File size: 31,891 Bytes
8ffc9f7
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
bd7cf06
8ffc9f7
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
410
411
412
413
414
415
416
417
418
419
420
421
422
423
424
425
426
427
428
429
430
431
432
433
434
435
436
437
438
439
440
441
442
443
444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460
461
462
463
464
465
466
467
468
469
470
471
472
473
474
475
476
477
478
479
480
481
482
483
484
485
486
487
488
489
490
491
492
493
494
495
496
497
498
499
500
501
502
503
504
505
506
507
508
509
510
511
512
513
514
515
516
517
518
519
520
521
522
523
524
525
526
527
528
529
530
531
532
533
534
535
536
537
538
539
540
541
542
543
544
545
546
547
548
549
550
551
552
553
554
555
556
557
558
559
560
561
562
563
564
565
566
567
568
569
570
571
572
573
574
575
576
577
578
579
580
581
582
583
584
585
586
587
588
589
590
591
592
593
594
595
596
597
598
599
600
601
602
603
604
605
606
607
608
609
610
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639
640
641
642
643
644
645
646
647
648
649
650
651
652
653
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
669
670
671
672
673
674
675
676
677
678
679
680
681
682
683
684
685
686
687
688
689
690
691
692
693
694
695
696
697
698
699
700
701
702
703
704
705
706
707
708
709
710
711
712
713
714
715
716
717
718
719
720
721
722
723
724
725
726
727
728
729
730
731
732
733
734
735
736
737
738
739
740
741
742
743
744
745
746
747
748
749
750
751
752
753
754
755
756
757
758
759
760
761
762
763
764
765
766
767
768
769
770
771
772
773
774
775
776
777
778
779
780
781
782
783
784
785
786
787
788
{
  "schema_version": "seqcolyte.spec.v1",
  "spec_id": "cel_seq2",
  "assay": "CEL-Seq2",
  "chemistry_version": "",
  "platform": "illumina",
  "platform_params": {
    "read_type": "short"
  },
  "source_docs": [
    {
      "doc_id": "CEL-seq2_detailed_protocol.docx",
      "title": "CEL-Seq2 detailed protocol (Additional file 4)",
      "url": "https://doi.org/10.1186/s13059-016-0938-8",
      "path": "/Users/seqmachines/playground/protocols-test/protocols/cel_seq2/CEL-seq2_detailed_protocol.docx",
      "retrieved_date": null
    },
    {
      "doc_id": "CEL-seq2_supp.pdf",
      "title": "CEL-seq2_supp.pdf",
      "url": null,
      "path": "/Users/seqmachines/playground/protocols-test/protocols/cel_seq2/CEL-seq2_supp.pdf",
      "retrieved_date": null
    },
    {
      "doc_id": "CEL-seq2.pdf",
      "title": "CEL-seq2.pdf",
      "url": null,
      "path": "/Users/seqmachines/playground/protocols-test/protocols/cel_seq2/CEL-seq2.pdf",
      "retrieved_date": null
    }
  ],
  "oligos": [
    {
      "oligo_id": "oligo_cel_seq2_rt_primer",
      "name": "CEL-Seq2 primer (barcoded RT primer)",
      "aliases": [],
      "role": "Barcoded reverse-transcription / capture primer with T7 promoter, Illumina 5' adapter, UMI, cell barcode and anchored poly(dT)",
      "kind": "assembled",
      "sequence": "GCCGGTAATACGACTCACTATAGGGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV",
      "direction": "5_to_3",
      "components": [
        {
          "name": "T7 promoter",
          "sequence": "GCCGGTAATACGACTCACTATAGGG",
          "role": "T7 RNA polymerase promoter (transcription start at the terminal GGG); GCCGG 5' leader"
        },
        {
          "name": "Partial Illumina 5' adapter (RA5)",
          "sequence": "AGTTCTACAGTCCGACGATC",
          "role": "Shortened Illumina small-RNA 5' adapter; becomes the Read 1 primer landing site"
        },
        {
          "name": "UMI",
          "sequence": "[UMI:6]",
          "role": "6-nt unique molecular identifier (NNNNNN), upstream of the barcode"
        },
        {
          "name": "Cell barcode",
          "sequence": "[CELL_BARCODE:6]",
          "role": "6-nt sample/cell barcode (Hamming distance >=2); one per well"
        },
        {
          "name": "Poly(dT)",
          "sequence": "TTTTTTTTTTTTTTTTTTTTTTTT",
          "role": "24-nt poly(dT) to prime poly(A) mRNA"
        },
        {
          "name": "Anchor (V)",
          "sequence": "V",
          "role": "3' anchor base (A/C/G) that seats the primer at the poly(A) junction"
        }
      ],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "82-nt primer (shortened from CEL-Seq's 92 nt). 96 barcoded variants are listed in the detailed protocol; e.g. barcode 1 = AGACTC, 4 = AGCTTC, 5 = CATGAG, 46 = TGCAGA. Recommended 10-primer pool: barcodes 1,4,5,9,10,23,25,26,31,46. 168 unique 6-nt barcodes were designed (GC 33-67%, last base != T)."
