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. 2017 Mar 21;6(1):12. doi: 10.3390/pathogens6010012

Table 1.

Description of the 97 putative protein kinases (PKs) identified in the kinome of Sarcocystis neurona. The putative PKs could be classified into eight groups. The amino acid coordinates of the conserved PK domains in the protein sequences and the PK homologies to other apicomplexan PKs are shown in Columns 7–12.

Description of the Putative Protein Kinases (PKs) in the Genome of S. neurona Description of Protein Kinase (PK) Homologies (BLASTp)
Protein ID a Sequence Annotations; Description b Family; (Subfamily) c Length (aa) pI MW (kDa) PK Domain Coordinates Sequence Name; (Apicomplexan) Bit Score E-Value Identity (%) Accession Number
1. Kinase Group AGC (Protein kinases A (PKA), G (PKG) and C (PKC) families)
SRCN_1312 AGC kinase 3-phosphoinositide dependent PK-1 (PDK1) 903 5.52 101.13 137–481 PDPK; (T. gondii RUB) 417 6.00 × 10−135 59 KFG61374.1
SRCN_3339 AGC kinase PKA 1428 8.85 147.52 1102–1417 Putative AGC kinase; (N. caninum L) 611 0.0 81 CEL65574.1
SRCN_4249 AGC kinase Nuclear dbf2-related (NDR) 152 8.91 17.47 6–141 Putative AGC kinase; (N. caninum L) 219 2.00 × 10−68 76 XP_003883757.1
SRCN_4518 PK G AGC kinase family member PKG Ciliate-E2 425 5.83 48.78 97–399 AGC kinase TgPKG1; (T. gondii ME49) 830 0.0 92 EPR61116.1
SRCN_3990 cAMP-dependent kinase CAMKL; (MELK) 1907 9.55 217.55 782–1634 cAMP-dependent protein kinase (T. gondii VEG) 75.9 1 × 10−12 30 ESS31194.1
SRCN_5165 cAMP-dependent PK, catalytic chain PKA 343 8.99 39.36 20–338 AGC kinase; (T. gondii ARI) 425 3.00 × 10−150 90 KYF43224.1
SRCN_4913 Putative PK PKD 2330 6.42 244.64 1093–1737 Putative PK; (E. tenella) 107 9.00 × 10−22 60 XP_013228294.1
SRCN_5430 AGC kinase Ribosomal protein S6 Kinases (RSK; (p70)) 1378 5.54 139.28 824–1344 AGC kinase; (T. gondii MAS) 223 3.00 × 10−60 59 KFH07588.1
SRCN_5610 cAMP-dependent PK, catalytic chain PKA 333 9.00 37.96 12–318 cAMP-dependent PK, catalytic subunit; (T. gondii ME49) 641 0.0 92 XP_002366464.1
2. Kinase Group calcium (Ca2+)-/calmodulin-regulated kinases (CAMK)
SRCN_1071 Ca2+-dependent kinase CAMK1 1495 9.47 152.66 1085–1401 Putative PK; (T. gondii VEG) 188 1.00 × 10−47 67 ESS31884.1
SRCN_2032 Putative PK Ciliate-C1 297 6.22 33.26 15–297 PK; (H. hammondi) 424 1.00 × 10−146 68 XP_008882026.1
SRCN_2165 Ca2+-dependent kinase CDPK2B CDPK 692 7.33 75.65 101–401 Ca2+-dependent PK CDPK2A; (T. gondii ARI) 674 0.0 90 KYF44522.1
SRCN_2257 Histone kinase CAMKL; (AMP-activated protein kinase (AMPK)) 1800 8.98 187.71 1159–1448 Putative CAM kinase, SNF1 family; (E. acervulina) 377 7.00 × 10−105 64 XP_013252246.1
