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RESEARCH ARTICLE
Received: 3 December 2012 / Revised: 13 April 2013 / Accepted: 7 May 2013 / Published online: 12 June 2013
The National Academy of Sciences, India 2013
Introduction
Castor (Ricinus communis L) is an important non-edible oil
seed crop belonging to the family Euphorbiaceae. The
genus Ricinus is monotypic with R. communis as the only
species [1]. Castor is diploid (2n = 20) and is presumed to
be a secondary-balanced polyploid with basic number of
chromosomes x = 5 [2]. The crop is extensively grown in
tropics and sub-tropics as well as temperate regions. Castor
is predominantly cultivated in India, China, Brazil, USSR,
the EEC and Japan. India accounts for nearly 60 % castor
production [3]. Castor bean oil besides its use as vegetable
and medicinal oil has several industrial applications.
Dehydrated castor oil is used in the paint and varnish
industry, manufacture of a wide range of products like
nylon fibre, jet engine lubricants, hydraulic fluids, plastics,
artificial leather, fibre optics, bullet proof glass and bone
prostheses [35]. High level (*90 %) of ricinoleic acid
present in castor oil contributes to the stable viscosity index
and high lubricity even under low temperature conditions
and hence castor oil is used in the manufacture of antifreeze fuels and lubricants for space rockets. The sequence
analysis established genome size of the castor as *320 Mb
with an estimated number of 31,000 genes [6].
Cytochrome P450s(CYPs) represents a large class of
enzymes which mediate diverse metabolic reactions in
almost all organisms such as bacteria, fungi, animals and
plants. Cytochrome P450s have been found to play vital
role in the metabolism and detoxification [7]. CYPs in
various plants are known to perform reactions of both
primary and secondary metabolism and are involved in the
production of fatty acids, sterols, plant hormones, flavonoids, terpenoids, lignin and signalling molecules etc.
Cytochrome P450s catalyze oxidation of various substrates
using oxygen and NAD (P) H [8, 9]. Furthermore, these
123
Results
Sequence analysis of castor genome disclosed 210 putative
CYP coding genes grouped into ten clans (CYP51, 71, 72, 74,
85, 86, 97, 710,711 and 727) consisting of 45 families represented by 77 subfamilies (Table 1). The highest number
(123) of the CYP genes are present in the CYP71 clan which
represents the whole set of A-type CYP genes belonging to
123
M. S. Kumar et al.
RcCYP72A15
RcCYP72A16
RcCYP72A17
RcCYP72A18
RcCYP709B2
RcCYP714A1
RcCYP714A2
RcCYP714A3
RcCYP714A4
RcCYP714A5
RcCYP715A1
RcCYP721A1
RcCYP734A1
RcCYP734A2
RcCYP734A3
RcCYP734A4
RcCYP734A5
RcCYP734A6
RcCYP735A1
29983.m003136
29739.m003779
29983.m003138
30076.m004534
29848.m004472
29794.m003364
27955.m000386
29907.m000622
27955.m000385
30169.m006256
30170.m014009
28320.m001078
30174.m009065
30174.m009066
30174.m009067
30174.m009070
30174.m009068
30205.m001577
RcCYP74B2
RcCYPN
29901.m000415
30910.m000018
RcCYP87A4
27985.m000878
RcCYP87A5
RcCYP87A2
30190.m011135
30147.m014292
RcCYP85A1
29790.m000806
CYP85 clan
RcCYP74A1
30170.m013972
CYP74 clan
RcCYP72A14
29633.m000932
RcCYP51G1
CYP
gene name
29739.m003612
CYP72 clan
30128.m008568
CYP51 clan
Castor gene ID
8,540
2,516
2,519
3,182
1,251
6,105
1,796
3,533
2,135
2,196
2,162
1,884
1,917
3,291
4,529
1,942
4,996
2,165
2,893
3,507
3,644
3,868
2,096
1,899
1,919
1,797
2,055
2,442
Length of
genomic
seqs (bp)
No. of
Intron (s)
197,325,150,252,
90,79,107,128,94
215,340,150,249,93,79,
118,126,94
194,322,150,252,90,79,
107,128,256,57
104,322,150,246,87,
79,107,125,88
1,251
471,382,638
1,557
274,227,242,379,426
280,224,233,379,429
94,224,233,379,429
169,221,233,376,429
289,224,233,376,429
274,224,236,379,429
295,224,242,376,453
492,245,364,385,356
606,426
301,159,95,168,88,417,
227,251,369,340
286,227,245,355,426
286,227,251,349,426
295,227,254,370,426
280,173,251,376,432
259,218,251,379,423
103,81,221,245,391,426
274,218,245,379,414
14,181,245,376,426
277,221,245,364,426
462,999