    },
    {
      "oligo_id": "oligo_cel_seq2_barcode_primer_1s",
      "name": "CEL-Seq2 primer 1s (barcode AGACTC)",
      "aliases": [],
      "role": "Representative fully-specified barcoded RT primer (well 1)",
      "kind": "single",
      "sequence": "GCCGGTAATACGACTCACTATAGGGAGTTCTACAGTCCGACGATCNNNNNNAGACTCTTTTTTTTTTTTTTTTTTTTTTTTV",
      "direction": "5_to_3",
      "components": [],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "NNNNNN = 6-nt UMI; AGACTC = 6-nt cell barcode for well 1. Concrete example of the CEL-Seq2 primer series (1s-96s) transcribed verbatim from the detailed protocol table."
    },
    {
      "oligo_id": "oligo_cel_seq_original_primer",
      "name": "CEL-Seq primer (original, 8-nt barcode)",
      "aliases": [],
      "role": "Original CEL-Seq barcoded RT primer (no UMI)",
      "kind": "assembled",
      "sequence": "CGATTGAGGCCGGTAATACGACTCACTATAGGGGTTCAGAGTTCTACAGTCCGACGATC[CELL_BARCODE:8]TTTTTTTTTTTTTTTTTTTTTTTTV",
      "direction": "5_to_3",
      "components": [
        {
          "name": "5' leader + T7 promoter",
          "sequence": "CGATTGAGGCCGGTAATACGACTCACTATAGGG",
          "role": "T7 promoter with extended 5' leader"
        },
        {
          "name": "Illumina 5' adapter (RA5)",
          "sequence": "GTTCAGAGTTCTACAGTCCGACGATC",
          "role": "Full-length Illumina small-RNA 5' adapter"
        },
        {
          "name": "Cell barcode",
          "sequence": "[CELL_BARCODE:8]",
          "role": "8-nt cell barcode (as previously published)"
        },
        {
          "name": "Poly(dT)",
          "sequence": "TTTTTTTTTTTTTTTTTTTTTTTT",
          "role": "24-nt poly(dT)"
        },
        {
          "name": "Anchor (V)",
          "sequence": "V",
          "role": "3' anchor base"
        }
      ],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Original CEL-Seq design (Table S2); 92 nt, longer T7 promoter and 5' adapter, no UMI."
    },
    {
      "oligo_id": "oligo_cel_seq_umi_primer",
      "name": "CEL-Seq + UMI primer (5-nt UMI, 6-nt barcode)",
      "aliases": [],
      "role": "Intermediate CEL-Seq primer with 5-nt UMI",
      "kind": "assembled",
      "sequence": "CGATTGAGGCCGGTAATACGACTCACTATAGGGGTTCAGAGTTCTACAGTCCGACGATC[UMI:5][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV",
      "direction": "5_to_3",
      "components": [
        {
          "name": "5' leader + T7 promoter",
          "sequence": "CGATTGAGGCCGGTAATACGACTCACTATAGGG",
          "role": "T7 promoter with extended 5' leader"
        },
        {
          "name": "Illumina 5' adapter (RA5)",
          "sequence": "GTTCAGAGTTCTACAGTCCGACGATC",
          "role": "Full-length Illumina small-RNA 5' adapter"
        },
        {
          "name": "UMI",
          "sequence": "[UMI:5]",
          "role": "5-nt UMI (NNNNN)"
        },
        {
          "name": "Cell barcode",
          "sequence": "[CELL_BARCODE:6]",
          "role": "6-nt cell barcode"
        },
        {
          "name": "Poly(dT)",
          "sequence": "TTTTTTTTTTTTTTTTTTTTTTTT",
          "role": "24-nt poly(dT)"
        },
        {
          "name": "Anchor (V)",
          "sequence": "V",
          "role": "3' anchor base"
        }
      ],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Table S2; the 'CEL-Seq performed as previously described with a 5-base UMI and 6-base barcode' variant used for comparison."