SRCN_2544 CAM SNF1 AMK1 family CAMKL;(AMPK-regulated kinase novel kinase (NUAK)) 333 5.71 37.85 62–333 CAM kinase, SNF1/AMK1 family ToxPK1; (N. caninum L) 480 9.00 × 10−167 79 XP_003882065.1
SRCN_2937 Ca2+-signaling kinase MARK CAMKL; (microtubule affinity regulating kinase (MARK)) 278 9.62 31.29 1–250 Putative Ca2+ signaling PK MARK; (T. gondii GT1) 356 5.00 × 10−124 72 EPR59053.1
SRCN_3011 Calmodulin-dependent PK (CAM) CDPK6 CDPK 1435 9.25 154.44 1238–1435 Cdpk kinase domain; (T. gondii) 179 6.00 × 10−49 75 3IS5_A
SRCN_3314 A Chain crystal Structure of TgCDPK1 with inhibitor bound CDPK1 519 5.99 58.89 37–335 Calmodulin-domain PK 1; (T. gondii) 536 0.0 97 3MA6_A
SRCN_3583 Ca2+-dependent kinase CDPK5 CDPK 454 6.09 50.38 35–308 Ca2+-dependent PK CDPK5; (T. gondii ARI) 776 0.0 89 KYF43137.1
SRCN_3701 Ca2+-dependent kinase CDPK3 CDPK 560 5.91 62.07 77–362 Ca2+-dependent Kinase; (T. gondii) 493 1.00 × 10−172 87 3DXN_A
SRCN_4076 CAM CDPK family CDPK 1701 5.93 181.42 1079–1663 CAM kinase, CDPK family; (H. hammondi) 239 2.00 × 10−62 69 XP_008884897.1
SRCN_4093 PK CAMKL; (AMP-activated protein kinase (AMPK)) 1155 9.02 118.33 1–261 Putative atypical MEK-related kinase; (N. caninum L) 222 1.00 × 10−59 70 XP_003880869.1
SRCN_4390 Ca2+-dependent kinase CDPK2 CDPK 790 6.24 85.47 280–556 Ca2+-dependent PK, related; (N. caninum L) 1130 0.0 74 XP_003884321.1
SRCN_4815 Histone kinase (partial) CAMKL; (AMP-activated protein kinase (AMPK)) 711 6.53 75.98 1–314 SNF1-related PK catalytic-α KIN10, 5 AMP-activated PK; (N. caninum L) 568 0.0 48 CEL67550.1
SRCN_5227 CAM CDPK CDPK8-like CDPK 2748 8.98 282.12 796–885 Putative CAM kinase, CDPK family; (N. caninum L) 117 3.00 × 10−24 59 XP_003881901.1
SRCN_5410 Calmodulin-dependent PK (CAM-SNF1 family) CAMK1 467 8.94 52.04 168–446 CAM kinase, SNF1 family; (H. hammondi) 431 5.00 × 10−134 78 XP_008883430.1
SRCN_5812 Ca2+-dependent kinase CDPK9 CDPK 760 8.37 84.23 254–573 Ca2+-dependent PK CDPK9; (H. hammondi) 1139 0.0 81 XP_008889286.1
SRCN_5948 Ca2+-dependent kinase CDPK8 CDPK 3298 7.11 345.85 208–860 EF-hand domain-containing protein; (T. gondii ME49) 114 1.00 × 10−23 58 XP_002368547.1
SRCN_6597 Ca2+ dependent kinase CDPK7 CAMK1 1374 9.09 138.28 365–623 PK-PH domain-containing protein; (T. gondii ME49) 813 0.0 80 XP_002366487.1
SRCN_6606 Ca2+-dependent kinase CDPK4 CDPK 1632 9.42 170.93 813–1236 Ca2+-dependent PK; (T. gondii) 731 0.0 58 CAD32376.2
3. Kinase Group casein kinase 1 (cell kinase 1)
SRCN_3445 Casein kinase I CK1-D 323 9.34 37.65 6–290 Casein kinase 1; (T. gondii ME49) 603 0.0 94 XP_002366683.1
SRCN_4587 Casein kinase I CK1-D 137 7.78 15.94 1–137 Casein kinase I; (H. hammondi) 226 3.00 × 10−71 81 XP_008883809.1
SRCN_4645 Casein kinase I CK1-D 229 9.51 25.66 38–229 Casein kinase I; (T. gondii GAB2-2007-GAL-DOM2) 259 1.00 × 10−86 79 KFG42638.1
