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FamD external comparator, not FamF. RNA/ATAC do not measure the in-vivo state." }, { "donor": "M4", "blood": 3.8, "culture": 16.59, "published_KLF1_genotype": "wt/p.K288X", "group": "moderate", "family": "FamF", "note": "Cultured HEP RNA/ATAC; blood and culture HbF are separate published measurements." }, { "donor": "M5", "blood": 7.26, "culture": 21.52, "published_KLF1_genotype": "wt/p.K288X", "group": "moderate", "family": "FamF", "note": "Cultured HEP RNA/ATAC; blood and culture HbF are separate published measurements." }, { "donor": "M3", "blood": 0.6, "culture": 7.22, "published_KLF1_genotype": "wt/wt", "group": "normal", "family": "FamF", "note": "Cultured HEP RNA/ATAC; blood and culture HbF are separate published measurements. Published table labels wt/wt; older project notes conflict. 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"sample": "M9_T48", "donor": "M9", "time": "T48", "alt_fraction": 0.0, "KLF1_exon_over_reference": 0.1734999743, "informative_weight": 20.0, "reference_fraction": 1.0, "allele_scaled_KLF1_proxy": 0.1734999743 }, { "gate": "BQ20", "cap": "cap5", "ref_T": 20.0, "alt_A": 0.0, "other": 0.0, "alignment_signatures": 16, "direct_canonical_junction_linked_records": 0, "sample": "M9_T48", "donor": "M9", "time": "T48", "alt_fraction": 0.0, "KLF1_exon_over_reference": 0.2919816812, "informative_weight": 20.0, "reference_fraction": 1.0, "allele_scaled_KLF1_proxy": 0.2919816812 }, { "gate": "BQ20", "cap": "cap1", "ref_T": 16.0, "alt_A": 0.0, "other": 0.0, "alignment_signatures": 16, "direct_canonical_junction_linked_records": 0, "sample": "M9_T48", "donor": "M9", "time": "T48", "alt_fraction": 0.0, "KLF1_exon_over_reference": 0.3273715771, "informative_weight": 16.0, "reference_fraction": 1.0, "allele_scaled_KLF1_proxy": 0.3273715771 }, { "gate": "BQ30_internal5", "cap": "uncapped", "ref_T": 17.0, "alt_A": 0.0, "other": 0.0, "alignment_signatures": 14, "direct_canonical_junction_linked_records": 0, "sample": "M9_T48", "donor": "M9", "time": "T48", "alt_fraction": 0.0, "KLF1_exon_over_reference": 0.1734999743, "informative_weight": 17.0, "reference_fraction": 1.0, "allele_scaled_KLF1_proxy": 0.1734999743 }, { "gate": "BQ30_internal5", "cap": "cap5", "ref_T": 17.0, "alt_A": 0.0, "other": 0.0, "alignment_signatures": 14, "direct_canonical_junction_linked_records": 0, "sample": "M9_T48", "donor": "M9", "time": "T48", "alt_fraction": 0.0, "KLF1_exon_over_reference": 0.2919816812, "informative_weight": 17.0, "reference_fraction": 1.0, "allele_scaled_KLF1_proxy": 0.2919816812 }, { "gate": "BQ30_internal5", "cap": "cap1", "ref_T": 14.0, "alt_A": 0.0, "other": 0.0, "alignment_signatures": 14, "direct_canonical_junction_linked_records": 0, "sample": "M9_T48", "donor": "M9", "time": "T48", "alt_fraction": 0.0, "KLF1_exon_over_reference": 0.3273715771, "informative_weight": 14.0, "reference_fraction": 1.0, "allele_scaled_KLF1_proxy": 0.3273715771 } ], "family_summary": { "primary_family": "FamF", "high_donors": 1, "moderate_donors": 2, "M8_used_in_primary": false, "primary_RNA": [ { "view": "NH_weighted_firststrand", "reference": "combined9", "time": "T0", "control": "M4M5", "gamma_log2": 0.6968842034585245, "beta_deficit_log2": 0.46785032693014816, "total_log2": 1.1647345303886727, "gamma_fold": 1.6210001265198664, "HBB_fold": 0.7230411556238157, "ratio_fold": 2.241919583569654, "gamma_algebraic_share": 0.5983202053999153, "beta_algebraic_share": 0.4016797946000847 }, { "view": "NH_weighted_firststrand", "reference": "combined9", "time": "T48", "control": "M4M5", "gamma_log2": 0.6577459821128713, "beta_deficit_log2": 0.8275120971613585, "total_log2": 1.4852580792742298, "gamma_fold": 1.5776158836090728, "HBB_fold": 0.5635001512860563, "ratio_fold": 2.799672511193717, "gamma_algebraic_share": 0.44284962410995826, "beta_algebraic_share": 0.5571503758900417 } ], "local_time_projection": [ { "view": "NH_weighted_firststrand", "control": "M4M5", "time": "T0", "interval_shift": -0.7438370103286336, "equivalent_hours_local_linear": -35.70417649577441, "squared_projection_fraction": 0.7849769163603297, "residual_log2_norm": 0.8081410875324998, "contrast_log2_norm": 1.7427883367742398, "status": "Six-gene CLR local-line approximation, not calibrated biological age or a causal fraction" }, { "view": "NH_weighted_firststrand", "control": "M4M5", "time": "T48", "interval_shift": -0.5563604525155631, "equivalent_hours_local_linear": -26.70530172074703, "squared_projection_fraction": 0.27313114120496945, "residual_log2_norm": 1.8840567401984003, "contrast_log2_norm": 2.209866032336225, "status": "Six-gene CLR local-line approximation, not calibrated biological age or a causal fraction" } ], "RNA_ATAC": [ { "width": 1000, "time": "T0", "control": "M4M5", "RNA_ratio_log2_contrast": 1.1647345303886727, "ATAC_ratio_log2_contrast": 0.3873927388393841, "RNA_to_ATAC_ratio_discordance": 0.7773417915492886, "RNA_fold": 2.241919583569654, "ATAC_fold": 1.3080273788990087, "discordance_fold": 1.7139699212234534, "M2_gamma_fragments": 21.0, "M4_gamma_fragments": 26.0, "M5_gamma_fragments": 20.0 }, { "width": 1000, "time": "T48", "control": "M4M5", "RNA_ratio_log2_contrast": 1.48525807927423, "ATAC_ratio_log2_contrast": 0.28202107930877607, "RNA_to_ATAC_ratio_discordance": 1.203236999965454, "RNA_fold": 2.7996725111937173, "ATAC_fold": 1.2158970486022322, "discordance_fold": 2.302557206148463, "M2_gamma_fragments": 40.0, "M4_gamma_fragments": 43.0, "M5_gamma_fragments": 44.0 } ], "KLF1_proxy_lower_in_both_moderates_by_time": { "T0": 6, "T48": 5 }, "no_valid_family_causal_pvalues_computed": true }, "genetics": { "WGS_retained_keys": 3225, "WGS_complete_M2_M4_M5": 454, "WGS_M2_distinct_both": 148, "WGS_M2_distinct_with_father": 50, "SNP_total": 208261, "SNP_M2_distinct_both": 43236, "SNP_M2_distinct_all15trios_compatible": 33450, "KLF1_two_marker_conditional_phase": [ { "M2_haplotypes": [ [ "G", "T" ], [ "C", "G" ] ], "compatible_child_transmission": [ [ "G", "T" ], [ "G", "G" ] ] } ], "phase_assumptions": [ "Correct published pedigree and genotype labels", "No recombination between markers", "No direct molecular phase claimed" ], "published_rs3817621_GC_carrier_HbF": [ { "person": "II-5", "published_historical_HbF": 20.0 }, { "person": "II-1", "published_historical_HbF": 17.0 }, { "person": "II-7", "published_historical_HbF": 3.5 } ], "qPCR_people": 10, "qPCR_primary_M4_M5_present": false, "qPCR_timing_matched_to_RNAseq": false, "qPCR_ratio_issue": "cMC-HbF!F2:F21 divides FLVCR1a delta-Ct by FLVCR1b delta-Ct. This is not an expression ratio. Correct cross-person relative isoform proxy uses differences of Ct and assumes amplification efficiencies. Original workbook preserved.", "qPCR_additional_constraint": "cMC-HbF column has no verified unit/specimen-time definition here; it is not assumed to be HbF%.", "qPCR_formulas_verified": 20, "KLF1_three_marker_conditional_phase": { "marker_order": [ "rs3817621", "source_rs112943513", "K288X_rs267607202" ], "M2_haplotypes": [ [ "C", "G", "WT" ], [ "G", "T", "K288X" ] ], "maternal_haplotype_in_both_daughters": [ "G", "T", "K288X" ], "paternal_intact_haplotype_in_both_daughters": [ "G", "G", "WT" ], "intact_maternal_haplotype": [ "C", "G", "WT" ], "scope": "Conditional Mendelian inference with no crossover/gene conversion, correct pedigree and calls. Not direct molecular phasing; promoter variants remain linked-haplotype candidates." } }, "arrays": [ { "gene": "KLF1", "M2_person": "II-5", "comparator": "III-9", "log2_RMA_difference": 0.2977082178, "fold_RMA": 1.2291902408, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "BCL11A", "M2_person": "II-5", "comparator": "III-9", "log2_RMA_difference": 0.5529487107, "fold_RMA": 