Table 1.
P-type ATPases included in the
phylogenetic analysis
Number useda | Accession numberb | Organism | Specificityc | Name |
---|---|---|---|---|
Type IA ATPases | ||||
20 | P03960 | Escherichia coli | K+ | kdpB / f682 |
21 | Z92539 |
Mycobacterium tuberculosis | kdpB | |
22 | D90910 | Synechocystis PCC6803 | kdpB / slr1729 | |
Type IB ATPases | ||||
23 | U78967 | Lactococcus lactis | cadA | |
24 | L28104 | Listeria monocytogenes | Cd2+ | cadA |
25 | P30336 | Bacillus firmus | cadA | |
26 | P37386 | Staphylococcus aureus | cadD | |
27 | D64005 | S. PCC6803 | cadA_2 / slr0798 | |
28 | D64005 | S. PCC6803 | cadA_1 / slr0797 | |
29 | L46864 | Helicobacter pylori | ||
30 | Q10866 | M. tuberculosis | MTCY39.27 | |
31 | P37617 | E. coli | yhh0 / o732 | |
32 | Z92771 | M. tuberculosis | MTCY71.10 | |
33 | P77871 | H. pylori | Cu2+ | copA |
34 | U32715 | Haemophilus influenzae | hi290 | |
35 | Q59385 | E. coli | f834 | |
36 | P05425 | Enterococcus hirae | Cu2+ | copB |
37 | P37385 | Synechococcus PCC7942 | Cu2+ | synA |
38 | D90915 | S. PCC6803 | slr1950 | |
39 | Z79700 | M. tuberculosis | MTCY10D7.05c | |
40 | P46840 | Mycobacterium leprae | ctpB | |
41 | Q10876 | M. tuberculosis | ctpA / MTCY251.11 | |
42 | Q10877 | M. tuberculosis | ctpB / MTCY251.22c | |
43 | P46839 | M. leprae | ctpA | |
44 | U42410 | Proteus mirabilis | ||
45 | D90904 | S. PCC6803 | sll1920 | |
46 | P37279 | S. PCC7942 | Cu2+ | pacS |
47 | U15554 | L. monocytogenes | Cu2+ | ctpA |
48 | P38995 | Saccharomyces cerevisiae | Cu2+ | CCC2 / ydr270w |
49 | Q04656 | Homo sapiens | Cu2+ | ATP7A / MNK |
50 | U08344 | Rattus norvegicus | Cu2+ | ATP7B |
51 | P35670 | H. sapiens | Cu2+ | ATP7B / WND |
52 | P38360 | S. cerevisiae | PCA1 / ybr295w | |
53 | X95634 | Bradyrhizobium japonicum | fixI | |
54 | P18398 | Rhizobium meliloti | fixI | |
55 | U16659 | Unknown d | hra2 | |
56 | P32113 | E. hirae | Cu2+ | copA |
57 | U16658 | Unknown d | hra1 | |
Type IIA ATPases | ||||
130 | Q10860 | M. tuberculosis | MTCY39.21c | |
131 | P37367 | S. PCC6803 | pma1 / sll1614 | |
132 | M93017 | R. norvegicus | ||
133 | P13586 | S. cerevisiae | Ca2+ | PMR1 |
134 | U65981 | Cryptosporidium parvum | CppA-E1 | |
135 | U39298 | Plasmodium falciparum | PfATPase4 | |
136 | U65066 | Trichomonas vaginalis | TVCA1 | |
137 | P54209 | Dunaliella bioculata | CA1 | |
138 | U70540 | Leishmania mexicana amazonensis | ||
139 | P35315 | Trypanosoma brucei | TBA1 | |
140 | L40328 | Schistosoma mansoni | SMA1 | |
141 | P16615 | H. sapiens | Ca2+ | ATP2A2 / SERCA2 |
142 | U65228 | Makaira nigricans | SERCA1A | |
143 | X63009 | Rana esculenta | Ca2+ | |
144 | M99223 | R. norvegicus | Ca2+ | |
145 | P18596 | R. norvegicus | Ca2+ | ATP2A3 / SERCA3 |
146 | P22700 | Drosophila melanogaster | DRSERCA / CA-60A | |
147 | P35316 | Artemia franciscana | ||
148 | Q08853 | P. falciparum | ||
149 | X55197 | Plasmodium yoelii | YEL6 | |
150 | M96324 | Lycopersicon esculentum | LCA1 | |
151 | U82966 | Oryza sativa | ||