4. Kinase Group CMGC (including cyclin-dependent kinases, mitogen-activated PKs, glycogen synthase kinases and CDK-like kinases)
SRCN_1104 Cyclin-dependent kinase family 5 Ca2+-dependent PK-L (CDKL) 372 9.23 42.71 1–318 Cyclin-dependent kinase family 5 protein; (H. hammondi) 490 2.00 × 10−173 76 XP_008884207.1
SRCN_1236 Cell-cycle-associated kinase (SRPK) Serine-arginine rich PK (SRPK) 2911 5.37 302.90 713–1837 PK; (T. gondii ME49) 476 9.00 × 10−141 76 XP_002369401.1
SRCN_1479 CMGC Lammer CLK 748 10.23 79.09 485–748 Cell-cycle-associated PK CLK; (T. gondii FOU) 288 2.00 × 10−81 74 KFG33061.1
SRCN_1611 CMGC Dual-specificity tyrosine-regulated kinase (DYRK) DYRK; (DYRKP) 1504 5.86 160.34 551–1498 Cell-cycle-associated PK DYRK; (T. gondii VEG) 223 2.00 × 10−57 63 ESS33160.1
SRCN_1731 Cell-cycle-associated kinase GSK Glycogen synthase kinase (GSK) 219 6.59 24.39 1–175 Cell-cycle-associated PK GSK; (H. hammondi) 330 2.00 × 10−112 82 XP_008887193.1
SRCN_1732 Cell-cycle-associated kinase GSK Glycogen synthase kinase (GSK) 203 10.78 20.72 86–203 CMGC kinase, GSK family TgPK3; (E. brunetti) 114 1.00 × 10−28 91 CDJ46527.1
SRCN_2759 Cell-cycle-associated kinase partial Ca2+-dependent PK (CDK); (CRK7) 1122 6.14 118.67 541–1122 Cell-cycle-associated PK CDK; (T. gondii VAND) 118 1.00 × 10−25 79 KFH12036.1
SRCN_2845 CMGC DYRK PRP4 kinase DYRK; (PRP4) 1665 9.76 177.08 1267–1596 Putative PK (CLK3); (P. malariae) 330 1.00 × 10−102 69 SBS85334.1
SRCN_3891 CMGC kinase DYRK; (DYRK2) 674 8.85 73.85 399–674 Putative CMGC kinase; (T. gondii ME49) 80.1 6.00 × 10−14 67 EPT25192.1
SRCN_4209 CMGC MAPK family (ERK) MAPK-1 Mitogen-activated PK (MAPK); (ERK)) 2361 6.73 247.71 94–754 CMGC, MAPK/ (ERK) TgMAPK-1; (E. brunetti) 137 1.00 × 10−30 74 CDJ49492.1
SRCN_4674 Cyclin-dependent kinase Ca2+-dependent PK (CDK); (CDK7) 138 7.80 15.50 1–138 Cyclin-dependent kinase; (T. gondii GT1) 108 7.00 × 10−27 58 EPR60430.1
SRCN_4801 Cell-cycle-associated kinase Ca2+-dependent PK (CDK); (CDK5) 300 6.08 34.33 1–289 CMGC kinase, CDK family TgPK2; (N. caninum L) 576 0.0 91 XP_003885801.1
SRCN_5365 Cell-cycle-associated kinase MAPK Mitogen-activated PK (MAPK; (ERK)) 417 6.77 48.32 7–363 Cell-cycle-associated PK MAPK; (H. hammondi) 823 0.0 93 XP_008886907.1
SRCN_6346 Cell-cycle-associated kinase CDK Ca2+-dependent PK (CDK); (CDK5) 690 9.55 80.90 208–603 Putative cell-cycle-associated PK CDK; (T. gondii ARI) 390 2.00 × 10−122 87 KYF45878.1
SRCN_6427 CMGC CK2 kinase Cell Kinase 2 (CK2) 1395 10.29 144.86 885–1356 CMGC kinase, CK2 family; (T. gondii MAS) 241 6.00 × 10−73 98 KFH07655.1
SRCN_6472 Cell-cycle-associated kinase ERK7 Mitogen-activated PK (MAPK; (ERK)) 983 9.28 104.95 7–317 Cell-cycle-associated PK ERK7; (T. gondii ARI) 647 0.0 81 KYF46268.1