    },
    {
      "oligo_id": "oligo_library_rt_primer",
      "name": "Library RT primer (randomhexRT)",
      "aliases": [],
      "role": "Random-hexamer RT primer with 5'-tail Illumina 3' adapter; converts fragmented aRNA back to cDNA",
      "kind": "assembled",
      "sequence": "GCCTTGGCACCCGAGAATTCCANNNNNN",
      "direction": "5_to_3",
      "components": [
        {
          "name": "Illumina 3' adapter (RA3, rev-comp orientation)",
          "sequence": "GCCTTGGCACCCGAGAATTCCA",
          "role": "5'-tail matching Illumina small-RNA 3' adapter (rev-comp of TGGAATTCTCGGGTGCCAAGGC)"
        },
        {
          "name": "Random hexamer",
          "sequence": "NNNNNN",
          "role": "Random-priming 3' hexamer that anneals across fragmented aRNA"
        }
      ],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "CEL-Seq2 change 5: inserts the Illumina 3' adapter at the RT step via a random hexamer, eliminating the ligation step of the original CEL-Seq. Transcribed verbatim from the protocol/Table S2."
    },
    {
      "oligo_id": "oligo_rna_pcr_primer_rp1",
      "name": "RNA PCR Primer (RP1)",
      "aliases": [],
      "role": "Forward library PCR primer; adds P5 and completes the Illumina 5' adapter",
      "kind": "assembled",
      "sequence": "AATGATACGGCGACCACCGAGATCTACACGTTCAGAGTTCTACAGTCCGACGATC",
      "direction": "5_to_3",
      "components": [
        {
          "name": "P5",
          "sequence": "AATGATACGGCGACCACCGAGATCTACAC",
          "role": "Illumina P5 flow-cell adapter"
        },
        {
          "name": "Illumina 5' adapter (RA5)",
          "sequence": "GTTCAGAGTTCTACAGTCCGACGATC",
          "role": "Full 5' small-RNA adapter; 3' end anneals to the AGTTCTACAGTCCGACGATC tag in the fragment"
        }
      ],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Standard Illumina TruSeq Small-RNA RP1; the documents state 'sequences available from Illumina' and are not printed. Sequence assembled from the verified P5 constant plus the in-document 5' adapter tag."
    },
    {
      "oligo_id": "oligo_rna_pcr_index_primer_rpi",
      "name": "RNA PCR Index Primer (RPIX)",
      "aliases": [],
      "role": "Reverse indexed library PCR primer; adds i7 sample index and P7 via the Illumina 3' adapter",
      "kind": "assembled",
      "sequence": "CAAGCAGAAGACGGCATACGAGAT[SAMPLE_INDEX:6]GTGACTGGAGTTCCTTGGCACCCGAGAATTCCA",
      "direction": "5_to_3",
      "components": [
        {
          "name": "P7",
          "sequence": "CAAGCAGAAGACGGCATACGAGAT",
          "role": "Illumina P7 flow-cell adapter"
        },
        {
          "name": "i7 sample index",
          "sequence": "[SAMPLE_INDEX:6]",
          "role": "6-nt sample index (read in the 7-cycle index read)"
        },
        {
          "name": "Illumina 3' adapter (RA3, rev-comp)",
          "sequence": "GTGACTGGAGTTCCTTGGCACCCGAGAATTCCA",
          "role": "3' portion whose 3' end anneals to the RA3 tag added by the library RT primer"
        }
      ],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Standard Illumina TruSeq Small-RNA RPIX; not printed in the docs ('sequences from Illumina kit'). Assembled from the verified P7 constant, a 6-nt index, and the in-document 3' adapter. A uniquely indexed RPIX is added per pooled library."