Exon(s)
size (bp)
4,265, 262
910
211,104, 100,140
402
Intron (s)
size (bp)
2,0,0,0,0,1,0,2
2,0,0,0,0,1,2,2
2,0,0,0,0,1,0,2,0
2,0,0,0,0,1,0,2
0,1
1,0,2,0
1,0,2,0
1,0,2,0
1,0,2,0
1,0,2,0
1,0,2,0
1,0,2,0
0,2,0,1
1,1,0,0,1,1,0,2,2
1,0,2,0
0,0,2,0
1,0,2,0
1,0,2,0
1,0,2,0
1,0,2,0
1,1,0,2,0
1,0,0,1
2,0,2,0
1,0,2,0
Intron (s)
phase
7,118
1,052
884
1,874
4,527
1,985
590
837
734
333
375
1,701
2,687
910
2,581
626
582
1,968
2,072
2,356
566
432
389
555
522
402
Sum of
intron
length
(bp)
473
487
544
435
416
496
518
515
514
452
475
516
513
529
613
343
445
512
493
512
523
503
509
488
509
413
510
486
No. of
amino
acids
123
123
RcCYP87A6
RcCYP87A7
RcCYP87A8
RcCYP87A9
RcCYP87A10
RcCYP88A4
RcCYP90B1
RcCYP90C1
RcCYP90D1
RcCYP707A1
RcCYP707A2
RcCYP707A3
RcCYP707A4
RcCYP716A1
RcCYP716A2
RcCYP716A3
RcCYP716A4
RcCYP716A5
RcCYP716B1
RcCYP716C1
RcCYP716C2
RcCYP716D1
RcCYP716E1
RcCYP716F1
RcCYP718A1
RcCYP720A1
30138.m003878
30170.m014356
27985.m000880
29609.m000602
29709.m001228
29634.m002158
29634.m002059
28694.m000680
30115.m001196
29801.m003223
30170.m013873
29801.m003183
28226.m000875
30018.m000548
30152.m002401
29728.m000795
30074.m001374
28226.m000853
29776.m000481
29776.m000483
30063.m001421
28842.m000941
29666.m001453
29993.m001052
30172.m000208
CYP
gene name
28448.m000359
Castor gene ID
Table 1 continued
2,014
1,805
2,302
2,914
No. of
Intron (s)
135,234,84,79,
122,128,115
961,179
501,442,188,306
502,372,188,306
88,456,189,179
931,188,303
940,188,297
946,188,309
958,188,306
961,188,294
943,188,303
451,504,188,300
928,188,300
206,322,150,249,87,
79,107,119,97
194,322,150,204,90,
79,107,119,103
206,319,387,175,
107,119,94
182,322,153,249,90,
79,107,119,112
221,325,150,246,90,
186,122,109
245,325,150,273,97,
125,128,100
203,325,153,234,93,
79,119,234
233,478,270,90,79,
107,122,100
200,331,150,249,87,
79,107,132
321,150,249,90,76,
107,128,100
212,331,150,249,90,
79,107,128,94
200,328,153,249,90,
79,107,128,94
197,325,150,252,90,
79,108,128,91
Exon(s)
size (bp)
665
387, 384, 94
329, 241
166, 142
183, 85
402, 81
108, 98
97, 86
94, 93
Intron (s)
size (bp)
0,0,0,1,0,2
0,1,0
1,1,0
1,1,1
1,0
1,0
1,0
1,0
1,0
1,0
1,1,0
1,0
2,0,0,0,0,1,0,2
2,0,0,0,0,1,0,2
2,0,0,1,0,2
2,0,0,0,0,1,0,2
0,1,0,0,0,0,2
2,0,0,0,1,0,2
2,0,0,0,0,1,0
2,0,0,0,1,0,2
2,0,0,0,0,1,0
0,0,0,0,1,0,2
2,0,0,0,0,1,0,2
2,0,0,0,0,1,0,2
2,0,0,0,0,1,0,2
Intron (s)
phase
1,117
665
865
1,546
391
570
308
268
483
206
183
1,671
187
994
1,422
756
1,756
3,444
5,065
1,918
1,969
1,790
1,345
1,059
888
807
Sum of
intron
length
(bp)
298
379
478
455
303
473
474
480
483
480
477
480
471
471
455
468
470
482
480
479
492
444
406
479
475
472
No. of
amino
acids
134
1,303
1,992
1,733
1,711
1,935
1,649
1,617
3,114
1,603
2,410
2,790
2,163
3,710
4,893
6,508
3,358
3,448
3,125
2,566
2,499
2,316
2,227
Length of
genomic
seqs (bp)
RcCYP722A2
RcCYP722A3
RcCYP722B1
RcCYP724A1
RcCYP724A2
29634.m002092
29863.m001089
29982.m000224
29633.m000931
30005.m001270
RcCYP86A3
RcCYP86B1
RcCYP86C1
RcCYP94B1
RcCYP94B2
RcCYP94B3
RcCYP94B4
RcCYP94C1
RcCYP94C2
RcCYP94D1
RcCYP94D2
RcCYP96A1
RcCYP96A2
RcCYP96B1
RcCYP96B2
RcCYP704A2
RcCYP704A3
RcCYP704A4
30174.m008617
30094.m000683
29929.m004790
30147.m014517
29883.m002015
30190.m010938
29883.m002017
29791.m000529
28779.m000137
29811.m000531
30078.m002275
29917.m002008
29917.m002010
29409.m000268
29660.m000772
30174.m008914
30174.m008915
30194.m000057
RcCYP704B1
RcCYP86A2
30190.m011234
29813.m001518
RcCYP86A1
29681.m001310
CYP86 clan
RcCYP722A1
CYP
gene name
30170.m014078
Castor gene ID
Table 1 continued
2,345
2,680
1,555
2,046
1,613
1,518
1,518
1,530
1,521
1,563
1,506
1,500
1,873
1,530
1,494
1,443
1,563
3,062
2,045
1,921
1,536
3,404
3,569
2,305
2,867
4,067
4,063
Length of
genomic
seqs (bp)
No. of
Intron (s)
330,216,297,
387,201,180
309,216,291,
321,45
277,26,144,92,
337,168
522,285,339,
201,192
1,086,324
1,518
1,518
1,530
1,521
1563
1506
1,500
1,524
1,530
1,494
1,443
1,563
1260,393
256,1313
1,638
1,536
221,325,153,246,87,
79,107,122,76,156
200,325,153,243,87,
79,111,136,115
209,319,153,267,
87,79,107,119,91
248,325,150,252,84,