SRCN_761 Cell-cycle-associated kinase Ca2+-dependent PK (CDK); (CDK7) 577 9.34 58.39 144–490 Cell-cycle-associated PK; (H. hammondi) 283 3.00 × 10−88 68 XP_008882409.1
SRCN_895 Cell-cycle-associated kinase Ca2+-dependent PK (CDK); (CDK10) 340 8.93 38.57 1–307 Cell-cycle-associated PK; (T. gondii ARI) 234 6.00 × 10−75 76 KYF44017.1
SRCN_977 Cell-cycle-associated kinase CDK Ca2+-dependent PK (CDK); (PITSLRE/CDK11) 1502 7.38 156.35 1114–1429 Cell-cycle-associated PK CDK; (T. gondii p89) 454 3.00 × 10−135 92 KFG28420.1
5. Kinase Group ‘Other’ (OPK; i.e., kinases with conventional PK (ePK) domains that do not fit into any of the other major groups of kinases)
SRCN_108 Unc-51-like autophagy activating kinase 1 (ULK1) ULK 343 7.13 38.90 1-223 ULK kinase; (T. gondii VAND) 376 2.00 × 10−130 75 KFH07419.1
SRCN_1606 eIF2 kinase IF2K-C PEK; (general control nonderepressible 2 (GCN2)) 4034 8.98 406.57 1235–2178 eIF2 kinase IF2K-C; (T. gondii VAND) 259 4.00 × 10−67 35 KFH07289.1
SRCN_2076 Rhoptry kinase family ROP30 Conserved hypothetical protein 1276 9.18 134.73 812–1260 ROP30 (T. gondii VEG) 230 3.00 × 10−63 53 CEL76436.1
SRCN_2123 Rhoptry kinase family ROP35 PLK; (PLK-Unclassified) 291 9.30 33.50 53–265 ROP35; (T. gondii RUB) 207 2.00 × 10−61 43 KFG59037.1
SRCN_3216 Rhoptry kinase family ROP32 CAMK-Unique 523 7.08 57.00 214–520 Putative PK; (T. gondii VAND) 167 1.00 × 10−42 30 KFH00232.1
SRCN_2183 Rhoptry kinase family ROP35 Aurora-like 226 6.36 25.84 1–212 ROP35; (T. gondii VEG) 198 8.00 × 10−59 48 ESS33297.1
SRCN_2271 Putative PK (incomplete catalytic triad) NimA (Never in mitosis gene A)-related Kinase (NEK) 1463 9.02 157.18 437–1145 Putative PK; (N. caninum L) 327 5.00 × 10−90 68 XP_003881849.1
SRCN_2403 Aurora kinase (incomplete catalytic triad) PLK; (SAK/Plk4) 778 9.79 79.92 492–778 Putative Aurora kinase; (N. caninum L) 127 3.00 × 10−28 44 XP_003880644.1
SRCN_2630 NimA related kinase (NEK) family protein NEK 351 8.70 38.38 1–336 NEK kinase; (T. gondii ME49) 242 8.00 × 10−75 52 XP_018638598.1
SRCN_286 Wee kinase Inhibitory regulator of the RAS-cAMP (IRA1) kinase suppressor (IKS) 1019 6.20 106.67 598–959 Wee kinase; (H. hammondi) 445 5.00 × 10−141 58 XP_008882669.1
SRCN_3075 Tyrosine kinase-like (TKL) protein Numb-associated kinase (NAK) 1571 8.41 164.18 16–500 TKL; (T. gondii TgCatPRC2) 138 1.00 × 10−32 73 KYK64203.1
SRCN_3142 PIK3R4 kinase-related Aurora 997 8.72 106.54 548–899 Putative PIK3R4 kinase-related protein; (N. caninum L) 449 2.00 × 10−137 60 XP_003885774.1
SRCN_3151 NimA related kinase (NEK) family protein NEK 3186 7.96 318.69 352–656 NEK kinase; (T. gondii VEG) 468 7.00 × 10−131 73 CEL78174.1
SRCN_3247 Rhoptry kinase family ROP27 Ciliate-D 345 8.94 38.81 23–325 ROP27; (T. gondii p89) 163 4.00 × 10−43 31 KFG37427.1