1.467081187, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "ZBTB7A", "M2_person": "II-5", "comparator": "III-9", "log2_RMA_difference": -0.039759678, "fold_RMA": 0.9728169843, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "SOX6", "M2_person": "II-5", "comparator": "III-9", "log2_RMA_difference": 0.6672773728, "fold_RMA": 1.5880731558, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "KLF1", "M2_person": "II-5", "comparator": "III-2", "log2_RMA_difference": -0.0561707542, "fold_RMA": 0.9618136098, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "BCL11A", "M2_person": "II-5", "comparator": "III-2", "log2_RMA_difference": 0.0289426255, "fold_RMA": 1.0202640836, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "ZBTB7A", "M2_person": "II-5", "comparator": "III-2", "log2_RMA_difference": -0.4398209763, "fold_RMA": 0.7372260852, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "SOX6", "M2_person": "II-5", "comparator": "III-2", "log2_RMA_difference": 0.1211221894, "fold_RMA": 1.0875804998, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "KLF1", "M2_person": "II-5", "comparator": "III-4", "log2_RMA_difference": 0.0348167005, "fold_RMA": 1.0244266577, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "BCL11A", "M2_person": "II-5", "comparator": "III-4", "log2_RMA_difference": -0.0669894636, "fold_RMA": 0.9546279861, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "ZBTB7A", "M2_person": "II-5", "comparator": "III-4", "log2_RMA_difference": -0.1125975673, "fold_RMA": 0.9249212443, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" }, { "gene": "SOX6", "M2_person": "II-5", "comparator": "III-4", "log2_RMA_difference": 0.110482974, "fold_RMA": 1.0795895925, "scope": "Historical CEL-derived accepted RMA; aliquot/time unmatched; no M4/M5 replication" } ], "development": { "version": "2026-10-23", "purpose": "Measured developmental-source panels for within-study comparisons; not a fetal-to-cord-to-adult calibrated trajectory.", "genes": [ "HBE1", "HBG2", "HBG1", "BGLT3", "HBBP1", "HBD", "HBB", "KLF1", "BCL11A", "ZBTB7A", "SOX6", "MYB", "LIN28B", "HIC2", "LDB1", "GATA1", "TAL1", "NR2C1", "NR2C2" ], "aliases": { "ZBTB7A": "LRF", "HBG2": "G\u03b3", "HBG1": "A\u03b3" }, "relative_value_definition": "Each source-normalized sample value divided by geometric mean of adult samples for the same gene, dataset and culture day. Calculated display normalization, not an additional measurement.", "limitations": [ "Culture day is erythroid maturation time, not gestational or postnatal age.", "Different studies are not pooled, cross-normalized, or stitched into one quantitative developmental curve.", "Adult references are dataset-specific and not equivalent to family M3/M7/M9.", "RNA values do not measure protein, transcription-factor activity, chromatin occupancy or HbF protein fraction.", "Sample labels establish libraries, not necessarily distinct donors or across-day pairing.", "Gene-count scales and FPKM scales are not interchangeable; compare relative values only within each selected study/day.", "BGLT3/HBBP1 RNA expression does not measure their local cis-regulatory DNA functions.", "GSE131809 adult LIN28B RNA counts are all zero; relative-to-adult folds are null, with exact values retained and no pseudocount inserted. Zero counts do not establish biological absence." ], "datasets": [ { "id": "GSE102182", "title": "Fetal liver vs adult erythroblasts", "geo_url": "https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE102182", "paper_url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC5647940/", "paper_doi": "10.1101/gad.303461.117", "build": "hg19", "comparison": "Fetal-liver versus adult-peripheral-blood CD34-derived erythroblasts, within matched culture day.", "culture_days": [ 11, 14 ], "samples_per_source_day": 2, "independent_donor_count_established": false, "input_file": "/workspace/scratch/910e18fd7392/graded_pattern_audit/legacy/nested/HPFH_Defensibility_v3_Reproducible/HPFH_Defensibility_v3/development/GSE102182_Gene_Values.csv", "input_sha256": "49381078ec8d8106af11f9a0cc1e570a6cc544f781071433235c0fdc8987594c", "normalization": "Supplied ERCC-normalized FPKM; STAR hg19. No fresh external raw-read realignment.", "gene_annotation_note": "BGLT3 was CTD-2643I7.1 / ENSG00000260629.1 in the source matrix; official Ensembl stable-ID bridge retained.", "records": [ { "gene": "KLF1", "sample": "FL.11.1", "geo_sample": "GSM2730041", "source": "fetal", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 395.065900810104, "adult_reference_geomean": 281.26144283068396, "relative_to_adult_geomean": 1.4046216105345408 }, { "gene": "KLF1", "sample": "FL.11.2", "geo_sample": "GSM2730042", "source": "fetal", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 301.541776796348, "adult_reference_geomean": 281.26144283068396, "relative_to_adult_geomean": 1.0721049204667294 }, { "gene": "KLF1", "sample": "FL.14.1", "geo_sample": "GSM2730043", "source": "fetal", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 217.384247742575, "adult_reference_geomean": 153.2782920074999, "relative_to_adult_geomean": 1.4182324508935578 }, { "gene": "KLF1", "sample": "FL.14.2", "geo_sample": "GSM2730044", "source": "fetal", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 139.072135382614, "adult_reference_geomean": 153.2782920074999, "relative_to_adult_geomean": 0.9073178827945786 }, { "gene": "KLF1", "sample": "AB.11.1", "geo_sample": "GSM2730037", "source": "adult", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 263.826820638252, "adult_reference_geomean": 281.26144283068396, "relative_to_adult_geomean": 0.938012754194227 }, { "gene": "KLF1", "sample": "AB.11.2", "geo_sample": "GSM2730038", "source": "adult", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 299.848207365041, "adult_reference_geomean": 281.26144283068396, "relative_to_adult_geomean": 1.0660835852482846 }, { "gene": "KLF1", "sample": "AB.14.1", "geo_sample": "GSM2730039", "source": "adult", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 126.771122089655, "adult_reference_geomean": 153.2782920074999, "relative_to_adult_geomean": 0.8270650750952527 }, { "gene": "KLF1", "sample": "AB.14.2", "geo_sample": "GSM2730040", "source": "adult", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 185.327970703934, "adult_reference_geomean": 153.2782920074999, "relative_to_adult_geomean": 1.2090947013871076 }, { "gene": "BCL11A", "sample": "FL.11.1", "geo_sample": "GSM2730041", "source": "fetal", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 4.71076710620029, "adult_reference_geomean": 9.709709647492716, "relative_to_adult_geomean": 0.48516045043805456 }, { "gene": "BCL11A", "sample": "FL.11.2", "geo_sample": "GSM2730042", "source": "fetal", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 2.49277027690725, "adult_reference_geomean": 9.709709647492716, "relative_to_adult_geomean": 0.2567296415038471 }, { "gene": "BCL11A", "sample": "FL.14.1", "geo_sample": "GSM2730043", "source": "fetal", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 1.6493528190243, "adult_reference_geomean": 4.785613150675545, "relative_to_adult_geomean": 0.34464817090187805 }, { "gene": "BCL11A", "sample": "FL.14.2", "geo_sample": "GSM2730044", "source": "fetal", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 0.622833377991169, "adult_reference_geomean": 4.785613150675545, "relative_to_adult_geomean": 0.13014703829607474 }, { "gene": "BCL11A", "sample": "AB.11.1", "geo_sample": "GSM2730037", "source": "adult", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 9.37788524151018, "adult_reference_geomean": 9.709709647492716, "relative_to_adult_geomean": 0.9658255068349834 }, { "gene": "BCL11A", "sample": "AB.11.2", "geo_sample": "GSM2730038", "source": "adult", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 10.053275233237, "adult_reference_geomean": 9.709709647492716, "relative_to_adult_geomean": 1.0353837136451347 }, { "gene": "BCL11A", "sample": "AB.14.1", "geo_sample": "GSM2730039", "source": "adult", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 3.61032006372716, "adult_reference_geomean": 4.785613150675545, "relative_to_adult_geomean": 0.7544111799378355 }, { "gene": "BCL11A", "sample": "AB.14.2", "geo_sample": "GSM2730040", "source": "adult", "culture_day": 14, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 6.34350772886198, "adult_reference_geomean": 4.785613150675545, "relative_to_adult_geomean": 1.325537089843236 }, { "gene": "ZBTB7A", "sample": "FL.11.1", "geo_sample": "GSM2730041", "source": "fetal", "culture_day": 11, "assay": "RNA-seq", "unit": "ERCC-normalized FPKM", "value": 12.4621616984549, "adult_reference_geomean": 8.863481751106628, 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"unit": "Median-ratio normalized RNA count", "value": 81702.76138895733, "raw_count": 103188.0, "adult_reference_geomean": 82415.72017744671, "relative_to_adult_geomean": 0.9913492379007993 }, { "gene": "HBD", "sample": "Adult4-RNA", "geo_sample": "GSM3820035", "source": "adult", "culture_day": 7, "assay": "RNA-seq", "unit": "Median-ratio normalized RNA count", "value": 83457.9237658451, "raw_count": 101346.0, "adult_reference_geomean": 82415.72017744671, "relative_to_adult_geomean": 1.012645689270863 }, { "gene": "HBD", "sample": "CordBlood1-RNA", "geo_sample": "GSM3820036", "source": "cord", "culture_day": 7, "assay": "RNA-seq", "unit": "Median-ratio normalized RNA count", "value": 16766.1953406458, "raw_count": 16187.0, "adult_reference_geomean": 82415.72017744671, "relative_to_adult_geomean": 0.20343443343754117 }, { "gene": "HBD", "sample": "CordBlood2-RNA", "geo_sample": "GSM3820037", "source": "cord", "culture_day": 7, "assay": "RNA-seq", "unit": "Median-ratio normalized RNA count", "value": 14751.861475771631, "raw_count": 15514.0, "adult_reference_geomean": 82415.72017744671, "relative_to_adult_geomean": 0.17899329695851543 }, { "gene": "HBD", "sample": "CordBlood3-RNA", "geo_sample": "GSM3820038", "source": "cord", "culture_day": 7, "assay": "RNA-seq", "unit": "Median-ratio normalized RNA count", "value": 18136.221084189223, "raw_count": 18438.0, "adult_reference_geomean": 82415.72017744671, "relative_to_adult_geomean": 0.2200577880668966 }, { "gene": "HBD", "sample": "CordBlood4-RNA", "geo_sample": "GSM3820039", "source": "cord", "culture_day": 7, "assay": "RNA-seq", "unit": "Median-ratio normalized RNA count", "value": 17656.653372144094, "raw_count": 16352.0, "adult_reference_geomean": 82415.72017744671, "relative_to_adult_geomean": 0.21423890168196194 } ], "relative_unavailable_genes": [ "LIN28B" ] } ], "validation": { "total_records": 304, "unique_sample_gene_within_dataset": true, "nonnegative_finite_source_values": true, "zero_adult_reference_handled_as_null": true, "generated_from_existing_accepted_panels": true } }, "modern": { "scope": "Selected primary mechanistic evidence through 2026-10-23; not an exhaustive systematic review. Literature edges are not automatically demonstrated in the Maltese samples.", "evidence_rule": "Solid edges may mean causal perturbation evidence in the named published system. Use a different indicator for observations in the Maltese family; absence of family occupancy data must not be hidden by literature evidence.", "edges": [ { "id": "klf1_bcl_hbb", "from": "KLF1", "to": "BCL11A and HBB", "effect": "activation", "level": "causal, human family and erythroid perturbation", "finding": "KLF1 haploinsufficiency disrupts the adult globin program; KLF1 activates BCL11A and adult HBB. This provides an established carrier-background mechanism.", "limit": "Does not by itself identify why one K288X carrier has higher HbF than another.", "source": "Borg et al. Nature Genetics 2010", "doi": "10.1038/ng.630", "url": "https://pubmed.ncbi.nlm.nih.gov/20676099/", "ref": "turn189search1" }, { "id": "klf1_lrf", "from": "KLF1", "to": "ZBTB7A/LRF", "effect": "activation", "level": "direct binding and perturbation", "finding": "KLF1 directly activates ZBTB7A through its proximal promoter in erythroid cells.", "limit": "LRF RNA abundance is not equivalent to its protein concentration, DNA occupancy or repressor-complex activity.", "source": "Norton et al. Blood Advances 2017", "doi": "10.1182/bloodadvances.2016002303", "url": "https://pubmed.ncbi.nlm.nih.gov/29296711/", "ref": "turn189search0" }, { "id": "bcl_promoter", "from": "BCL11A", "to": "HBG promoters", "effect": "repression", "level": "direct binding, protein-domain and promoter-editing evidence", "finding": "BCL11A binds the distal TGACCA element near -115 and directly represses gamma promoters; perturbing the site relieves repression.", "limit": "Motif presence or ATAC accessibility alone does not demonstrate BCL11A occupancy in a family sample.", "source": "Liu et al. Cell 2018", "doi": "10.1016/j.cell.2018.03.016", "url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC5889339/", "ref": "turn193search2" }, { "id": "lrf_promoter", "from": "ZBTB7A/LRF", "to": "HBG promoters", "effect": "repression", "level": "direct binding and natural-variant recreation", "finding": "LRF binds near -200; naturally occurring HPFH promoter mutations can prevent its binding and increase gamma expression. BCL11A-associated HPFH mutations cluster near -115.", "limit": "Not every promoter HPFH allele works by removing a repressor; some create activator sites.", "source": "Martyn et al. Nature Genetics 2018", "doi": "10.1038/s41588-018-0085-0", "url": "https://pubmed.ncbi.nlm.nih.gov/29610478/", "ref": "turn207search2" }, { "id": "lcr_loop", "from": "LCR contact promoted by tethered LDB1", "to": "HBG transcription", "effect": "activation", "level": "causal engineered looping", "finding": "Forced LCR-gamma contact activates gamma and reduces beta expression in primary adult human erythroblasts; related embryonic-gene experiments were performed in mice.", "limit": "An artificial looping perturbation establishes capacity, not a measured contact trajectory for each Maltese donor.", "source": "Deng et al. Cell 2014", "doi": "10.1016/j.cell.2014.05.050", "url": "https://pubmed.ncbi.nlm.nih.gov/25126789/", "ref": "turn204view2" }, { "id": "competition", "from": "Adult HBB promoter activity", "to": "LCR availability for HBG", "effect": "competition", "level": "causal promoter mutagenesis and contact assays", "finding": "Disabling the HBB promoter increases HBG expression and redirects LCR contacts. Removing coding sequence while retaining the promoter has a much weaker effect.", "limit": "Competition is a major component; enhancer rearrangement, local silencing and deletion geometry also matter. Do not equate low HBB RNA with promoter deletion.", "source": "Topfer et al. Blood 2022", "doi": "10.1182/blood.2021014205", "url": "https://pubmed.ncbi.nlm.nih.gov/35090172/", "ref": "turn193search0", "geo": "GSE194064" }, { "id": "hbbp1_dna", "from": "HBBP1 DNA region", "to": "Adult repressive locus architecture", "effect": "supports repression", "level": "developmental contact comparison and deletion", "finding": "Removing the HBBP1 region shifts adult erythroid chromatin contacts toward a fetal-like configuration, increases LCR-gamma contact and raises gamma transcription.", "limit": "DNA deletion and RNA depletion are different perturbations; this evidence cannot assign a repressive sign to HBBP1 RNA abundance.", "source": "Huang et al. Genes & Development 2017", "doi": "10.1101/gad.303461.117", "url": "https://pubmed.ncbi.nlm.nih.gov/28916711/", "ref": "turn193search4" }, { "id": "hbbp1_rna", "from": "HBBP1 