79,107,119,112
260,325,150,252,87,
79,107,119,100
146,322,150,184,80,90,
79,107,119,100
Exon(s)
size (bp)
203
1,409
110
Intron (s)
size (bp)
0,0,0,0,0
0,0,0,0
1,0,0,2,0
0,0,0,0
2,0,0,0,0,1,0,2,0
2,0,0,0,0,1,1,2
2,0,0,0,0,1,0,2
2,0,0,0,0,1,0,2
2,0,0,0,0,1,0,2,
2,0,0,1,0,0,1,0,2
Intron (s)
phase
734
1,498
511
507
203
1,409
110
1,832
2,120
874
1,391
2,588
2,686
Sum of
intron
length
(bp)
536
394
347
512
469
505
505
509
506
520
501
499
507
509
497
480
520
550
522
545
511
523
482
476
491
492
458
No. of
amino
acids
123
123
RcCYP97C1
30078.m002224
28223.m000100
RcCYP71A24
RcCYP71A25
RcCYP71A26
RcCYP71A27
RcCYP71A28
RcCYP71B1
RcCYP71B4
RcCYP71B8
RcCYP71B9
RcCYP71B10
RcCYP71B11
RcCYP71B13
RcCYP71B14
RcCYP71B22
RcCYP71B23
RcCYP71B24
RcCYP71B25
RcCYP71B26
RcCYP71B27
RcCYP71B28
30147.m013842
30147.m013843
30147.m013847
30147.m013848
30129.m000355
29724.m000821
30169.m006288
29785.m000965
29785.m000959
29785.m000962
29826.m000757
30169.m006277
29887.m000239
29910.m000943
29878.m000239
29887.m000240
29887.m000241
30169.m006273
30169.m006275
RcCYP727A1
RcCYP711A1
30147.m013846
CYP71 clan
29686.m000867
CYP727 clan
29739.m003566
CYP711 clan
RcCYP710A1
RcCYP97B3
29724.m000853
CYP710 clan
RcCYP97A3
CYP
gene name
30128.m009010
CYP97 clan
Castor gene ID
Table 1 continued
1,608
1,616
1,647
1,754
1,810
13
15
No. of
Intron (s)
891,615
906,615
894,621
882,621
477,26,199,615
897,627
882,621
996,609
945,654
906,624
906,624
666,153,624
157,638,627
123,466,359
942,615
963,621
945,621
915,612
915,618
915,612
208,200,642,
148,127,136,78
248,153,784,
96,330
903,615
501,297,198,102,
141,108,83,163,66
71,178,108,85,239,
174,48,60,87,89,
92,62,150,225
139,163,217,72,111,
192,162,96,102,33,
81,83,132,61,66,189
Exon(s)
size (bp)
102
95
132
251
268
112
129
79
892
856
79, 611
39, 99
117, 97
88
477
975
338
187
918
630, 593,290, 77
143
0, 2, 0
0,0
1,0
0,1
1,0,0,1,2,0
2,2,0,0
0,0,0,0,0,0,2,0
2,0,0,1,0,0,0,
0,0,2,1,0,0
1,2,0,0,0,0,0,
0,0,0,0,2,2,0,0
Intron (s)
phase
Intron (s)
size (bp)
102
95
132
251
493
268
112
129
79
892
856
690
138
214
88
477
975
338
187
918
3,347
1,590
143
2,670
17,801
3,247
Sum of
intron
length
(bp)
501
506
504
500
438
507
500
534
532
509
509
480
473
315
518
527
521
508
510
508
512
536
159
552
555
632
No. of
amino
acids
136
1,792
1,615
1,734
1,678
2,475
2,501
2,185
1,560
1,662
1,645
2,061
2,541
1,865
1,720
2,445
4,886
3,201
480
4,329
19,469
5,146
Length of
genomic
seqs (bp)
RcCYP71B29
RcCYP71B30
RcCYP71B31
RcCYP71B36
RcCYP71B37
RcCYP71B38
RcCYP71B39
RcCYP71B40
RcCYP71B41
RcCYP71B42
RcCYP71C1
RcCYP71C2
RcCYP71C3
RcCYP71D1
RcCYP73A5
RcCYP73A6
RcCYP75B1
RcCYP75B2
RcCYP75B3
RcCYP75B4
RcCYP76C3
RcCYP76C4
RcCYP76D1
RcCYP76D2
RcCYP76D3
RcCYP76D4
RcCYP76E1
RcCYP76G1
RcCYP76G2
RcCYP76H1
RcCYP76H2
RcCYP76I1
RcCYP76I2
RcCYP76I3
RcCYP76I4
RcCYP77A4
RcCYP77B1
30169.m006282
30169.m006285
29792.m000625
29929.m004561
29629.m001350
29629.m001392
29929.m004562
29792.m000626
30142.m000643
29792.m000624
30206.m000783
29785.m000966
29929.m004748
29976.m000504
43540.m000048
30138.m003983
30146.m003563
29706.m001271
29739.m003754
30190.m011068
30190.m011069
30147.m014296
30169.m006290
30169.m006295
30169.m006291
30190.m011130
29815.m000508
29815.m000509
29815.m000510
29815.m000512
29815.m000515
29815.m000516
29815.m000519
29815.m000520
29428.m000318
29842.m003625
CYP
gene name
30169.m006279
Castor gene ID
Table 1 continued
1,518
No. of
Intron (s)
1,518
1,596
930,615
930,618
939,591
960,615
473,445,621
900,609
915,630
927,651
879,627
906,51
885,606
882,609
900,606
889,245
942,624
906,630
942,624
282,453,651
875, 736
927,636
903, 510, 87
903,12
906,618
894,174,21
906,609
951,624
900,624
891,609
903,612
862,63,614
990,612
885,609
957,615
Exon(s)
size (bp)
128
153
149
107
93, 312
556
468
1,037
176
97, 113
499
142
121
84
179
156
547
573
891, 853
362
176, 1,525
723
280, 1,035
66
1,472
250, 52
824
150
108
132
269
124, 84
93
122
157
Intron (s)
size (bp)
2,0
1,1
0,0
2,1
0,0
0,0
1,1
Intron (s)
phase
128
153
149
107
405
556
468
1,037
176
210
499
142
121
84
179
156
547
573
1,744
362
1,701
723
1,315
66
1,472