    },
    {
      "oligo_id": "oligo_illumina_ra5_adapter",
      "name": "Illumina 5' small-RNA adapter (RA5)",
      "aliases": [],
      "role": "5' adapter / Read 1 primer landing region of the final library",
      "kind": "single",
      "sequence": "GTTCAGAGTTCTACAGTCCGACGATC",
      "direction": "5_to_3",
      "components": [],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "The shortened form AGTTCTACAGTCCGACGATC is carried on the CEL-Seq2 primer; full length restored by RP1 during PCR."
    },
    {
      "oligo_id": "oligo_illumina_ra3_adapter",
      "name": "Illumina 3' small-RNA adapter (RA3)",
      "aliases": [],
      "role": "3' adapter / Read 2 primer landing region of the final library",
      "kind": "single",
      "sequence": "TGGAATTCTCGGGTGCCAAGGC",
      "direction": "5_to_3",
      "components": [],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Reverse complement of the library RT primer 5'-tail (GCCTTGGCACCCGAGAATTCCA)."
    },
    {
      "oligo_id": "oligo_illumina_p5_adapter",
      "name": "Illumina P5 adapter",
      "aliases": [],
      "role": "P5 flow-cell binding sequence",
      "kind": "single",
      "sequence": "AATGATACGGCGACCACCGAGATCTACAC",
      "direction": "5_to_3",
      "components": [],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Standard Illumina constant; contributed by RP1."
    },
    {
      "oligo_id": "oligo_illumina_p7_adapter",
      "name": "Illumina P7 adapter",
      "aliases": [],
      "role": "P7 flow-cell binding sequence",
      "kind": "single",
      "sequence": "CAAGCAGAAGACGGCATACGAGAT",
      "direction": "5_to_3",
      "components": [],
      "provenance": "document",
      "derivation": null,
      "sequence_source": "llm_extracted_from_docs",
      "evidence": [
        {
          "source_doc": "protocol_docs",
          "locator": "oligo / final library",
          "method": "claude_llm_extraction"
        }
      ],
      "notes": "Standard Illumina constant; contributed by RPIX. Appears at the 3' end of the top strand as its reverse complement (ATCTCGTATGCCGTCTTCTGCTTG)."
    }
  ],
  "final_library": {
    "source_label": "CEL-Seq2 final Illumina small-RNA library (5'->3' top strand)",
    "annotated_library_sequence": "AATGATACGGCGACCACCGAGATCTACACGTTCAGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC[SAMPLE_INDEX:6]ATCTCGTATGCCGTCTTCTGCTTG",
    "library_sequence": "AATGATACGGCGACCACCGAGATCTACACGTTCAGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC[SAMPLE_INDEX:6]ATCTCGTATGCCGTCTTCTGCTTG",
    "strands": [
      {
        "direction": "5_to_3",
        "source_html": "AATGATACGGCGACCACCGAGATCTACACGTTCAGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC[SAMPLE_INDEX:6]ATCTCGTATGCCGTCTTCTGCTTG",
        "source_sequence": "AATGATACGGCGACCACCGAGATCTACACGTTCAGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC[SAMPLE_INDEX:6]ATCTCGTATGCCGTCTTCTGCTTG"
      }
    ],
    "annotation_lines": [
      "AATGATACGGCGACCACCGAGATCTACAC = P5",
      "GTTCAGAGTTCTACAGTCCGACGATC = Illumina 5' small-RNA adapter (RA5 / Read 1 primer site)",
      "[UMI:6] = UMI",
      "[CELL_BARCODE:6] = Cell barcode",
      "TTTTTTTTTTTTTTTTTTTTTTTT = poly(dT)",
      "V = anchor base",
      "[CDNA] = cDNA insert",
      "TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC = Illumina 3' small-RNA adapter (RA3 / Read 2 primer site)",
      "[SAMPLE_INDEX:6] = i7 sample index (reverse complement)",
      "ATCTCGTATGCCGTCTTCTGCTTG = reverse complement of P7"
    ],
    "evidence": [
      {
        "source_doc": "protocol_docs",
        "locator": "CEL-Seq2 final Illumina small-RNA library (5'->3' top strand)",
        "method": "claude_llm_extraction"
      }
    ]
  },
  "read_structure": {
    "reads": [
      {
        "read": "R1",
        "primer": "Illumina small-RNA Read 1 sequencing primer",
        "template": "bottom",
