usability and performance. Among them were:
Improved battery technology. The heavy lead-acid batteries were replaced with lighter and more efficient technologies, first nickel cadmium or NiCd, then nickel metal hydride (NiMH) and then Lithium-ion battery and lithium polymer.
Power-saving processors. While laptops in 1991 were limited to the 80286 processor because of the energy demands of the more powerful 80386, the introduction of the Intel 386SL processor, designed for the specific power needs of laptops, marked the point at which laptop needs were included in CPU design. The 386SL integrated a 386SX core with a memory controller and this was paired with an I/O chip to create the SL chipset. It was more integrated than any previous solution although its cost was higher. It was heavily adopted by the major notebook brands of the time. Intel followed this with the 486SL chipset which used the same architecture. However, Intel had to abandon this design approach as it introduced its Pentium series. Early versions of the mobile Pentium required TAB mounting (also used in LCD manufacturing) and this initially limited the number of companies capable of supplying notebooks. However, Intel did eventually migrate to more standard chip packaging. One limitation of notebooks has always been the difficulty in upgrading the processor which is a common attribute of desktops. Intel did try to solve this problem with the introduction of the MMC for mobile computing. The MMC was a standard module upon which the CPU and external cache memory could sit. It gave the notebook buyer the potential to upgrade his CPU at a later date, eased the manufacturing process somewhat, and was also used in some cases to skirt U.S. import duties as the CPU could be added to the chassis after it arrived in the U.S. Intel stuck with MMC for a few generations but ultimately could not maintain the appropriate speed and data integrity to the memory subsystem through the MMC connector. A more specialized power saving CPU variant for laptops is the PowerPC 603 family.[11] Derived from IBM's 601 series for laptops (while the 604 branch was for desktops), it found itself used on many low end Apple desktops before it was wildly used in laptops, starting with PowerBook models 5300, 2400, 500 upgrades. What started out as a laptop processor was eventually used across all platforms in its follow up PPC 750.
Improved Liquid-crystal displays, in particular active-matrix TFT (Thin-film transistor) LCD technology. Early laptop screens were black and white, blue and white, or grayscale, STN (Super Twist Nematic) passive-matrix LCDs prone to heavy shadows, ghosting and blurry movement (some portable computer screens were sharper monochrome plasma displays, but these drew too much current to be powered by batteries). Color STN screens were used for some time although their viewing quality was poor. By about 1991, two new color LCD technologies hit the mainstream market in a big way; Dual STN and TFT. The Dual STN screens solved many of the viewing problems of STN at a very affordable price and the TFT screens offered excellent viewing quality although initially at a steep price. DSTN continued to offer a significant cost advantage over TFT until the mid-90s before the cost delta dropped to the point that DSTN was no longer used in notebooks. Improvements in production technology meant displays became larger, sharper, had higher native resolutions, faster response time and could display color with great accuracy, making them an acceptable substitute for a traditional CRT monitor.
OLPC XO-1 laptop in Ebook-Mode.
Improved storage technology. Early laptops and portables had only floppy disk drives. As thin, high-capacity hard disk drives with higher reliability and shock resistance and lower power consumption became available, users could store their work on laptop computers and take it with them. The 3.5" HDD was created initially as a response to the needs of notebook designers that needed smaller, lower power consumption products. With continuing pressure to shrink the notebook size even further, the 2.5" HDD was introduced. One Laptop Per Child (OLPC) and other new laptops use Flash RAM (non volatile, non mechanical memory device) instead of the mechanical hard disk.
Improved connectivity. Internal modems and standard serial, parallel, and PS/2 ports on IBM PC-compatible laptops made it easier to work away from home; the addition of network adapters and, from 1997, USB, as well as, from 1999, Wi-Fi, made laptops as easy to use with peripherals as a desktop computer. Many newer laptops are also available with built-in 3G Broadband wireless modems.
Other peripherals may include:
an integrated video camera for video communication
a fingerprint sensor for implementing a restriction of access to a sensitive data or the computer itself.
Netbooks[edit]
Main article: Netbook
In June 2007, Asus announced the Eee PC 701 to be released in October, a small lightweight x86 Celeron-M ULV 353 powered laptop with 4 GB SDHC disk and a 7-inch screen.[12] Despite previous attempts to launch small lightweight computers such as ultra-portable PC, the Eee was the first success story largely due to its low cost, small size, low weight and versatility. The term 'Netbook' was later dubbed by Intel. Asus then extended the Eee line with models with features such as a 9-inch screen and other brands, including Acer, MSI and Dell followed suit with similar devices, often built on the fledgling low-power Intel Atom processor architecture.
Smartbooks[edit]
Main article: Smartbook
In 2009, Qualcomm introduced a new term "smartbook", which stands for a hybrid device between smartphone and laptop.[13]
See also[edit]
Timeline of portable computers
Portal laptop computer
References[edit]
Jump up ^ Bob Armstrong, http://cosy.com/language/cosyhard/cosyhard.htm
Jump up ^ Japanese PCs (1984) (13:13), Computer Chronicles
Jump up ^ "OLD-COMPUTERS.COM : The Museum". Retrieved 11 August 2015.
Jump up ^ See TRS-80 Model 100 / 102 at old-computers.com
Jump up ^ "Commodore SX-64 Portable". Retrieved 11 August 2015.
Jump up ^ "Commodore SX-64 portable computer". Retrieved 11 August 2015.
Jump up ^ "World's First Laptop PC". Toshiba Science Museum. Retrieved 21 May 2017.
Jump up ^ "Business Desktops/Calcs". HP Computer Museum. Retrieved 7 April 2014.
Jump up ^ Lewis, Peter H. (23 October 1988). "THE EXECUTIVE COMPUTER; Compaq Finally Makes a Laptop". The New York Times.
Jump up ^ Somerson, Paul (July 1986). "Is It On Yet?". PC. pp. 122–123. Retrieved 9 January 2015.
Jump up ^ "IBM Power Systems archived white papers". Retrieved 11 August 2015.
Jump up ^ Paul Miller. "Asus' new Eee PC 701 joins the laptop-lite fray with a bang". Engadget. AOL. Retrieved 11 August 2015.
Jump up ^ Hachman, Mark (1 June 2009). "Qualcomm Shows Off Snapdragon Smartbooks". PC Magazine.
External links[edit]
"Laptops, 360 degree models", Russian Vintage Laptop Museum (museum)
Further reading[edit]
Wilson, James E. (2006). Vintage Laptop Computers: First Decade: 1980–89. Outskirts Press. p. 132. ISBN 978-1-59800-489-2.
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