302
824
150
108
132
269
208
93
122
157
Sum of
intron
length
(bp)
505
531
514
515
509
524
512
502
514
525
501
426
318
496
496
501
377
521
511
521
461
536
505
520
499
304
507
362
504
524
507
499
504
512
533
497
523
No. of
amino
acids
1,596
1,673
1,701
1,679
1,682
1,944
2,065
2,013
2,615
1,682
1,491
1,456
1,633
1,612
1,794
1,313
1,722
2,210
2,139
3,130
1,973
3,576
2,399
2,815
981
2,996
1,391
2,339
1,725
1,632
1,632
1,784
1,747
1,695
1,616
1,729
Length of
genomic
seqs (bp)
123
123
RcCYP77B2
RcCYP78A6
RcCYP78A7
RcCYP78A8
RcCYP78A9
RcCYP78A10
RcCYP78A11
RcCYP79A2
RcCYP79B3
RcCYP79B4
RcCYP79B7
RcCYP81D2
RcCYP81D3
RcCYP81D4
RcCYP81D8
RcCYP81D9
RcCYP81D10
RcCYP81D11
RcCYP81H1
RcCYP81K1
RcCYP82C2
RcCYP82C3
RcCYP82C4
RcCYP82C5
RcCYP82C6
RcCYP82C7
RcCYP82C8
RcCYP82C9
RcCYP82C10
RcCYP82C11
RcCYP82C12
RcCYP82C13
RcCYP82G1
RcCYP83B1
RcCYP83B3
RcCYP83B4
RcCYP84A1
28014.m000118
28256.m000134
28644.m000933
30068.m002578
30138.m003950
29929.m004656
29910.m000911
29910.m000917
29910.m000914
28438.m000050
29970.m001003
29910.m000948
30170.m014208
29970.m001002
30170.m013774
30170.m013773
30170.m013780
29970.m000998
30170.m014207
29851.m002485
30170.m013958
30170.m013953
30120.m000372
30170.m013957
30170.m013963
29676.m001679
30170.m013949
30170.m013950
30170.m013960
30170.m013964
30170.m013965
30120.m000371
30170.m014153
30170.m014151
30174.m009168
30131.m007121
CYP
gene name
29842.m003626
Castor gene ID
Table 1 continued
942
2,331
1,797
1,611
2,684
No. of
Intron (s)
942
714,612
879,612
885,609
942,630
963,624
939,630
945,621
948,624
948,633
939,627
945,621
927,666
951,627
400,464,624
954,633
963
897,621
327,633
903,612
882,621
903,627
891,621
930,615
918,636
272,469,618
1,017
1,113
981,9
1,173
963,597
783,606
960,636
996,618
999,603
405,618
1,518
Exon(s)
size (bp)
1,005
306
117
1,112
145
654
1,542
122
267
721
580
53
1,158
90, 207
112
187
12,217
158
624
406
1,072
766
1,472
149, 1,745
81
1,166
533
349
94
92
157
Intron (s)
size (bp)
1,0
2,0
Intron (s)
phase
1,005
306
117
1,112
145
654
1,542
122
267
721
580
53
1,158
297
112
187
12,217
158
624
406
1,072
766
1,472
1,894
81
1,166
533
349
94
92
157
Sum of
intron
length
(bp)
313
441
496
497
523
528
522
521
523
526
521
521
530
525
495
528
320
505
319
504
500
509
503
514
517
452
338
370
329
390
519
462
531
537
533
340
505
No. of
amino
acids
138
1,732
2,223
3,108
1,694
1,848
2,287
2,146
1,646
2,736
1,785
1,699
963
1,790
13,177
1,673
2,127
1,936
2,584
2,311
3,026
3,253
1,017
1,113
1,071
1,173
2,726
2,045
1,950
1,708
1,694
1,180
1,529
Length of
genomic
seqs (bp)
RcCYP84A2
RcCYP84A3
RcCYP84A4
RcCYP89A4
RcCYP89A5
RcCYP89A6
RcCYP89A7
RcCYP89A8
RcCYP89A9
RcCYP89A10
RcCYP89A11
RcCYP89A12
RcCYP89B1
RcCYP93D1
RcCYP93D2
RcCYP93D3
RcCYP93D4
RcCYP93D5
RcCYP98A3
RcCYP98A4
RcCYP98A5
RcCYP701A3
RcCYP703A2
RcCYP706A4
RcCYP706A5
RcCYP706A7
RcCYP712A1
RcCYP712A2
RcCYP712B1
30138.m003926
30174.m008711
29827.m002605
30148.m001481
30073.m002236
30148.m001477
30148.m001478
30148.m001476
30148.m001482
30148.m001475
30148.m001483
29083.m000045
29216.m000256
29216.m000255
30152.m002423
29788.m000321
29788.m000323
29940.m000401
30147.m014117
29940.m000400
30170.m013942
29742.m001406
30190.m011008
30190.m011007
30147.m014189
29216.m000258
27647.m000174
29216.m000257
CYP
gene name
28196.m000205
Castor gene ID
Table 1 continued
2,151
3,153
1,967
1,694
3,820
1,705
1,888
2,988
4,472
1,536
3,583
2,230
1,296
3,031
2,434
2,380
1,738
1,545
1,503
1,568
1,551
1,542
1,575
1,601
1,551
1,551
1,783
2,629
1,661
Length of
genomic
seqs (bp)
No. of
Intron (s)
921,630
939,618
912,630
894,235,347
936,621
960,633
909,639
178,398,612
1,536
493,398,645
609,231,681
891,111
16,959,666
802,428
811,671
1,173
1,545
1,503
1,557
1,545
1,542
1,575
1,563
1,551
1,551
675,828
714,822
903,615
Exon(s)
size (bp)
600
1,281
425
172, 46
2,116
112
340
2,311, 973
1,574, 335
70, 639
294
157, 1,233
1,204
898
277
1,093
143
Intron (s)
size (bp)
0,1
1,2,0,2,2,1,1
1,0
1,0
0,0
1,0
Intron (s)
phase
600
1,281
425
218
2,116
112
340
1,456
3,284
1,909
709
294
1,390
1,204
898
277
1,093
143
Sum of
intron
length
(bp)