RNA / ELK1 pathway", "to": "HBG expression", "effect": "activation in tested models", "level": "cell-model overexpression, interaction and promoter assays", "finding": "HBBP1 overexpression increased gamma RNA and protein in HUDEP-2; the study implicates ELK1, which also binds and activates the gamma promoter.", "limit": "Keep distinct from HBBP1 DNA's architectural role. This does not establish the sign of changing endogenous HBBP1 RNA in every developmental context.", "source": "Ma et al. Biochemical and Biophysical Research Communications 2021", "doi": "10.1016/j.bbrc.2021.03.051", "url": "https://pubmed.ncbi.nlm.nih.gov/33744764/", "ref": "turn205search7" }, { "id": "bglt3", "from": "BGLT3 locus transcription and RNA", "to": "HBG transcription", "effect": "activation", "level": "deletion, RNA depletion, overexpression and looping assays", "finding": "Both transcription through BGLT3 and its RNA support gamma expression. Locus transcription supports local looping; the RNA interacts with Mediator. Overexpression increased gamma in K562.", "limit": "RNA alone did not rescue deletion or transcription blockade. Epsilon was not reduced by BGLT3 deletion. Do not label BGLT3 a universal embryonic-globin switch.", "source": "Ivaldi et al. Blood 2018", "doi": "10.1182/blood-2018-07-862003", "url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC6213316/", "ref": "turn183search0" }, { "id": "bglt3_repression", "from": "BCL11A/LRF, G9a and CHD4", "to": "BGLT3", "effect": "repression", "level": "human erythroid knockout, knockdown, drug and chromatin experiments", "finding": "G9a-dependent chromatin repression and CHD4 recruitment connect BCL11A/LRF with BGLT3 repression. BGLT3 knockdown attenuated gamma induction after repressor knockout. BGLT3 overexpression increased gamma in HUDEP-2.", "limit": "The authors' universal wording refers to tested systems and inducers. RK-701 mouse embryonic-globin induction is not direct human BGLT3 gene-therapy evidence.", "source": "Takase et al. Nature Communications 2023", "doi": "10.1038/s41467-022-35404-0", "url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC9837035/", "ref": "turn185search0", "archive": "DRA014673" }, { "id": "hif_bglt3", "from": "HIF1alpha/HIF1beta", "to": "BGLT3 regulatory DNA", "effect": "activation", "level": "binding, base-edited motifs, primary-cell expression and contact assays", "finding": "HIF1 binds two BGLT3 hypoxia-response motifs, recruits activators, opens chromatin and promotes LCR-gamma interactions. Disrupting the motifs impairs gamma induction.", "limit": "HIF1A RNA is not a proxy for stabilized HIF1alpha protein. This pathway did not generate a uniform embryonic program: HBZ decreased in the VHL-disruption study.", "source": "Feng et al. Nature 2022", "doi": "10.1038/s41586-022-05312-w", "url": "https://pubmed.ncbi.nlm.nih.gov/36224385/", "ref": "turn188search0" }, { "id": "distance", "from": "Enhancer proximity", "to": "HBG/HBE/HBZ activation", "effect": "activation after engineered rearrangement", "level": "causal deletion/inversion in adult erythroid cells and ex-vivo differentiated HSPCs", "finding": "Bringing strong enhancers closer through intervening-sequence deletion or inversion can override developmental silencing at beta-like and alpha-like globin loci.", "limit": "This is not evidence of such rearrangements in the Maltese family and does not identify a single shared transcription factor.", "source": "Felder et al. Blood 2025", "doi": "10.1182/blood.2024028128", "url": "https://pubmed.ncbi.nlm.nih.gov/40456162/", "ref": "turn197search4", "geo": "GSE274030; GSE274029" }, { "id": "lrf_reversible", "from": "LRF protein loss and restoration", "to": "HBG repression state", "effect": "reversible", "level": "controlled protein degradation and restoration in HUDEP-2", "finding": "Gamma activation after LRF depletion can reverse when LRF returns, including RNA and chromatin changes; LRF-NuRD association persists without BCL11A.", "limit": "Demonstrated in immature erythroid models; not a direct measurement of family T0-to-T48 mechanism.", "source": "Miyazaki et al. Blood Red Cells & Iron 2026", "doi": "10.1016/j.brci.2026.100087", "url": "https://www.sciencedirect.com/science/article/pii/S3050598426000429", "ref": "turn197search0", "geo": "GSE309998; GSE309999" }, { "id": "igf2bp1", "from": "IGF2BP1", "to": "HIC2/BCL11A and HBG RNA translation", "effect": "supports fetal hemoglobin", "level": "human erythroid perturbation and RNA-binding/translation assays", "finding": "IGF2BP1 supports HIC2, a BCL11A repressor, and directly supports HBG1/2 translation through stop-codon-proximal m6A-containing RNA elements.", "limit": "A new 2026 mechanistic layer; family mediation is untested. This specifically cautions against equating RNA fractions with hemoglobin protein fractions.", "source": "Coyne et al. Blood 2026", "doi": "10.1182/blood.2025031840", "url": "https://pubmed.ncbi.nlm.nih.gov/42341077/", "ref": "turn205search1", "geo": "GSE303945", "publication_date": "2026-09-24" } ], "optional_sources": [ { "source": "Masuda et al. Science 2016", "doi": "10.1126/science.aad3312", "url": "https://pubmed.ncbi.nlm.nih.gov/26816381/", "finding": "LRF and BCL11A independently repress fetal hemoglobin; avoids treating LRF as merely a downstream BCL11A component.", "ref": "turn195search2" }, { "source": "Martyn et al. Blood 2019", "doi": "10.1182/blood-2018-07-863951", "url": "https://pubmed.ncbi.nlm.nih.gov/30617196/", "finding": "The -113A>G HPFH allele creates an activating GATA1 site without disrupting BCL11A binding; natural HPFH can add activation as well as remove repression.", "ref": "turn207search0" }, { "source": "Rahaman et al. Life Science Alliance 2026", "doi": "10.26508/lsa.202603620", "url": "https://pubmed.ncbi.nlm.nih.gov/42373543/", "finding": "UCA1/miR-148b modulates BCL11A and gamma in HUDEP-2 and CD34-derived cells; optional additional RNA regulatory layer, not a Maltese-family finding.", "ref": "turn197search3" } ], "family_interpretation": "KLF1 K288X supplies an established background affecting the adult program. BGLT3 is a plausible local gamma amplifier because the family RNA pattern is graded and independent studies demonstrate function. Greater BGLT3 could be both a readout of reduced repression and a mediator. M2 versus M4/M5 cannot currently distinguish those roles or establish ZFP57 interaction. Known cell perturbations justify mechanism examples, not fitted family-specific causal coefficients.", "deletion_caution": "Do not claim Antoniani's 13.6-kb HBD/HBB deletion proves BGLT3 dispensability: the BGLT3/HBBP1 region is retained. Hb Kenya is a potential natural counterexample involving deletion between Agamma and beta and elevated intact Ggamma, but verify its breakpoints and hybrid transcript context before using it to argue against universal necessity." }, "history": { "prepared_for": "HBB interactive locus model; historical evidence and scientific lineage", "scope": "Curated discovery timeline: legacy haemoglobin research, the Augusta\u2013Malta collaborations, Joseph Borg\u2019s verified globin publications and complementary mechanistic studies. Not an exhaustive systematic review.", "key_interpretations": [ "HBG2 encodes G\u03b3 and HBG1 encodes A\u03b3. Their relative contributions change with development and genetic background; a single undifferentiated gamma curve hides informative biology.", "HbF percent of total hemoglobin, G\u03b3 percent within HbF, F-cell percentage, and the fraction of a hybrid hemoglobin such as Hb Kenya are distinct measurements and must never share an unlabeled axis.", "The historical evidence supports multiple routes to persistent gamma