152 | U93845 | Arabidopsis thaliana | ACA3 | |
153 | D64005 | S. PCC6803 | pacL / sll0672 | |
154 | P37278 | S. PCC7942 | pacL | |
155 | D90911 | S. PCC6803 | pacL / slr0822 | |
Type IIB ATPases | ||||
118 | U05880 | Paramecium tetraurelia | ||
119 | X99972 | Brassica oleracea | Ca2+ | BCA1 |
120 | L08468 | A. thaliana | PEA1 | |
121 | U20321 e | Entamoeba histolytica | ||
122 | U15408 | R. norvegicus | ||
123 | P23634 | H. sapiens | Ca2+ | ATP2B4 / PMCA4 |
124 | P20020 | H. sapiens | Ca2+ | ATP2B1 / PMCA1 |
125 | Q16720 | H. sapiens | Ca2+ | ATP2B3 / PMCA3 |
126 | Q01814 | H. sapiens | Ca2+ | ATP2B2 / PMCA2 |
127 | Z68221 | Caenorhabditis elegans | ||
128 | P54678 | Dictyostelium discoideum | patA | |
129 | P38929 | S. cerevisiae | Ca2+ | PMC1 / ygl006w |
Type IIC ATPases | ||||
97 | U70316 f | D. discoideum | IonA | |
98 | Z81457 g | C. elegans | ||
99 | P35317 | Hydra vulgaris | Na+/K+ | |
100 | P28774 | A. franciscana | ||
101 | U18546 | C. elegans | Na+/K+ | eat-6 |
102 | P13607 | D. melanogaster | Na+/K+ | NA-P |
103 | S66043 | Ctenocephalides felis | ||
104 | P05023 | H. sapiens | Na+/K+ | ATP1A1 / NaK1 |
105 | P05025 | Torpedo californica | Na+/K+ | ATP1A1 / NaK1 |
106 | P13637 | H. sapiens | Na+/K+ | ATP1A3 / NaK3 |
107 | Q92030 | Anguilla anguilla | ||
108 | P25489 | Catostomus commersoni | ||
109 | P50993 | H. sapiens | Na+/K+ | ATP1A2 / NaK2 |
110 | U15176 | R. norvegicus | ATP1A4 / NaK4 | |
111 | P17326 | A. franciscana | Na+/K+ | |
112 | P20648 | H. sapiens | H+/K+ | ATP4A |
113 | U17249 | Xenopus laevis | H+/K+ | |
114 | D21854 | Cavia cobaya | ||
115 | P54707 | H. sapiens | H+/K+ | ATP1AL1 |
116 | P54708 | R. norvegicus | H+/K+ | ATP1AL1 |
117 | Z25809 | Bufo marinus | H+/K+ | |
Type IID ATPases | ||||
93 | P22189 | Schizosaccharomyces pombe | Ca2+ | CTA3 |
94 | P13587 | S. cerevisiae | Na+ | ENA1 / ydr040c |
95 | D78567 | Zygosaccharomyces rouxii | Z-ENA1 | |
Type IIIA ATPases | ||||
61 | U67563 | Methanococcus jannaschii | mj1226 | |
62 | P54210 | Dunaliella acidophila | H+ | dha1 |
63 | P54211 | D. bioculata | PMA1 | |
64 | P54679 | D. discoideum | ||
65 | D88424 | Cyanidium caldarium | ||
66 | X76535 | Solanum tuberosum | PHA2 | |
67 | Q03194 | Nicotiana plumbaginifolia | PMA4 | |
68 | P19456 | A. thaliana | H+ | AHA2 |
69 | P20431 | A. thaliana | H+ | AHA3 |
70 | S79323 | Vicia faba | ||
71 | D45189 | Zostera marina | ZHA1 | |
72 | X85805 | Zea mays | MHA2 | |
73 | U84891 | Mesembryanthemum crystallinum | PMA | |
74 | X73676 | A. thaliana | AHA9 | |
75 | D31843 | O. sativa | OSA2 | |
76 | U09989 | Z. mays | MHA1 | |
77 | D10207 | O. sativa | OSA1 | |
78 | M27888 | N. plumbaginifolia | H+ | PMA2 |
79 | X85804 | Phaseolus vulgaris | BHA1 | |
80 | S74033 | A. thaliana | AHA10 | |
81 | P12522 | Leishmania donovani | ATPase1b | |
82 | D26472 | Heterosigma akashiwo | ||
83 | P24545 | Z. rouxii | H+ | |
84 | P28877 | Candida albicans | PMA1 | |
85 | P49380 | Kluyveromyces lactis | PMA1 | |
86 | P05030 | S. cerevisiae | H+ | PMA1 / ygl008c |