516
518
513
491
518
530
515
513
395
511
511
506
333
546
409
493
390
514
500
518
514
513
524
520
516
516
501
511
505
No. of
amino
acids
123
M. S. Kumar et al.
914 1 7
000
9
000
10.m
-299 9910.m
9B 4
-2
911
YP7
9B3
000 00050
RcC
YP7
0.m
0
RcC 2-2991 438.m
79A B7-28
YP
79
YP
RcC
CY
2
79A
YP 3
AtC P79B 2
06 2
14 A
Y 9B
00 703
AtC YP7
.m P
42 Y
AtC 2
97 AtC
9F
2-2
P7 F1
3A
1
CY 79
At CYP 79C2 79C P70
At YP CYP cCY
C
R
At At
0.1
AtCYP71B21
AtCYP71B22
AtCYP71B38P
AtCYP71B5
AtCYP71B30P
AtCYP71B
AtCYP71B 32
31
AtCYP7 AtCYP71B8
1B23
AtCYP7
1B7
AtCY AtCYP7
1B15
P71B
AtCY AtCYP71 28
P71
B29
AtC B2
YP7
1
A
AtC tCYP7 B27
AtC YP71B 1B16
17
Y
AtCAtCYP P71B1
9
Y
7
AtC P71B 1B20
AtC AtC YP71 24
YP YP7 B3
71 1B
A B4 25
A tCY
A t tCYP P 7 1 B
CY 71 11
A P B
A t 71 12
A tC CY B1
A tC YP P71 3
At tCY YP7 71B B14
At CY P71 1B2 6
B 6
C P
A Y 71 33
At At tCY P71 B34
B
At CY CY P71 35
P P
R CY 71B 71B B9
cC P7 3 3
Y 1B 7 6
P7 1
1B 0
36
-2
97
92
.m
00
06
25
Rc C
76
14
00 78 7
.m 014 147 1
8
48
36
01 .m0 00 14
22
9-3 48 8.m 00
00
9A 301 014 8.m 3
P8
8-3 14 48 3.m 75
CY 9A A7 -30 01 07 14
Rc YP8 P89 A5 .m0 6-30 00 482
C CY P89 148 A 8.m 01
9
4
0
Rc
c
0
R
P8 301 .m
CY 2-3
Rc A1 cCY 11- 148
9
R 9A -30
P8
P8 A10
CY 89
Rc YP
cC
R
Rc
A AtC
At tCY YP
Rc
At CY P8 89
P 9 A
Rc CY
C
P8
At YP 89A A2 3
C
Rc
CY YP8 9A4
At At CYP 89A 7
9
P7
7B B1-2 2982 CYP CYP 89A 6
1
9
8
7
8
0
.