expression: trans regulation, promoter variants, large rearrangements, and culture-dependent erythroid states. It does not support one universal obligate upstream molecule.", "The 1980 paper coauthored by Huisman and Felice directly warns against equating cultured erythroid output with circulating blood. It does not identify the explanation for M8 specifically.", "For Hb Kenya, the demonstrated A\u03b3\u2013\u03b2 fusion and deletion spanning 3\u2032 A\u03b3 through delta to 5\u2032 beta imply removal of the intervening region on the rearranged chromosome. Including BGLT3 in that removed interval is an annotation/gene-order inference; BGLT3 was not assayed in these historical papers.", "Hb Kenya heterozygotes still have the other chromosome. The historical studies cannot establish complete cellular absence of BGLT3 RNA, nor do the cited phenotype measurements prove the allele of origin of retained G\u03b3 expression. Avoid calling this proof that all BGLT3 activity is dispensable.", "Use a schematic ~22\u201323 kb Kenya deletion. Retrieved IthaGenes precise coordinates and stated size were inconsistent, and historical protein-junction conventions differ; do not hard-code an unverified base-pair breakpoint." ], "sources": [ { "id": "history_pmid_13909332", "pmid": "13909332", "year": 1962, "title": "Further sequences in the gamma-chain of human fetal hemoglobin", "authors": [ "SCHROEDER WA", "SHELTON JR", "SHELTON JB", "CORMICK J" ], "author_string": "SCHROEDER WA, SHELTON JR, SHELTON JB, CORMICK J.", "journal": "Proceedings of the National Academy of Sciences of the United States of America", "journal_abbreviation": "Proc Natl Acad Sci U S A", "volume": "48", "issue": null, "pages": "284-287", "doi": "10.1073/pnas.48.2.284", "url": "https://pubmed.ncbi.nlm.nih.gov/13909332/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/13909332/", "https://europepmc.org/article/MED/13909332" ], "headline": "Reading the fetal gamma-chain sequence", "finding": "Schroeder and colleagues reported additional amino-acid sequences in the fetal gamma chain.", "model_relevance": "Protein chemistry established the molecular substrate for later developmental comparisons.", "limitation": "Sequence determination alone does not identify developmental regulation.", "timeline_type": "supporting", "tags": [ "W. A. Schroeder" ], "felice_author": false, "huisman_author": false, "schroeder_author": true, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "research-article", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_14087393", "pmid": "14087393", "year": 1963, "title": "THE AMINO ACID SEQUENCE OF THE GAMMA CHAIN OF HUMAN FETAL HEMOGLOBIN", "authors": [ "SCHROEDER WA", "SHELTON JR", "SHELTON JB", "CORMICK J", "JONES RT" ], "author_string": "SCHROEDER WA, SHELTON JR, SHELTON JB, CORMICK J, JONES RT.", "journal": "Biochemistry", "journal_abbreviation": "Biochemistry", "volume": "2", "issue": null, "pages": "992-1008", "doi": "10.1021/bi00905a016", "url": "https://pubmed.ncbi.nlm.nih.gov/14087393/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/14087393/", "https://europepmc.org/article/MED/14087393" ], "headline": "The gamma-chain sequence", "finding": "Schroeder and colleagues determined the amino-acid sequence of human fetal gamma globin.", "model_relevance": "A foundational structural reference for distinguishing fetal globin chains and variants.", "limitation": "A sequence is not a regulatory mechanism.", "timeline_type": "milestone", "tags": [ "W. A. Schroeder" ], "felice_author": false, "huisman_author": false, "schroeder_author": true, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_5248810", "pmid": "5248810", "year": 1968, "title": "Evidence for multiple structural genes for the gamma chain of human fetal hemoglobin", "authors": [ "Schroeder WA", "Huisman TH", "Shelton JR", "Shelton JB", "Kleihauer EF", "Dozy AM", "Robberson B" ], "author_string": "Schroeder WA, Huisman TH, Shelton JR, Shelton JB, Kleihauer EF, Dozy AM, Robberson B.", "journal": "Proceedings of the National Academy of Sciences of the United States of America", "journal_abbreviation": "Proc Natl Acad Sci U S A", "volume": "60", "issue": "2", "pages": "537-544", "doi": "10.1073/pnas.60.2.537", "url": "https://pubmed.ncbi.nlm.nih.gov/5248810/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/5248810/", "https://europepmc.org/article/MED/5248810", "https://authors.library.caltech.edu/records/xqpep-v1q60" ], "headline": "Two gamma-chain products", "finding": "Schroeder, Huisman and colleagues found glycine or alanine at gamma-chain residue 136, supporting multiple nonallelic structural genes.", "model_relevance": "Established the G\u03b3/A\u03b3 distinction underlying separate HBG2 and HBG1 outputs.", "limitation": "The protein evidence predates the complete DNA sequence and modern locus annotation.", "timeline_type": "milestone", "tags": [ "T. H. J. Huisman", "W. A. Schroeder" ], "felice_author": false, "huisman_author": true, "schroeder_author": true, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "research-article", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_5792729", "pmid": "5792729", "year": 1969, "title": "Haemoglobin F(Malta): a new foetal haemoglobin variant with a high incidence in Maltese infants", "authors": [ "Cauchi MN", "Clegg JB", "Weatherall DJ" ], "author_string": "Cauchi MN, Clegg JB, Weatherall DJ.", "journal": "Nature", "journal_abbreviation": "Nature", "volume": "223", "issue": "5203", "pages": "311-313", "doi": "10.1038/223311a0", "url": "https://pubmed.ncbi.nlm.nih.gov/5792729/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/5792729/", "https://europepmc.org/article/MED/5792729" ], "headline": "Hb F-Malta enters the literature", "finding": "Cauchi, Clegg and Weatherall described Hb F(Malta) in Maltese cord blood at an unusually appreciable frequency.", "model_relevance": "Maltese newborn screening supplied a natural marker for fetal-globin biology.", "limitation": "Felice was not an author of this 1969 discovery; attribution belongs to its listed authors.", "timeline_type": "milestone", "tags": [ "Malta" ], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "huisman_1970_greek_hpfh", "year": 1970, "title": "Nature of fetal hemoglobin in the Greek type of hereditary persistence of fetal hemoglobin with and without concurrent beta-thalassemia", "authors": [ "Huisman TH", "Schroeder WA", "Stamatoyannopoulos G", "Bouver N", "Shelton JR", "Shelton JB", "Apell G" ], "journal": "The Journal of clinical investigation", "doi": "10.1172/jci106303", "pmid": "5441539", "url": "https://www.jci.org/articles/view/106303", "pubmed_url": "https://pubmed.ncbi.nlm.nih.gov/5441539/", "web_refs": [ "turn194search1", "turn194search4", "turn194search6" ], "finding": "In three Greek families, HPFH heterozygotes had A\u03b3-only fetal hemoglobin, whereas concurrent beta-thalassemia increased HbF and introduced G\u03b3 as well.", "model_relevance": "Separate HBG1 and HBG2 and represent genotype-dependent cis context rather than a single gamma output.", "limitation": "Protein composition and family inheritance suggested chromosome-specific contributions; the study did not identify modern regulatory factors or molecular chromatin contacts.", "huisman_author": true, "felice_author": false, "borg_author": false, "schroeder_author": true, "volume": "49", "issue": "5", "pages": "1035-1040", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/5441539/", "https://europepmc.org/article/MED/5441539" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [ "T. H. J. Huisman", "W. A. Schroeder" ], "indexed_publication_types": [ "research-article", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": false }, { "id": "schroeder_1971_postnatal", "year": 1971, "title": "Postnatal Changes in the Chemical Heterogeneity of Human Fetal Hemoglobin", "authors": [ "W A Schroeder", "T H J Huisman", "Audrey K Brown", "Nicole G Bouver", "P O Lerch", "J Roger Shelton", "Joan Balog Shelton", "Gerald Apell" ], "journal": "Pediatric Research. 