87 | P19657 | S. cerevisiae | H+ | PMA2 / ypl036w |
88 | U65004 | Pneumocystis carinii | PCA1 | |
89 | P28876 | S. pombe | H+ | PMA2 |
90 | P09627 | S. pombe | H+ | PMA1 |
91 | P07038 | Neurospora crassa | H+ | PMA1 |
92 | Q07421 | Histoplasma capsulatum | PMA1 | |
Type IIIB ATPases | ||||
58 | P22036 | Salmonella typhimurium | Mg2+ | mgtB |
59 | P39168 | E. coli | mgtA / o898 | |
60 | P36640 | S. typhimurium | Mg2+ | mgtA |
Type IV ATPases | ||||
8 | Z81078 | C. elegans | ||
9 | P40527 | S. cerevisiae | yil048w | |
10 | Q10309 | S. pombe | SPAC6C3.05C | |
11 | U28940 h | C. elegans | T24H7.5 | |
12 | U16955 | P. falciparum | PfATPase2 | |
13 | Q12674 | S. cerevisiae | ymr162c | |
14 | U51100 | Bos taurus | PL | |
15 | P39524 | S. cerevisiae | PL | DRS2 / yal026c |
16 | Q09891 | S. pombe | SPAC24B11.12C | |
17 | Q12675 | S. cerevisiae | ydr093w | |
18 | P32660 | S. cerevisiae | yer166w | |
Type V ATPases | ||||
1 | Q12697 | S. cerevisiae | yor291w | |
2 | Z70271 | C. elegans | W08D2.5 | |
3 | U41552 i | C. elegans | ||
4 | X65738 | P. falciparum | PfATPase1 | |
5 | Z83217 | C. elegans | ||
6 | P39986 | S. cerevisiae | yel031w | |
7 | U41063 | Tetrahymena thermophila | TPA9 | |
Others | ||||
158 | Q10900 | M. tuberculosis | MTCY251.26C | |
96 | D90905 | S. PCC6803 | pacL / sll1076 | |
157 | P47317 | Mycoplasma genitalium | pacL / mg071 | |
159 | Z84724 | M. tuberculosis | MTCY22G10.22c | |
19 | Z95210 | M. tuberculosis | MTCY21C12.02 | |
156 | P78036 | Mycoplasma pneumoniae | mgtA / G07_872V |
a | The number and the groupings refer to the sequence numbers and groups used in all figures |
b | The accession numbers starting with a Q or P are from the SWISS-PROT database. All other accession numbers are from the EMBL database |
c | The ion specificity of the P-type ATPases is shown, when it has been experimentally shown. PL: phospholipids |
d | The sequences were found in a human cDNA library. They were suggested to originate from E. coli, but could not be identified in the genome |
e | The N-terminus is missing. Sequence alignments with other type IIB ATPases indicate that the sequence is probably 80-130 amino acids longer |
f | Both termini are missing. Sequence alignments with other type IIC ATPases indicate that the sequence is probably 30-45 amino acids longer |
g | Only genomic sequence data are available and no proposed translation. The conserved segments have therefore been extracted directly from the genomic clone |
h | The proposed translation of the genomic DNA sequence lacks the region around the conserved segment D (Fig. 2). The sequence can be corrected by inserting two additional exons from nucleotide 9,394 to 9,573 and from nucleotide 10,127 to 10,223. An expressed sequence tag (D68066) covering this area of the protein further supports the corrections |
i | The proposed translation of the genomic DNA sequence lacks several of the conserved regions. The conserved segments have therefore been extracted directly from the genomic clone |
Table 2.