        "cycles": 15,
        "segments": [
          {
            "name": "UMI",
            "type": "umi",
            "order": 0,
            "scored": true,
            "provenance": null,
            "whitelist_ref": null,
            "constant_ref": null,
            "notes": null,
            "length": 6
          },
          {
            "name": "Cell barcode",
            "type": "barcode",
            "order": 1,
            "scored": true,
            "provenance": null,
            "whitelist_ref": null,
            "constant_ref": null,
            "notes": null,
            "length": 6
          },
          {
            "name": "poly(dT)",
            "type": "constant",
            "order": 2,
            "scored": false,
            "provenance": null,
            "whitelist_ref": null,
            "constant_ref": null,
            "notes": null,
            "length": 3
          }
        ]
      },
      {
        "read": "I1",
        "primer": "Illumina index sequencing primer",
        "template": "top",
        "cycles": 7,
        "segments": [
          {
            "name": "i7 sample index",
            "type": "index",
            "order": 0,
            "scored": false,
            "provenance": null,
            "whitelist_ref": null,
            "constant_ref": null,
            "notes": null,
            "length": 6
          }
        ]
      },
      {
        "read": "R2",
        "primer": "Illumina small-RNA Read 2 sequencing primer",
        "template": "top",
        "cycles": 36,
        "segments": [
          {
            "name": "cDNA insert",
            "type": "insert",
            "order": 0,
            "scored": true,
            "provenance": null,
            "whitelist_ref": null,
            "constant_ref": null,
            "notes": null
          }
        ]
      }
    ]
  },
  "library_generation": [
    {
      "step": 1,
      "title": "Anneal barcoded CEL-Seq2 primer & reverse transcription",
      "summary": "The barcoded poly(dT) CEL-Seq2 primer captures a single cell's poly(A) mRNA and SuperScript II reverse-transcribes first-strand cDNA.",
      "note": "Each cell/well gets a unique 6-nt barcode; the primer also carries a T7 promoter, the Illumina 5' adapter and a 6-nt UMI. SuperScript II extends from the anchored poly(dT) into the transcript body.",
      "product": "5'- GCCGGTAATACGACTCACTATAGGGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]------->\n                                                                    3'- AAAAAAAAAAAAAAAAAAAAAAAA...(mRNA)...-5'"
    },
    {
      "step": 2,
      "title": "Second-strand synthesis (dsDNA with T7 promoter)",
      "summary": "RNase H, E. coli DNA Pol I and DNA ligase convert the RNA:cDNA hybrid into double-stranded DNA carrying an intact T7 promoter.",
      "note": "SuperScript II Double-Stranded cDNA Synthesis Kit reagents. Barcoded samples are then pooled and bead-purified (AMPure XP) before amplification.",
      "product": "5'- GCCGGTAATACGACTCACTATAGGGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA] -3'\n3'- CGGCCATTATGCTGAGTGATATCCCTCAAGATGTCAGGCTGCTAG[UMI:6][CELL_BARCODE:6]AAAAAAAAAAAAAAAAAAAAAAAAB[CDNA] -5'"
    },
    {
      "step": 3,
      "title": "In vitro transcription (T7 linear amplification)",
      "summary": "T7 RNA polymerase transcribes from the promoter, linearly amplifying each molecule into many antisense aRNA copies.",
      "note": "IVT (MEGAscript/SuperScript II kit) avoids exponential PCR bias; transcription starts at the terminal GGG so the T7 promoter itself is not copied into the aRNA.",
      "product": "5'- GGGAGUUCUACAGUCCGACGAUC[UMI:6][CELL_BARCODE:6]UUUUUUUUUUUUUUUUUUUUUUUUV[CDNA] -3'   (aRNA, antisense to mRNA)"
    },
    {
      "step": 4,
      "title": "aRNA fragmentation",
      "summary": "The amplified RNA is chemically fragmented (Mg2+, heat) to ~200-500 nt.",