1971;5:493", "doi": "10.1203/00006450-197109000-00008", "url": "https://www.nature.com/articles/pr1971360", "web_refs": [ "turn212search5" ], "finding": "Among 11 infants of varied genotypes and 16 adults, newborn G\u03b3:A\u03b3 was about 3:1 and adult composition about 2:3, with a marked postnatal change around months three to four.", "model_relevance": "A developmental animation should show changing composition within gamma output as well as the overall gamma-to-beta switch.", "limitation": "These are G\u03b3:A\u03b3 protein ratios within HbF, not HbF as a percentage of total hemoglobin; small heterogeneous cohorts do not define a universal exact developmental curve.", "usa_context": "Caltech, Pasadena; Medical College of Georgia and Veterans Administration Hospital, Augusta.", "huisman_author": true, "felice_author": false, "borg_author": false, "schroeder_author": true, "timeline_type": "milestone", "source_urls": [ "https://www.nature.com/articles/pr1971360" ], "verified_at": "2026-10-23", "tags": [ "T. H. J. Huisman", "W. A. Schroeder" ], "joseph_borg_author": false, "verification_note": "Exact author list and developmental interpretation rechecked against Pediatric Research publisher page.", "indexed_publication_types": [], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": false }, { "id": "history_pmid_5050916", "pmid": "5050916", "year": 1972, "title": "Evidence for four nonallelic structural genes for the chain of human fetal hemoglobin", "authors": [ "Huisman TH", "Schroeder WA", "Bannister WH", "Grech JL" ], "author_string": "Huisman TH, Schroeder WA, Bannister WH, Grech JL.", "journal": "Biochemical genetics", "journal_abbreviation": "Biochem Genet", "volume": "7", "issue": "2", "pages": "131-139", "doi": "10.1007/bf00486084", "url": "https://pubmed.ncbi.nlm.nih.gov/5050916/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/5050916/", "https://europepmc.org/article/MED/5050916", "https://link.springer.com/article/10.1007/BF00486084" ], "headline": "Maltese variants challenge early gene models \u2014 historical hypothesis", "finding": "Huisman, Schroeder, Bannister and Grech used fetal variant proportions, including Malta variants, to propose an early four-gene interpretation.", "model_relevance": "Shows how Maltese observations informed\u2014and tested\u2014the developing globin model.", "limitation": "The four-nonallelic-gene hypothesis is historical and is not the modern normal HBG1/HBG2 locus map. Do not present it as current fact.", "timeline_type": "historical-hypothesis", "tags": [ "T. H. J. Huisman", "W. A. Schroeder" ], "felice_author": false, "huisman_author": true, "schroeder_author": true, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "interpretation_status": "Superseded four-nonallelic-gamma-gene hypothesis; retain only with explicit historical qualification.", "landmark": false, "borg_author": false, "seminal": false }, { "id": "huisman_1972_kenya", "year": 1972, "title": "Hemoglobin Kenya, the product of fusion of \u03b3 and \u03b2 polypeptide chains", "authors": [ "Huisman TH", "Wrightstone RN", "Wilson JB", "Schroeder WA", "Kendall AG" ], "journal": "Archives of biochemistry and biophysics", "doi": "10.1016/0003-9861(72)90408-0", "pmid": "4676910", "url": "https://pubmed.ncbi.nlm.nih.gov/4676910/", "web_refs": [ "turn198search0" ], "finding": "The original structural study identified Hb Kenya as a gamma\u2013beta fusion hemoglobin.", "model_relevance": "Introduce a separate hybrid protein output and a rearranged locus rather than displaying Kenya as ordinary elevated HbF alone.", "limitation": "Protein characterization does not provide a verified modern genome assembly breakpoint or a BGLT3 functional assay; PubMed's Greek-letter rendering of the title is corrupted.", "huisman_author": true, "felice_author": false, "borg_author": false, "schroeder_author": true, "volume": "153", "issue": "2", "pages": "850-853", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/4676910/", "https://europepmc.org/article/MED/4676910" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "title_note": "Greek symbols restored from publisher/references because PubMed rendering is corrupted.", "tags": [ "T. H. J. Huisman", "W. A. Schroeder" ], "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": true }, { "id": "history_pmid_4512230", "pmid": "4512230", "year": 1972, "title": "World-wide occurrence of nonallelic genes for the -chain of human foetal haemoglobin in newborns", "authors": [ "Schroeder WA", "Shelton JR", "Shelton JB", "Apell G", "Huisman TH", "Bouver NG" ], "author_string": "Schroeder WA, Shelton JR, Shelton JB, Apell G, Huisman TH, Bouver NG.", "journal": "Nature: New biology", "journal_abbreviation": "Nat New Biol", "volume": "240", "issue": "104", "pages": "273-274", "doi": "10.1038/newbio240273a0", "url": "https://pubmed.ncbi.nlm.nih.gov/4512230/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/4512230/", "https://europepmc.org/article/MED/4512230" ], "headline": "Gamma heterogeneity across populations", "finding": "Schroeder and Huisman\u2019s collaborators extended evidence for nonallelic gamma-chain products to newborn samples from multiple populations.", "model_relevance": "Places the G\u03b3/A\u03b3 distinction in an international developmental context.", "limitation": "Historical protein analysis does not map all regulatory DNA.", "timeline_type": "milestone", "tags": [ "T. H. J. Huisman", "W. A. Schroeder" ], "felice_author": false, "huisman_author": true, "schroeder_author": true, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "nute_1976_kenya_phenotype", "year": 1976, "title": "The Kenya form of hereditary persistence of fetal haemoglobin: structural studies and evidence for homogeneous distribution of haemoglobin F using fluorescent anti-haemoglobin F antibodies", "authors": [ "Nute PE", "Wood WG", "Stamatoyannopoulos G", "Olweny C", "Failkow PJ" ], "author_note": "Last author spelling follows PubMed metadata as retrieved.", "journal": "British journal of haematology", "doi": "10.1111/j.1365-2141.1976.tb01875.x", "pmid": "1259926", "url": "https://pubmed.ncbi.nlm.nih.gov/1259926/", "web_refs": [ "turn213search1", "turn212search1", "turn210search0" ], "finding": "Ugandan Hb Kenya heterozygotes had 20.68\u201323.35% Hb Kenya and 5\u20138% true HbF, exclusively G\u03b3; a specific fluorescent antibody demonstrated HbF in all red cells.", "model_relevance": "The retained gamma product is HBG2-derived G\u03b3, and Hb Kenya percentage is not the HbF percentage.", "limitation": "Heterozygotes retain the second chromosome; the study did not measure BGLT3 or directly establish the allele of origin of G\u03b3 transcripts.", "huisman_author": false, "felice_author": false, "inclusion_reason": "Independent primary phenotype evidence required for interpreting Huisman's Kenya structural work.", "borg_author": false, "schroeder_author": false, "volume": "32", "issue": "1", "pages": "55-63", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/1259926/", "https://europepmc.org/article/MED/1259926" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article", "Case Reports" ], "publication_type": "Case report", "publication_type_basis": "PubMed/Europe PMC case-report indexing; may include original molecular characterization.", "seminal": false }, { "id": "history_pmid_834242", "pmid": "834242", "year": 1977, "title": "Anomaly in the gamma chain heterogeneity of the newborn", "authors": [ "Huisman TH", "Schroeder WA", "Felice A", "Powars D", "Ringelhann B" ], "author_string": "Huisman TH, Schroeder WA, Felice A, Powars D, Ringelhann B.", "journal": "Nature", "journal_abbreviation": "Nature", "volume": "265", "issue": "5589", "pages": "63-65", "doi": "10.1038/265063a0", "url": "https://pubmed.ncbi.nlm.nih.gov/834242/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/834242/", "https://europepmc.org/article/MED/834242", "https://www.nature.com/articles/265063a0" ], "headline": "Gamma-chain diversity in newborns", "finding": "Huisman, Schroeder, Felice and colleagues documented newborns with unusually high G\u03b3:A\u03b3 ratios in a study of nearly 700 cord-blood samples.", "model_relevance": "Shows that composition within HbF varies between individuals before the adult switch.", "limitation": "G\u03b3:A\u03b3 is a ratio within HbF; it is not total HbF percentage.