SRCN_3417 Aurora kinase Aurora 438 7.65 48.49 14–289 Aurora kinase; (T. gondii TgCatPRC2) 490 3.00 × 10−155 76 KYK63669.1
SRCN_3444 Unc-51-like Autophagy activating kinase 1 (ULK1) ULK 406 6.52 44.69 12–406 ULK kinase; (T. gondii RUB) 232 3.00 × 10−71 61 KFG59767.1
SRCN_3669 CMGC kinase ULK 1803 8.41 189.65 736–1200 Putative CMGC kinase; (N. caninum L) 624 0.0 62 CEL65030.1
SRCN_4410 Rhoptry kinase family ROP35 PKA-like 204 9.44 23.50 1–166 ROP35; (H. hammondi) 107 1.00E × 10−25 39 XP_008885989.1
SRCN_4503 eIF2 kinase IF2K-B PEK; (general control nonderepressible 2 (GCN2)) 158 5.76 17.59 1–158 eIF2 kinase IF2K-B (T. gondii TgCatPRC2) 149 5.00 × 10−41 74 KYK69938.1
SRCN_4528 NimA related kinase (NEK) family protein NEK 187 8.20 21.23 1–186 NEK kinase; (H. hammondi) 177 4.00 × 10−54 64 XP_008885186.1
SRCN_2404 Aurora kinase (incomplete catalytic triad) Serum and glucocorticoid induced Kinase (SGK) 295 8.81 31.42 1–249 Putative Aurora kinase; (N. caninum L) 126 7.00 × 10−31 43 CEL65223.1
SRCN_5653 PEK kinase Aurora 626 8.27 60.75 513–626 PEK kinase (T. gondii TgCatPRC2) 251 1.00 × 10−76 60 KYK62422.1
SRCN_5943 NIMA-related kinase NIMA1 NEK 2842 9.04 295.44 73–383 NIMA-related PK NIMA1; (T. gondii MAS) 486 2.00 × 10−140 67 KFH05809.1
SRCN_6157 Unc-51-like autophagy activating kinase 1 (ULK1) ULK 2420 9.38 250.58 1380–1672 ULK kinase (T. gondii ME49) 99.8 3 × 10−21 38 XP_018635814.1
SRCN_6184 Myosin-light-chain kinase Ciliate-E2-Unclassified 478 5.42 53.65 177–474 ROP19A (T. gondii ME49) 127 3.00 × 10−30 27 XP_018637476.1
SRCN_6572 Tyrosine kinase-like (TKL) ULK 622 6.13 68.80 1–345 TKL; (T. gondii VAND) 181 1.00 × 10−46 74 KFH00338.1
SRCN_6812 PK ULK 199 6.74 22.72 1–183 PK; (H. hammondi) 172 3.00 × 10−48 53 XP_008887491.1
SRCN_7083 Rhoptry kinase family ROP35 PKA-like 262 9.62 30.26 1–242 ROP35; (H. hammondi) 127 6.00 × 10−32 39 XP_008885989.1
SRCN_4310 Rhoptry kinase family ROP33 Kinase Homologous to SPS1/STE20 (KHS) 1591 9.85 169.63 1265–1578 ROP33; (H. hammondi) 306 2.00 × 10−87 39 XP_008887632.1
SRCN_7082 Rhoptry kinase family ROP33 Kinase Homologous to SPS1/STE20 (KHS) 403 9.59 45.92 77–390 ROP33 (T. gondii p89) 277 3.00 × 10−89 40 KFG45248.1
SRCN_7084 Rhoptry kinase family ROP37 Ribosomal protein S6 Kinases (RSK; (RSK)) 339 5.41 38.09 19–334 ROP37; (N. caninum L) 144 1.00 × 10−36 36 CEL64242.1
6. Kinase Group “Sterile” serine/threonine kinase, or sterile-phenotype kinases (STE)
SRCN_1328 Serine threonine kinase Conserved hypothetical protein 1461 9.29 158.74 559–671 Hypothetical protein, conserved; (E. maxima) 88.6 5.00 × 10−16 68 XP_013335801.1
SRCN_5172 “Sterile” serine/threonine kinase (STE) Mammalian Sterile 20-like (MST)) 6552 6.14 671.51 3410–4122 STE kinase; (T. gondii TgCatPRC2) 412 1.00 × 10−114 54 KYK71951.1