9
Rc
29
9 5
84 83.m m00 A9 A4
CY
2.m
P
0 26
Rc 77B
At 003 0004 05
CY
2
C
P7 -298 YP 625 5
4
7A
7
4-2 2.m0 7B1
9
0
AtC 428.m 362
RcC
YP
00 6
YP
701
AtC AtC 77A9 0318
A3-3
YP YP7
017
7A
77
A
0.m
7
013 tCYP A6
77A
AtC 942
4
Y
RcC P701A
YP7
3
3A5
-299
RcC
76.m
YP7
000
3A6
A
5
-435
40.m tCYP7 04
0000 3A5
48
RcCY
AtC
Y
4-3014 AtCYP P98A9
98A8
RcC 7.m0141
RcCY YP98A5- 17
P98A329
29940. 940.m0004
m
00
00
RcCYP7
AtCYP90401
8A8-286
8A3
RcCYP7
44.m00
09
8A7-282
56.m0001 33
34
RcCYP78A1 AtCYP78A7
0-30138.m0
03950
AtCYP78A10
AtCYP78A5
RcCYP78A11-29929.m004656
RcCYP78A9-30068.m002578
RcCYP78A6-28014.m000118
AtCYP78A8
AtCYP78A9
AtCYP78A6
29815.m000520
6I4RcCYP7
5.m000519
981
3-2
RcCYP76I
000516
-29815.m
15
RcCYP76I2 6I1-29815.m0005 000512
815.m
RcCYP7
6H2-29
10
RcCYP7 815.m0005 6G1
29
tCYP7 09
P76H1A
Y
cC
R
0005 08
05
815.m
G2-29 815.m00
76
P
30
RcCY P76G1-290.m0111 006291
RcCY E1-3019 0169.m 96
-3
4
76
6D
0142
YP
7.m
YP7
RcC
RcC 1-3014
6D
069
290
YP7
006 .m011
.m
RcC
169 0190
C7
2-30 4-3
P76 3
76D P76C 8
CY P76C
295
t
P
6
Y
0
A
Y
6
0
Y C6
RcC RcC 0110
9.m
AtC P76 C5 6C1
016
.m
Y 76 P7
190
D3-3
C
t
6
0
7
P
A Y tCY 6C4 6C2
3-3
YP
C A P7 7
76C
RcC
At
P
YP
CY Y
RcC
At AtC
89
41
A2
01
.m 1008 706 A1 07
7
14 01 YP 06 0
-30 .m tC P7 011
A7 0190 A tCY 0.m A7 A3 A4
6
0
A 19 706 706 706 6
P7 4-3
30 P P P
CY 6A
5- Y Y Y 06A 5
Rc P70
6A AtC AtC AtC P7 6A
70
CY
0
CY 7
Rc
YP
C
At YP
Rc
tC
A
2
2
A A3 28
05 05 5A 30
P7 P7 0 5A
Y Y P7 70 2
tC tC Y P A2
A A AtC tCY 705 3
A P 5A
0
CY P7 A6 8
At CY 705 5A1 33
At CYP P70 05A 3
At tCY YP7 5A1 2
24
A AtC P70 A1 A1
5
5 705A
CY 70 70
At CYP YP CYP
At AtC At 25
05A
258 00257
P7 A27
000
0
CY 05
321
At YP7 A1 216.m 216.m
000
2 -29
-29
AtC 71
88.m
74
YP 2A1 12B1
001
-297
AtC YP71 CYP7 712A2 647.m0 P93D4
P
RcC Rc AtCY 2A2-27 RcCY
1
YP7
RcC
3
2
0003
88.m 02423
-297
0
3D5 0152.m
YP9
-3
56
RcC P93D3 D1
m0002
Y
93
0255
RcC AtCYP D1-29216. 216.m00
P93
D2-29
RcCYRcCYP93
cC
Y
P7
5B
At
130
C
A
13
A tC YP
8.
At tCY YP 81D
m
At CY P8 81D 3 At 00
CY 3
CY P8 1D 2
1D 8
P
P7 983
At AtC AtC 81D 10
5B
At CY YP YP 5 P
1
Rc
C
P
8
8
YP 81 1D 1D
C
Rc
81 D7 11 4
CY YP8
D
1
P8
1D H1-2 AtCY AtC 6
3-2
997 P8 YP
Rc
9
C
1
RcC YP 910 0.m H 81
D1
YP 81D .m0 000 1
81D 8-2 00
9
2-2 997 948 98
997 0.m
0
0
AtC .m001 0100
YP 00 2
AtC 81F4 3
YP
A
RcC
AtC tCYP8 81F3
YP8
YP8 1F2
1D
RcC 4-30170 AtCYP 1F1
Y
.m
8
RcCY P81D11-3 014208 1G1
P81D
0170
9-30
.m01
R
RcCY cCYP81D10 170.m01 3780
37
P81K
1-3017 -30170.m 74
01
0.
AtCYP8 m014207 3773
1K2
AtCY
RcCYP8
2C8-296 P81K1
76.m00
AtCYP82F 1679
1
RcCYP82C5
RcCYP82C2-298
-30120.m000
51.m002485
372
RcCYP82G1-30120.m0
00371
AtCYP82G1
RcCYP82C13-30170.m013965
RcCYP82C12-30170.m013964
RcCYP82C7-30170.m013963
RcCYP82C11-30170.m013960
13957
RcCYP82C6-30170.m0
70.m013958
RcCYP82C3-301 0170.m013949
9-3
RcCYP82C
m013950
10-30170.
RcCYP82C 0170.m013953
2C4
2C4-3
AtCYP83
RcCYP8
2C
AtCYP8 2C2
P8
AtCY
P98A
3
64
00
6 m0
62 42.
0
83
00 01
07
m -3
00
2. 42 24
6.m
79 B 06
20
29 P71 00 65 9
0
3
41 Y 2.m 09 095
21B cC 79 00 00 962 71C 6
P7 R1-29 5.m 5.m 000 YP 096
Y
C 78 78 .m cC 00
cC
71 9-29 -29 785 R 5.m
R
P
0
8
9
1
B 1
97 A
CY 71 1B 1-2
3-2 83
Rc YP P7 1B1
3
1C YP
15
C Y 7
P7 AtC 3B1 014 4151
5
Rc RcC CYP
8
.m 1
8
CY
035
YP 0170 0.m0 0916 .m00
Rc
Rc
C
At 1-3 3017 4.m0 0129 842
B
3
3
7
3
3
P8 83B 4-301 71B1 7.m01 3843
Y
P B
01
4
C
P
Rc cCY P83 cCY 5-301 147.m 3846
1
R CY R A2 -30
.m0 13847 13848
1
Rc
P7 A26 147
0
0
7.m
CY 71 -30 7.m
Rc cCYP 1A24 -3014 8-3014
7
7
2
R YP A2
1
71A
C
7
P
c
P
Y
R CY
1
RcCP71A2 2
Rc
2
Y
AtC YP71A 4
AtC P71A2
Y
6
AtC P71A2
Y
AtC P71A23
AtCY P71A25
AtCY 71A12
P
AtCY YP71A13
AtC 1A18
AtCYP7 1A19
AtCYP7 20
1A
AtCYP7 YP71A16
AtC
27
AtCYP71A
AtCYP71A28
AtCYP71A14
AtCYP71A15
RcCYP71D1-29929.m004748
RcCYP71B13-29826.m000757
RcCYP71B38-29629.m001350
RcCYP71B39-29629.m001392
RcCYP71B
RcCYP71B24-29 23-29910.m000943
878.m000239
RcCYP71B
RcCYP71B 8-30169.m006288
37-29929.