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice", "T. H. J. Huisman", "W. A. Schroeder" ], "felice_author": true, "huisman_author": true, "schroeder_author": true, "joseph_borg_author": false, "verified_affiliations": [ { "author": "A. Felice", "affiliation": "Department of Physiology and Biochemistry, Royal University of Malta, Msida, Malta" }, { "author": "T. H. J. Huisman", "affiliation": "Laboratory of Protein Chemistry and Comprehensive Sickle Cell Center, Medical College of Georgia, Augusta, Georgia" }, { "author": "W. A. Schroeder", "affiliation": "California Institute of Technology, Pasadena, California" } ], "verification_note": "Metadata and abstract checked against publisher page; Felice is listed with Malta affiliation, not Augusta.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_588240", "pmid": "588240", "year": 1977, "title": "Detection and quantitation of the fetal hemoglobin variant Hb F-Malta-I in adults", "authors": [ "Altay G", "Garver F", "Bannister WH", "Grech JL", "Felice A", "Huisman TH" ], "author_string": "Altay G, Garver F, Bannister WH, Grech JL, Felice A, Huisman TH.", "journal": "Biochemical genetics", "journal_abbreviation": "Biochem Genet", "volume": "15", "issue": "9-10", "pages": "915-923", "doi": "10.1007/bf00483988", "url": "https://pubmed.ncbi.nlm.nih.gov/588240/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/588240/", "https://europepmc.org/article/MED/588240" ], "headline": "Hb F-Malta-I persists into adulthood", "finding": "Sensitive immunoassay detected the Maltese G\u03b3 variant in adult blood, showing that expression of its gene was reduced after birth rather than completely extinguished.", "model_relevance": "A naturally occurring Maltese protein marker traces developmental G\u03b3 expression.", "limitation": "Very low protein abundance does not establish its upstream transcriptional regulator.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_750552", "pmid": "750552", "year": 1978, "title": "Delta beta-thalassemia in a Mexican family: clinical differences among homozygotes", "authors": [ "Reyes GR", "Pi\u00f1a-Camara A", "Felice AE", "Gravely ME", "Huisman TH" ], "author_string": "Reyes GR, Pi\u00f1a-Camara A, Felice AE, Gravely ME, Huisman TH.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "2", "issue": "6", "pages": "513-529", "doi": "10.3109/03630267809005353", "url": "https://pubmed.ncbi.nlm.nih.gov/750552/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/750552/", "https://europepmc.org/article/MED/750552" ], "headline": "A shared delta-beta defect, different severity", "finding": "Three homozygotes in a Mexican family differed substantially in clinical severity and globin-chain imbalance despite sharing a delta-beta-thalassemia classification.", "model_relevance": "Early family evidence that a globin genotype alone need not determine the full phenotype.", "limitation": "The proposed F-cell precursor explanation remained a hypothesis.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_747178", "pmid": "747178", "year": 1978, "title": "Is the trimodality of Hb Leslie (alpha 2 beta 2 131 Gln---O) in heterozygotes the result of a variable number of active alpha-chain genes? Evidence for posttranslational control of hemoglobin synthesis", "authors": [ "Felice A", "Abraham EC", "Miller A", "Stallings M", "Huisman TH" ], "author_string": "Felice A, Abraham EC, Miller A, Stallings M, Huisman TH.", "journal": "American journal of hematology", "journal_abbreviation": "Am J Hematol", "volume": "5", "issue": "1", "pages": "1-9", "doi": "10.1002/ajh.2830050102", "url": "https://pubmed.ncbi.nlm.nih.gov/747178/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/747178/", "https://europepmc.org/article/MED/747178" ], "headline": "Globin abundance also reflects protein assembly", "finding": "Studies of Hb Leslie carriers connected alpha-chain availability with differential incorporation and degradation of variant beta chains.", "model_relevance": "Distinguishes measured hemoglobin proportions from transcription alone.", "limitation": "These protein-assembly observations are not evidence for a common transcriptional switch factor.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_681817", "pmid": "681817", "year": 1978, "title": "Variability in the interaction of beta-thalassemia with the alpha-chain variants Hb G-Philadelphia and Hb Rampa", "authors": [ "Huisman TH", "Gravely ME", "Henson J", "Felice A", "Wilson JB", "Abraham EC", "Vella F", "Little MW" ], "author_string": "Huisman TH, Gravely ME, Henson J, Felice A, Wilson JB, Abraham EC, Vella F, Little MW.", "journal": "The Journal of laboratory and clinical medicine", "journal_abbreviation": "J Lab Clin Med", "volume": "92", "issue": "2", "pages": "311-320", "doi": null, "url": "https://pubmed.ncbi.nlm.nih.gov/681817/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/681817/", "https://europepmc.org/article/MED/681817" ], "headline": "Alpha\u2013beta interactions alter variant output", "finding": "Family and recombination studies showed that beta-chain deficiency affected Hb Rampa and Hb G-Philadelphia differently.", "model_relevance": "A quantitative model should allow chain assembly to alter the protein readout.", "limitation": "Variant-specific assembly effects cannot be generalized to all hemoglobins.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article", "Case Reports" ], "publication_type": "Case report", "publication_type_basis": "PubMed/Europe PMC case-report indexing; may include original molecular characterization.", "borg_author": false, "seminal": false }, { "id": "history_pmid_438612", "pmid": "438612", "year": 1979, "title": "Hb P-Nilotic in association with beta0-thalassemia: cis-mutation of a hemoglobin betaA chain regulatory determinant?", "authors": [ "Abu-Sin A", "Felice AE", "Gravely ME", "Wilson JB", "Reese AL", "Miller HL", "Huisman TH" ], "author_string": "Abu-Sin A, Felice AE, Gravely ME, Wilson JB, Reese AL, Miller HL, Huisman TH.", "journal": "The Journal of laboratory and clinical medicine", "journal_abbreviation": "J Lab Clin Med", "volume": "93", "issue": "6", "pages": "973-982", "doi": null, "url": "https://pubmed.ncbi.nlm.nih.gov/438612/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/438612/", "https://europepmc.org/article/MED/438612" ], "headline": "A hybrid locus suggests cis regulation", "finding": "A family carrying Hb P-Nilotic and beta-zero-thalassemia motivated a model involving altered regulation of the beta gene on the hybrid-bearing chromosome.", "model_relevance": "Connects rearranged globin loci with the distinction between cis and trans effects.", "limitation": "The regulatory-sequence mechanism was proposed from family and biosynthesis data, not demonstrated by modern perturbation.