P-type ATPases, which are excluded from the phylogenetic analysis.
Accession number a | Organism (Open reading frame) |
ATPase type |
>90% identity with seq. no.b |
Ion speci- ficityc |
Name |
---|---|---|---|---|---|
P20021 | Staphylococcus aureus | IB | 26 | Cd2+ | cadA |
Q59467 | Helicobacter pylori | IB | 33 | copA2 | |
P49015 | Cricetulus griseus | IB | 49 | ATP7A | |
U03434 | Mus musculus | IB | 49 | Cu2+ | ATP7A / MNK |
U59245 | Rattus norvegicus | IB | 49 | Cu2+ | ATP7A / MNK |
U38477 | M. musculus | IB | 51 | ATP7B / WND | |
Z69881 | Homo sapiens | IIA | 145 | Ca2+ | ATP2A3 / SERCA3 |
P13585 | Gallus gallus | IIA | 144 | Ca2+ | ATP2A1 / SERCA1 |
P04191 | Oryctolagus cuniculus | IIA | 144 | Ca2+ | ATP2A1 / SERCA1 |
P11719 | O. cuniculus | IIA | 144 | Ca2+ | ATP2A1 / SERCA1 |
P16614 | H. sapiens | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
Q00779 | Felis silvestris catus | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
M66385 | G. gallus | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
U49393 | M. musculus | IIA | 141 | Ca2+ | ATP2A3 / SERCA3 |
P04192 | O. cuniculus | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
P20647 | O. cuniculus | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
P11508 | R. norvegicus | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
P11507 | R. norvegicus | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
P11606 | Sus scrofa | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
P11607 | S. scrofa | IIA | 141 | Ca2+ | ATP2A2 / SERCA2 |
Q00804 | O. cuniculus | IIB | 124 | Ca2+ | ATP2B1 / PMCA1 |
P11505 | R. norvegicus | IIB | 124 | Ca2+ | ATP2B1 / PMCA1 |
P23220 | S. scrofa | IIB | 124 | Ca2+ | ATP2B1 / PMCA1 |
J05087 | R. norvegicus | IIB | 125 | Ca2+ | ATP2B3 / PMCA3 |
P11506 | R. norvegicus | IIB | 126 | Ca2+ | ATP2B2 / PMCA2 |
P30714 | Bufo marinus | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P50997 | Canis familiaris | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P18907 | Equus caballus | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P09572 | G. gallus | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P04074 | Ovis aries | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P06685 | R. norvegicus | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P05024 | S. scrofa | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P18874 | S. scrofa | IIC | 104 | Na+/K+ | ATP1A3 / NaK3 |
U10108 | Xenopus laevis | IIC | 104 | Na+/K+ | ATP1A1 / NaK1 |
P24798 | G. gallus | IIC | 106 | Na+/K+ | ATP1A3 / NaK3 |
P06687 | R. norvegicus | IIC | 106 | Na+/K+ | ATP1A3 / NaK3 |
P24797 | G. gallus | IIC | 109 | Na+/K+ | ATP1A2 / NaK2 |
P06686 | R. norvegicus | IIC | 109 | Na+/K+ | ATP1A2 / NaK2 |
P50996 | C. familiaris | IIC | 112 | H+/K+ | ATP4A |
U17282 | M. musculus | IIC | 112 | H+/K+ | |
P27112 | O. cuniculus | IIC | 112 | H+/K+ | ATP4A |
P09626 | R. norvegicus | IIC | 112 | H+/K+ | ATP4A |
P19156 | S. scrofa | IIC | 112 | H+/K+ | ATP4A |