7. Kinase Group Tyrosine Kinase-Like (TKL)
SRCN_1435 Tyrosine kinase-like (TKL) Mixed lineage kinase (MLK); (Leucine Zipper-bearing Kinase (LZK)) 3064 8.05 306.74 2540–3060 Tyrosine kinase-like (TKL) protein; (N. caninum L) 278 3.00 × 10−73 72 CEL64955.1
SRCN_1571 Tyrosine kinase-like (TKL) Microtubule-associated S/T kinase (MAST) 550 9.76 59.91 135–501 Conserved hypothetical protein; (E. praecox) 76.3 6.00 × 10−13 53 CDI87140.1
SRCN_3466 Tyrosine kinase-like (TKL) TKL-Unique 3002 9.87 320.97 2342–2997 Tyrosine kinase-like (TKL) protein; (H. hammondi) 202 3.00 × 10−50 65 XP_008887506.1
SRCN_3928 Tyrosine kinase-like (TKL) LISK - LIMK (LIM kinase) and TESK (Testicular protein Kinase); (DD1) 5842 8.78 608.80 3639–4268 Tyrosine kinase-like (TKL) protein; (T. gondii TgCatPRC2) 216 2.00 × 10−61 79 KYK63216.1
SRCN_4277 Kinase domain-containing protein TKL-ciliate1 2256 8.43 240.25 1570–2256 Tyrosine kinase-like (TKL) protein; (N. caninum L) 204 8.00 × 10−51 61 CEL67693.1
SRCN_811 Tyrosine kinase-like (TKL) TKL-Unique 1099 9.18 119.26 814–1083 Putative tyrosine kinase-like (TKL) protein; (E. acervulina) 403 6.00 × 10−125 59 XP_013252162.1
8. Kinase Group Atypical (aPKs)
SRCN_3601 Atypical MEK-related kinase Muscle-associated kinase TRIO 950 7.20 103.14 381–850 Atypical MEK-related kinase; (T. gondii GT1) 171 1.00 × 10−42 32 EPR62774.1
SRCN_5962 Atypical MEK-related kinase Rho-associated protein kinase (ROCK)-like 805 5.01 87.71 525–805 Atypical MEK-related kinase; (H. hammondi) 127 4.00 × 10−29 56 XP_008884362.1
SRCN_3988 Phosphatidylinositol 3-/4-kinase (PI3K) Atypical/PIKK/ATM 4251 5.95 440.5 3671–3772 PI3K, ; (H. hammondi) 268 1 × 10−69 55 XP_008886631.1
SRCN_6465 Phosphatidylinositol 3-/4-kinase (PI3K) Atypical PIKK/ATM 207 5.2 23.24 5-122 Phosphatidylinositol 4-kinase, partial; (T. gondii p89) 272 4 × 10−93 96 KFG28404.1
SRCN_1259 Phosphatidylinositol 3-/4-kinase (PI3K) Atypical/PIKK/FRAP 1362 9.56 142.85 1140–1311 PI3K; (T. gondii MAS) 317 3 × 10−94 76 KFH10008.1
SRCN_6464 Phosphatidylinositol 3-/4-kinase (PI3K) No hits found 2108 8.95 209.68 1626–1722 Phosphatidylinositol 3-4-kinase; (T. gondii p89) 244 1 × 10−63 66 KFG28409.1
SRCN_1743 Pyruvate dehydrogenase kinase Atypical/PDHK/BCKDK 930 6.50 101.41 291–426 PDHK, isoenzyme-2; (N. caninum L) 458 6 × 10−146 45 CEL70411.1

a The protein sequences and their corresponding identified were obtained from the Toxoplasma Genomics Resource database (Release 28; Version May 2016) [42]; b the descriptions of the protein sequence are based on BLASTp annotations using Blast2GO (see the text for details); c the kinase classification is based on BLASTp on the kinase database.