RcCY
m0
RcCYP7 P71B40-29929.m 04561
1B
RcCYP7 27-30169.m 004562
006273
1B
RcCY 29-30169.m
P7
00
1B22
6279
RcC
RcCY YP71B25 -29887.m00
-2
P71B
0239
26-2 9887.m
R
RcC cCYP71 9887.m00 000240
B
YP7
1B31 14-30169 0241
-3
.m
RcC RcCY RcCY 0169.m0 006277
P
0
AtCYP71B 71B28 P71B4-2 6285
YP 30-3 -301
9724
84A
6
016
Rc
1
9.m 9.m006 .m00082
C
2
006
AtC
1
282 75
R YP
Rc Rc cCY 84A2 YP84
CY CY P8 -28 A4
Rc P75B P84 4A3- 196.m
CY 3- A4 301 00
RcC
Rc P75 297 -301 38.m 0205
YP
CY B2 06.m 74.m 00
84A
P7 -30 00 00 3926
1-3
5B 14 12 87
013
4-2 6.m 71 11
1.m
97 00
007
121
At 39.m 3563
C
At YP 0037
C
5
7
A
4
0
Y
At A tCY P7 5A1
CY tC P7 05A 6
Y
0
P P
5 19
A 705 705 A23
At At tCY A21 A20
CY CY P7
A
P P7 05
tC
Y At 705 05A A15
P7 C A
05 YP 8 4
A 70
5
5A
9
Fig. 1 Phylogenetic tree of A-type Cytochrome P450 proteins of castor and Arabidopsis. Rc and At represent CYP proteins of castor and
Arabidopsis, respectively
123
CY
P8
P8
Rc
0.m
7A
P8
7A
10
8-3
-29
01
CY
P8
70
9.m
42
11
At
60
014517
RcCYP94B3
-30190.m010
938
AtCYP94B
2
AtCYP94B1
RcCYP94B1-30147.m
03
59
92
35
CY
7A
CY
CY
P8
P87
-27
01
9-2
P8
0C1
00
.m
94
00
-3
B1
790
004
1
9.m
992
1-2
6C
8
P
2
CY
86C
Rc
YP
3
AtC
772
86C
000
YP
60.m
AtC
C4
-296
P86
6B2
Y
08
YP9
0020
AtC
.m
RcC
7
1
-299
6A1
10
YP9
0020
RcC
917.m
A2-29
68
YP96
m0002
RcC
29409.
96B1P
Y
RcC
3
P96A
AtCY
6A4
AtCYP9
6A15
AtCYP9
P8
CY
6C
P8
CY
At
AtCYP9
B2
86
P
CY
At
83
06
6B
P8
Rc
798
.m
81
96
-2
A1
CY
At
10
13
00
A1
86
YP
C
At
CY
7-3
7A
A4
A7
86
Rc
P8
7A
.m
00
01
2
06
43
02
56
38
.m
00
38
5.m
7
8
985
000
.m0
880
008
78
AtC
YP
RcC
85A
YP8
AtC
2
5A1
YP
-297
85A
90.m
1
RcC
000
YP7
806
20A
1-30
172.m
RcCY
AtC
P90D
0002
YP7
1-2869
08
20A1
4.m00
0680
AtCY
RcCY
P90D
P90C11
29634.
m0020
59
Rc
Rc
6A
P
CY
At
0.
19
0
-3
P8
CY
Rc
00
01
01
P8
CY
Rc
8.m
7.m
34
12
01
cC
A2
44
14
19
CY
08
28
30
30
3-
6
P8
P7
6-
5-
2-
17
86
00
4.
7
01
CY
7A
7A
7A
Rc
P8
0.2
RcCYP9
4B2-298
83.m00
2015
RcCY
P94B429883.
m0020
AtCY
17
P94C
1
RcCY
P94C
1-2979
1.m00
RcC
05
29
YP9
4C2
RcC
-287
YP9
79.m
4D1
RcC
0001
-298
YP9
37
11.m
4D2
AtC
0005
-3
007
YP
31
8.m
94D
AtC
002
1
275
YP
94D
A tC
2
YP
704
Rc
B1
CY
P7
04B
1-2
At
981
CY
Rc
3.m
CY
P7
001
At
04
P7
518
CY
A1
04
A4
P
70
Rc
-30
4A
CY
19
2
4.m
P7
At
00
04
00
CY
A2
At
57
-30
P8
CY
Rc
6A
At
17
CY
4.m
CY P86
2
P7
A4
00
04
P8
89
A3
6A
1
4
30
8
17
4.m
00
89
15
Rc
At
CY
CY
415
000
01.m
4B2
-299
YP7
4B2
AtC
YP 7
RcC
972
013
74A
0.m
YP
017
6
A tC
018
1-3
02 A
000
74A
P7
5
YP
10.m
CY
2A
2
R cC
At
70
309
A
P
02
PN
CY
P7
CY
At
A1
Rc
CY
02
8
At
P7
2A
CY P7 0
3
At
A1
8
A
CY
70
02
At
P7
YP
A4
CY AtC
08
At
P7
3
CY 8A
0
At
P7
Y
tC
A
Rc
Rc
141
AtCYP94B3
10
AtCYP90A
1
-29634.m002
158
AtCYP96A
RcCYP90B1
AtCYP96A9
AtCYP90B1
AtCYP96A1
RcCYP724A2-30005.m001270
AtCYP96A5
RcCYP724A1-29633.m000931
AtCYP96A7
AtCYP724A1
AtCYP96A8
AtCYP96A13
AtCYP96A
12
AtCYP9
AtCY
P97A
3
P97A
3-3012
8.m00
AtC
9010
YP9
7B3
RcC
YP9
7B3
-297
24.m
AtC
0008
YP9
5
7C1
RcC
YP
97C
1-3
007
8.m
AtC
RcC
002
YP
224
YP
711
Rc
710
A1
CY
A1-2
P7
822
11A
3.m
At
1-2
000
CY
973
100
P7
9.m
At
Rc
003
CY 14A1
CY
566
P7
P
Rc
Rc
727
14
CY
CY
A2
A
1-2
Rc
P7
P7
Rc
968
14
CY
14
A1
6.m
P7
A3 CYP
-29
000
14
-2
71
84
867
A2
79
4A
8.m
-2
55
A
4
9
00
-2
tC
.m
79
99
4
4
00
Y
4
.m
72
07
Rc
P7
03
0
.