Q01896 | Saccharomyces cerevisiae | IID | 94 | ENA2 / ydr039c | |
Z74334 | S. cerevisiae | IID | 94 | ENA5 / ydr038c | |
U72148 | Lycopersicon esculentum | IIIA | 67 | LHA4 | |
P20649 | Arabidopsis thaliana | IIIA | 68 | H+ | AHA1 |
P22180 | L. esculentum | IIIA | 78 | LHA1 | |
Q08435 | Nicotiana plumbaginifolia | IIIA | 78 | PMA1 | |
Q08436 | N. plumbaginifolia | IIIA | 78 | PMA3 | |
X76536 | Solanum tuberosum | IIIA | 78 | PHA1 | |
P11718 | Leishmania donovani | IIIA | 81 | ATPase1a | |
U75321 | M. musculus | IV | 14 | PL |
a | The accession numbers starting with a Q or P are from the SWISS-PROT database. All other accession numbers are from the EMBL database |
b | The sequence numbers refer to the numbers defined in Table 1 |
c | The ion specificity of the P-type ATPases is shown, when it has been experimentally shown |
Table 3.
Distribution of cloned P-type ATPases according to phylogenetic relationship a
ATPase type Organism |
Total |
IA K |
IB HM |
IIA Ca |
IIB Ca |
IIC Na/K |
IIC H/K |
IID Na |
IIIA H |
IIIB Mg |
IV PL |
V NAS |
Othersb |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Bacteria | |||||||||||||
Escherichia coli | 4 | 1 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 |
Haemophilus influenzae | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Mycobacterium tuberculosis | 10 | 1 | 5 | 1 | - | - | - | - | - | - | - | - | 3 |
Mycoplasma genitalium | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
Mycoplasma pneumoniae | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
Synechocystis PCC6803 | 9 | 1 | 4 | 3 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
Archaea | |||||||||||||
Methanococcus jannaschii | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
Eukarya | |||||||||||||
protozoa | |||||||||||||
Plasmodium falciparum | 4 | - | - | 2 | - | - | - | - | - | - | 1 | 1 | - |
fungi | |||||||||||||
Saccharomyces cerevisiae | 16 | 0 | 2 | 1 | 1 | 0 | 0 | 3(2) | 2 | 0 | 5 | 2 | 0 |
Schizosaccharomyces pombe | 5 | - | - | - | - | - | - | 1 | 2 | - | 2 | - | - |
plants | |||||||||||||
Arabidopsis thaliana | 8 | - | - | 1 | 1 | - | - | - | 6(2) | - | - | - | - |
animals | |||||||||||||
Caenorhabditis elegans | 8 | - | - | - | 1 | 2 | - | - | - | - | 3 | 2 | - |
Rattus norvegicus | 14c | - | 2 | 3(2c) | 4 | 3 | 2 | - | - | - | - | - | - |
Homo sapiens | 17c | - | 2 | 5(2c) | 4(3) | 4(3) | 2(1) | - | - | - | - | - | - |
a | The categories in the table correspond to the branches found in the phylogenetic tree in Fig. 6. The figures in parentheses give the number of closely related ATPases from the groups which are not included in the phylogenetic analysis. Abbreviations: HM: heavy metals; PL: phospholipids; NAS: no assigned specificity |
b | This group contains bacterial ATPases, for which the ion specificity is not known and which do not group into any of the major branches indicated in Fig. 6 |
c | Two of the proteins in each of these groups originate from one gene by different splicing |
Table 4.