      "note": "Fragmentation is stopped with EDTA and the aRNA is bead-purified (RNAClean XP). Only fragments retaining the 5' adapter + UMI + barcode end carry the demultiplexing information.",
      "product": "5'- GGGAGUUCUACAGUCCGACGAUC[UMI:6][CELL_BARCODE:6]UUUU...V[CDNA frag] -3'   +   internal [CDNA] aRNA fragments"
    },
    {
      "step": 5,
      "title": "RT of aRNA with random-hexamer / 3'-adapter primer",
      "summary": "A random hexamer bearing a 5'-tail Illumina 3' adapter reverse-transcribes the fragmented aRNA into cDNA, appending the 3' adapter (ligation-free).",
      "note": "SuperScript II RT. This CEL-Seq2 change replaces the inefficient adapter ligation of the original CEL-Seq, improving read mapping (93.8% vs 60.9%).",
      "product": "5'- ...GAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]TGGAATTCTCGGGTGCCAAGGC -3'\n                                                                                  (Illumina 3' adapter from randomhexRT)"
    },
    {
      "step": 6,
      "title": "Library PCR (RP1 + RPIX) \u2014 final indexed library",
      "summary": "Phusion PCR with RP1 (adds P5 + full 5' adapter) and a uniquely indexed RPIX (adds i7 index + P7) produces the sequenceable Illumina small-RNA library.",
      "note": "11-15 cycles. Double AMPure XP cleanup; expected 200-400 bp peak. Handled as an Illumina Small-RNA library on HiSeq 2500 rapid mode.",
      "product": "5'- AATGATACGGCGACCACCGAGATCTACACGTTCAGAGTTCTACAGTCCGACGATC[UMI:6][CELL_BARCODE:6]TTTTTTTTTTTTTTTTTTTTTTTTV[CDNA]TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC[SAMPLE_INDEX:6]ATCTCGTATGCCGTCTTCTGCTTG -3'"
    }
  ],
  "library_sequencing": [
    {
      "read": "R1",
      "primer": "Illumina small-RNA Read 1 sequencing primer (= RA5 sense; anneals to the bottom strand, extends 5'->3')",
      "template": "bottom",
      "cycles": 15,
      "note": "Reads the 6-nt UMI then the 6-nt cell barcode (UMI is 5' of the barcode in the RT primer), followed by ~3 poly(dT) bases. The barcode demultiplexes wells; the UMI is carried onto the R2 molecule for counting.",
      "diagram": "5'- GTTCAGAGTTCTACAGTCCGACGATC----------------->   Read 1 primer (= RA5)\n5'- GTTCAGAGTTCTACAGTCCGACGATCNNNNNNNNNNNNTTTTTTTTTT...V[cDNA] -3'\n3'- CAAGTCTCAAGATGTCAGGCTGCTAGNNNNNNNNNNNNAAAAAAAAAA...B[cDNA] -5'\n                              ^^^^^^^^^^^^\n                              UMI(6) + cell barcode(6)  (read 5'->3')"
    },
    {
      "read": "I1",
      "primer": "Illumina i7 index sequencing primer (anneals over the P7 region of the top strand, extends 5'->3' toward the insert)",
      "template": "top",
      "cycles": 7,
      "note": "Reads the 6-nt i7 sample index (7 cycles run). The index is stored as its reverse complement on the top strand, so the read reproduces the designed index sequence.",
      "diagram": "                     <------ i7 index read (7 cycles)\n                        3'-TAGAGCATACGGCAGAAGACGAAC-5'   i7 index primer\n5'- ...GGAACTCCAGTCACNNNNNNATCTCGTATGCCGTCTTCTGCTTG -3'\n3'- ...CCTTGAGGTCAGTGNNNNNNTAGAGCATACGGCAGAAGACGAAC -5'\n                     ^^^^^^ i7 index (6 nt; stored as rev-comp on top strand)"
    },
    {
      "read": "R2",
      "primer": "Illumina small-RNA Read 2 sequencing primer (= RA3 antisense; anneals to the RA3 region of the top strand, extends 5'->3' into the cDNA)",
      "template": "top",
      "cycles": 36,
      "note": "Reads 36 bases of the 3'-biased cDNA insert (used for genome mapping); paired with the R1 UMI/barcode.",
      "diagram": "    <--------------------------- read (36 nt, into cDNA)\n          3'-ACCTTAAGAGCCCACGGTTCCTTGAGGTCAGTG-5'   Read 2 primer\n5'- ...[cDNA]TGGAATTCTCGGGTGCCAAGGAACTCCAGTCAC[i7]ATCTCGTATGCCGTCTTCTGCTTG -3'\n3'- ...[cDNA]ACCTTAAGAGCCCACGGTTCCTTGAGGTCAGTG[i7]TAGAGCATACGGCAGAAGACGAAC -5'"
    }
  ],
  "whitelists": {},
  "build": {