m
0
86
CY
15
33
00
A
64
06
P7
1
22
14
A5
-2
79
55
.m
00
03
85
53
07
00
A3
8.m
-30
16
A2
16
16A
P7
P7
CY
CY
Rc
Rc
Rc
C
YP
5
P7
CY
Rc
01
-30
15
At
At
CY
.m
28
97
-2
A4
16
P7
CY
Rc
003779
RcCYP72A17-29739.m
AtCYP72A7
AtCYP72A8
AtCYP72A9
AtCYP72A
14
AtCYP7
2A10
AtCY
P7
2A
15
AtCY
P72A11
AtCY
P72A
13
RcCYP72A1
95
P
CY 716A
P7
2
2.m 16A
1
00
24
01
1-2
1G
00
1-3
822
05
012
48
6.m
8.m
000
008
875
568
AtC
YP
51G
AtC
1
YP
AtC
51G
YP7
2
10A
AtC
4
YP7
10A
3
AtC
YP7
10A
2
AtCY
P710A
1
74
13
00
00
.m
74
00
-3
A5
16
16
P7
CY
Rc
CY
P7
cC
B1
P7
-2
16
82
26
2-
29
.m
77
6.
08
00
04
83
RcCY
RcCYP72A15-29633.m000932
Rc
6A11
AtCYP9
6A2
56
62
00
.m
77
69
15
01
00
-3
.m
A1 5A1 A2
05
3
15
5
02
-3
P7 YP7 P73
A1
CY tC tCY
35
B1
A
Rc
A
P7
09
CY YP7
534
B2
Rc
C
004
09
67
0
At
P7
3
6.m
009
CY 709B
007
.m
At
2-3
P
068
174
009
-30
CY
09B
A4
At
P7
4.m
4
3
017
CY
P7
70
6-3
Rc
090
CY
4A
Rc
.m0
P73
174
CY
65
-30
Rc
A5
090
.m0
734
4
6
6
YP
017
090
RcC
A2-3
.m0
174
734
YP
3-30
34A
RcC
1
YP7
21A
9
0
RcC
YP7
0140
AtC
70.m
-301
21A1
YP7
RcC
1
78
10
P734A
0.m00
AtCY
1-2832
P734A
003136
RcCY
9983.m
2A16-2
3138
RcCYP7
983.m00
2A18-29
RcCYP7
1
AtCYP72C
12
4-29739.m0036
.m001089
RcCYP722A3-29863
092
29634.m002
RcCYP722A20.m014078
22A1-3017
RcCYP7
22A1
AtCYP7
000224
9982.m
22B1-2
P7
Y
RcC
88A4
P
Y
AtC
3
P88A
AtCY
1228
83
.m00
9
0
7
0031
-29
73
801.m
8A4
138
YP8
4-29
.m0
RcC
07A
7A4
170
YP7
P70
3-30
Y
A
RcC
7
6
AtC
P70
119
Y
0
C
Rc
.m0 7A2
5
1
1
70
223
-30
YP
A1
003 7A3
AtC
.m
707
1
P
70
0
Y
98
YP
1
2
C
RcC
t
7A
A
A2
70
707
YP
YP
C
t
C
A
Rc
18
P7
3
52
CY 145
10
At
00
00
21
m
.
m
.
6
41 014
3
66
99
09
0
9
9
0
0
-2
.m
1-2
.m
F1
63
8A
42 300
16
71
8
7
P
8
P
1
-2
CY
D1 048
CY
E1
Rc
16
Rc
00
16
P7
.m
P7
CY 776
CY
9
Rc
Rc
2
1C
16
P7
Y
cC
R
Fig. 2 Phylogenetic tree of non-A-type Cytochrome P450 proteins of castor and Arabidopsis. Rc and At represent CYP proteins of castor and
Arabidopsis, respectively
Discussion
The present study on the genome analysis of castor revealed
the presence of 210 putative CYP genes belonging to 45
families, grouped into ten clans. CYP450 genes identified in
castor represent *0.68 % of the predicted *31,000 protein
coding genes. The observation is in conformity with CYPs
representing 0.571.07 % of the protein coding genes of
Arabidopsis (246/23,000), rice (356/37,544) [14], poplar
123
Conclusion
Castor genome disclosed 210 putative CYP coding genes
grouped into 10 clans consisting of 45 families represented
by 77 subfamilies. The highest number (123) of the CYP
genes are A-type and the remaining 87 genes are non-A
123
M. S. Kumar et al.
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