Expressed sequence tags in three
eukaryotic organisms with similarity to P-type ATPases
a
Arabidopsis thaliana | ||||||
---|---|---|---|---|---|---|
IB HM 3-6 b | IIA Ca 3 | IIB Ca 4-8 | IIC Na/K+H/K - | III H 5-6 | IV PL 2-3 | V NAS 1 |
N96835 T43098 T43560 T46552 W43689 Z33730 |
R64945 Z37285 Seq. 152 [2 ;-] |
H37540 N65090 N96705 R65015 (T46834) c R84090 (T43417) T41649 (T41650) Z25618 Seq .120 d [2;-] e |
|
F13883 N38709 N96803 P20649 [24;3] Seq. 68 [32;10] Seq. 69 [2;0] Seq. 74 [0;-] Seq. 80 [0;-] |
N96084 T45045 T88130 |
R65211 (R65219) |
Caenorhabditis elegans | ||||||
IB HM 1 | IIA Ca 3-4 | IIB Ca 1 | IIC Na/K+H/K 5-7 | III H - | IV PL 5 | V NAS 4-5 |
D36097 (D37525; D67367) |
C11879 C13112 (C13651; C13948; D34934; D68093; D68790; D70495; D74177; D74205; D75938) C13654 D67762 (M89247) |
Seq. 127 [13;0] |
D27090 D27840 (D64489) D32312 (D64530) D35686 (D68187) D72536 (Z14852) Seq. 98 [4;0] Seq. 101 [13;-] |
|
D27733 (D67512) D34761 (C12815; D34857; D36536; D68551) D35311 (D36265; D36990; D37399) Seq. 8 [4;0] Seq. 11 [16;0] |
C12075 (D37337) D35844 D69289 Seq. 2 [6;0] Seq. 3 [0;0] Seq. 5 [7;0] |
Homo sapiens | ||||||
IB HM 3 | IIA Ca 5-10 | IIB Ca 4 | IIC Na/K+H/K 5-8 | III H - | IV PL 3-6 | V NAS 2-4 |
Z16260 Seq. 49 [4;1] Seq. 51 [2;-] |
F00478 F00875 F01249 (F18895) H14052 H82605 N31343 (N56068) N40047 Seq. 141 f [58;24] P16614 f [3;0] Z69881 [5;0] |
Seq. 123 [6;-] Seq. 124 [25;9] Seq. 125 [0;-] Seq. 126 [6;1] |
R88986 T19577 W22378 W22714 Seq. 104 [51;12] Seq. 106 [21;0] Seq. 109 [32;13] Seq. 112 [4;0] Seq. 115 [0;-] |
|
D44622 H13657 (R17452; R50093; R20441) H26328 H30693 (R51412; R87856) H97619 (R81633; T81806) W26763 |
T19151 H16899 (R12888; R48894; R86015) R50051 T72002 |
a | The expressed sequence tags (ESTs) are placed in the groups of ATPases to which they show the highest similarity on amino acid basis |
b | The figures give an estimate of how many independent genes the ESTs are representing. The lower figure is based on the differences in similarity among the ESTs to the known P-type ATPases and the higher figure is the maximal possible number of independent genes with the available ESTs. |
c | EST clones placed in parenthesis depict EST clones overlapping with the EST clone placed in the above position. Abbreviations are as in Table 3 |
d | The accession number or sequence number show an already cloned P-type ATPase. The sequence numbers are as in Table 1. Database accession numbers are given for sequences not included in the phylogenetic analysis |
e | The figures in square brackets are the total number of ESTs aligning to the mRNA and the ESTs aligning to the 3' untranslated region of the mRNA respectively. A dash indicates that the 3' untranslated region was not available |
f | Seq. 141 and P16614 are both derived from the same gene as two different splicing products. The ESTs which are identical to both sequences are shown under seq. 141 |