    "builder_version": "llm-generic-1.0",
    "deterministic": false,
    "source_html_sha256": null,
    "extraction_method": "claude_llm_generic",
    "model": "claude-opus-4-8"
  },
  "title": "CEL-Seq2",
  "description": "CEL-Seq2 is a 3'-end-tag, early-barcoding single-cell RNA-seq method that linearly amplifies transcripts by in vitro transcription (IVT). A barcoded RT primer carrying a T7 promoter, a shortened Illumina 5' (small-RNA) adapter, a 6-nt UMI and a 6-nt cell barcode captures poly(A) mRNA at the reverse-transcription step. After second-strand synthesis and T7 IVT (linear amplification), the amplified antisense RNA (aRNA) is fragmented, reverse-transcribed with a random hexamer bearing the Illumina 3' adapter, and PCR-amplified with the Illumina small-RNA RP1/RPI primers into a sequenceable library. Read 1 reads the UMI + cell barcode, Read 2 reads the 3'-biased cDNA, enabling accurate molecule counting across many multiplexed cells.",
  "reference": {
    "kind": "paper",
    "label": "CEL-Seq2 detailed protocol (Additional file 4)",
    "path": null,
    "url": "https://doi.org/10.1186/s13059-016-0938-8",
    "doi": "10.1186/s13059-016-0938-8"
  },
  "publication": {
    "year": 2016,
    "original_publication": {
      "title": "CEL-Seq2: sensitive highly-multiplexed single-cell RNA-Seq",
      "journal": "Genome Biology",
      "doi": "10.1186/s13059-016-0938-8",
      "url": "https://doi.org/10.1186/s13059-016-0938-8"
    },
    "authors": [
      {
        "name": "Tamar Hashimshony",
        "affiliation": "Department of Biology, Technion - Israel Institute of Technology, Haifa, Israel"
      },
      {
        "name": "Naftalie Senderovich",
        "affiliation": "Department of Biology, Technion - Israel Institute of Technology, Haifa, Israel"
      },
      {
        "name": "Gal Avital",
        "affiliation": "Department of Biology, Technion - Israel Institute of Technology, Haifa, Israel"
      },
      {
        "name": "Agnes Klochendler",
        "affiliation": "Department of Developmental Biology and Cancer Research, The Hebrew University-Hadassah Medical School, Jerusalem, Israel"
      },
      {
        "name": "Yaron de Leeuw",
        "affiliation": "Department of Biology, Technion - Israel Institute of Technology, Haifa, Israel"
      },
      {
        "name": "Leon Anavy",
        "affiliation": "Department of Biology, Technion - Israel Institute of Technology, Haifa, Israel"
      },
      {
        "name": "Dave Gennert",
        "affiliation": "Klarman Cell Observatory, Broad Institute of Harvard and MIT; Department of Biology, MIT; HHMI, Massachusetts Institute of Technology, Cambridge, MA, USA"
      },
      {
        "name": "Shuqiang Li",
        "affiliation": "Fluidigm Corporation, South San Francisco, CA, USA"
      },
      {
        "name": "Kenneth J. Livak",
        "affiliation": "Fluidigm Corporation, South San Francisco, CA, USA"
      },
      {
        "name": "Orit Rozenblatt-Rosen",
        "affiliation": "Klarman Cell Observatory, Broad Institute of Harvard and MIT; Department of Biology, MIT; HHMI, Massachusetts Institute of Technology, Cambridge, MA, USA"
      },
      {
        "name": "Yuval Dor",
        "affiliation": "Department of Developmental Biology and Cancer Research, The Hebrew University-Hadassah Medical School, Jerusalem, Israel"
      },
      {
        "name": "Aviv Regev",
        "affiliation": "Klarman Cell Observatory, Broad Institute of Harvard and MIT; Department of Biology, MIT; HHMI, Massachusetts Institute of Technology, Cambridge, MA, USA"
      },
      {
        "name": "Itai Yanai",
        "corresponding": true,
        "email": "yanai@technion.ac.il",
        "affiliation": "Department of Biology, Technion - Israel Institute of Technology, Haifa, Israel"
      }
    ],
    "throughput": {
      "summary": "Highly-multiplexed single cells: 24 fibroblasts (CEL-Seq), 20 (CEL-Seq2), 72 captured on the Fluidigm C1, and dendritic cells in a 384-well plate; 96 barcoded primers per run (168 6-nt barcodes designed).",
      "cells": "Up to 96 cells per plate / C1 run (168 unique barcodes available)"
    },
    "statistical_model": "Binomial statistics to convert UMI counts into transcript counts (UMI-based molecule counting; efficiency estimated by linear fit on ERCC spike-in log-log plots)",
    "other": [
      {
        "label": "RT/detection efficiency",
        "value": "19.7% manual, 22% on Fluidigm C1, vs 5.8% for original CEL-Seq (ERCC spike-in estimate)"
      },
      {
        "label": "Primer length",
        "value": "Shortened from 92 nt (CEL-Seq) to 82 nt while adding a 6-nt UMI"
      },
      {
        "label": "Ligation-free prep",
        "value": "Illumina adapter inserted at RT via random hexamer; raised barcoded-read mapping from 60.9% to 93.8%"
      },
      {
        "label": "GEO accession",
        "value": "GSE78779"
      },
      {
        "label": "Pipeline",
        "value": "https://github.com/yanailab/CEL-Seq-pipeline (GPLv3); demultiplex R1 barcode/UMI, Bowtie2 map, UMI-aware htseq-count"
      }
    ]
  },
  "modality": "RNA",
  "method_type": "plate-based",
  "data_processing": {
    "summary": "Read 1 is used to demultiplex the per-well cell barcode and extract the UMI; Read 2 (3'-biased cDNA) is mapped to the genome and reads are assigned to genes, then collapsed by UMI to give a per-cell molecule (UMI) count matrix.",
    "stages": [
      {
        "id": "demux",
        "label": "Barcode & UMI processing"
      },
      {
        "id": "align",
        "label": "Alignment"
      },
      {
        "id": "quant",
        "label": "Quantification"
      }
    ],
    "nodes": [
      {
        "id": "demux_bc",
        "label": "Demultiplex by cell barcode",
        "tool": "CEL-Seq-pipeline",
        "stage": "demux",
        "scope": "per_cell",
        "terminal": false,
        "viz_only": false
      },
      {
        "id": "extract_umi",
        "label": "Extract UMI",
        "tool": "CEL-Seq-pipeline",
        "stage": "demux",
        "scope": "per_cell",
        "terminal": false,
        "viz_only": false
      },
      {
        "id": "map_cdna",
        "label": "Map cDNA to genome",
        "tool": "Bowtie2",
        "stage": "align",
        "scope": "per_cell",
        "terminal": false,
        "viz_only": false
      },
      {
        "id": "assign_genes",
        "label": "Assign reads to genes",
        "tool": "htseq-count",
        "stage": "quant",
        "scope": "per_cell",
        "terminal": false,
        "viz_only": false
      },
      {
        "id": "collapse_umi",
        "label": "Collapse duplicate UMIs",
        "tool": "CEL-Seq-pipeline",
        "stage": "quant",
        "scope": "per_cell",
        "terminal": false,
        "viz_only": false
      },
      {
        "id": "count_matrix",
        "label": "Build UMI count matrix",
        "tool": "CEL-Seq-pipeline",
        "stage": "quant",
        "scope": "bulk",
        "terminal": true,
        "viz_only": false
      }
    ],
    "edges": [
      {
        "from": "demux_bc",
        "to": "map_cdna",
        "kind": "sequential"
      },
      {
        "from": "map_cdna",
        "to": "assign_genes",
        "kind": "sequential"
      },
      {
        "from": "assign_genes",
        "to": "collapse_umi",
        "kind": "sequential"
      },
      {
        "from": "extract_umi",
        "to": "collapse_umi",
        "kind": "sequential"
      },
      {
        "from": "collapse_umi",
        "to": "count_matrix",
        "kind": "fan_in"
      }
    ],
    "statistical_model": "Binomial (UMI-based molecule counting; observed UMIs converted to transcript counts, efficiency estimated from ERCC spike-ins)"
  }
}