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_474574", "pmid": "474574", "year": 1979, "title": "The association of sickle cell anemia with heterozygous and homozygous alpha-thalassemia-2: in vitro HB chain synthesis", "authors": [ "Felice AE", "Webber B", "Miller A", "Mayson SM", "Harris HF", "Henson JB", "Gravely ME", "Huisman TH" ], "author_string": "Felice AE, Webber B, Miller A, Mayson SM, Harris HF, Henson JB, Gravely ME, Huisman TH.", "journal": "American journal of hematology", "journal_abbreviation": "Am J Hematol", "volume": "6", "issue": "2", "pages": "91-106", "doi": "10.1002/ajh.2830060202", "url": "https://pubmed.ncbi.nlm.nih.gov/474574/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/474574/", "https://europepmc.org/article/MED/474574" ], "headline": "Alpha-thalassemia modifies sickle-cell biosynthesis", "finding": "Felice and colleagues investigated in-vitro globin-chain synthesis in sickle-cell anemia with heterozygous or homozygous alpha-thalassemia.", "model_relevance": "Adds a cross-locus modifier study to the Felice\u2013Huisman research programme.", "limitation": "PubMed metadata supplies no abstract; no additional quantitative claim is made here.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Historical Article", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "huisman_felice_1980_culture", "year": 1980, "title": "Adult and fetal hemoglobin production in erythroid colonies from subjects with beta-thalassemia or with hereditary persistance of fetal hemoglobin (HPFH)", "title_note": "PubMed indexes 'persistance'; readable title normalized to 'persistence'.", "authors": [ "Huisman TH", "Reese AL", "Gravely ME", "Wilson JB", "Webber B", "Felice AE" ], "journal": "Hemoglobin", "doi": "10.3109/03630268008996226", "pmid": "6158501", "url": "https://pubmed.ncbi.nlm.nih.gov/6158501/", "web_refs": [ "turn213search0", "turn198search1" ], "finding": "Cultured colonies from beta-thalassemia heterozygotes averaged 12.6% HbF compared with 12.2% in normal adult colonies; the authors also documented culture-related effects on globin-chain balance.", "model_relevance": "Display culture state separately from the subject's circulating blood phenotype; do not equate M8 HEP RNA/ATAC with in-vivo HbF.", "limitation": "Historical culture conditions differ from modern HEP protocols, and this paper does not establish the specific cause of the M8 signal.", "huisman_author": true, "felice_author": true, "borg_author": false, "schroeder_author": false, "volume": "4", "issue": "3-4", "pages": "449-467", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6158501/", "https://europepmc.org/article/MED/6158501" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": false }, { "id": "history_pmid_7383748", "pmid": "7383748", "year": 1980, "title": "Hb S, Hb G-Philadelphia and alpha-thalassemia-2 in a Black family", "authors": [ "Felice AE", "Mayson SM", "Webber BB", "Miller A", "Gravely ME", "Huisman TH" ], "author_string": "Felice AE, Mayson SM, Webber BB, Miller A, Gravely ME, Huisman TH.", "journal": "Pediatric research", "journal_abbreviation": "Pediatr Res", "volume": "14", "issue": "3", "pages": "266-267", "doi": "10.1203/00006450-198003000-00019", "url": "https://pubmed.ncbi.nlm.nih.gov/7383748/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/7383748/", "https://europepmc.org/article/MED/7383748" ], "headline": "Development changes the phenotype of a shared genotype", "finding": "A family with Hb S, Hb G-Philadelphia and alpha-thalassemia showed age-dependent differences in chain synthesis and red-cell indices.", "model_relevance": "Developmental stage matters when comparing a cord-blood sample with later blood.", "limitation": "A small family report does not establish a population-wide developmental curve.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_7295286", "pmid": "7295286", "year": 1981, "title": "Alpha-thalassemia and the production of different alpha chain variants in heterozygotes", "authors": [ "Felice AE", "Webber BB", "Huisman TH" ], "author_string": "Felice AE, Webber BB, Huisman TH.", "journal": "Biochemical genetics", "journal_abbreviation": "Biochem Genet", "volume": "19", "issue": "5-6", "pages": "487-498", "doi": "10.1007/bf00484621", "url": "https://pubmed.ncbi.nlm.nih.gov/7295286/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/7295286/", "https://europepmc.org/article/MED/7295286" ], "headline": "Gene dosage and assembly both shape alpha variants", "finding": "Biosynthesis and protein studies distinguished alpha-gene dosage effects from impaired assembly of particular alpha-chain variants.", "model_relevance": "Connects the alpha locus to quantitative hemoglobin phenotypes without equating all effects with transcription.", "limitation": "The work does not prove a shared developmental controller of the alpha and beta loci.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_6163753", "pmid": "6163753", "year": 1981, "title": "Clinical and hematological evaluation of two delta 0 beta 0-thalassemia homozygotes", "authors": [ "Dincol G", "Altay C", "Aksoy M", "Gurgey A", "Felice AE", "Huisman TH" ], "author_string": "Dincol G, Altay C, Aksoy M, Gurgey A, Felice AE, Huisman TH.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "5", "issue": "2", "pages": "153-164", "doi": "10.3109/03630268108996921", "url": "https://pubmed.ncbi.nlm.nih.gov/6163753/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6163753/", "https://europepmc.org/article/MED/6163753" ], "headline": "Gamma composition distinguishes delta-beta thalassemias", "finding": "Clinical and biosynthetic comparison of two delta-beta-zero-thalassemia homozygotes showed that different gamma compositions could accompany severe chain imbalance.", "model_relevance": "High HbF, balanced synthesis and benign HPFH are distinct outcomes.", "limitation": "Two homozygotes do not define all deletional thalassemia or HPFH phenotypes.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article", "Case Reports" ], "publication_type": "Case report", "publication_type_basis": "PubMed/Europe PMC case-report indexing; may include original molecular characterization.", "borg_author": false, "seminal": false }, { "id": "history_pmid_6166190", "pmid": "6166190", "year": 1981, "title": "In vitro synthesis of hemoglobin and hemoglobin chains in the BFUe-derived colonies form person with alpha- or beta-thalassemia", "authors": [ "Huisman TH", "Reese AL", "Webber B", "Okonjo K", "Altay C", "Felice AE" ], "author_string": "Huisman TH, Reese AL, Webber B, Okonjo K, Altay C, Felice AE.", "journal": "American journal of hematology", "journal_abbreviation": "Am J Hematol", "volume": "10", "issue": "3", "pages": "227-237", "doi": "10.1002/ajh.2830100302", "url": "https://pubmed.ncbi.nlm.nih.gov/6166190/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6166190/", "https://europepmc.org/article/MED/6166190" ], "headline": "Cultured colonies differ from circulating reticulocytes", "finding": "In thalassemia and variant carriers, BFUe-derived colonies showed more balanced chain synthesis and increased HbF and HbA2 compared with reticulocytes.", "model_relevance": "An early direct reason to keep culture state separate from in-vivo phenotype.", "limitation": "The authors proposed rapid proteolysis as an explanation; this was not a measurement of the present family or modern HEP cultures.", "timeline_type": "milestone", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_7258153", "pmid": "7258153", "year": 1981, "title": "The occurrence and identification of alpha-thalassemia-2 among hemoglobin S heterozygotes", "authors": [ "Felice AE", "Altay CA", "Milner PF", "Huisman TH" ], "author_string": "Felice AE, Altay CA, Milner PF, Huisman TH.", "journal": "American journal of clinical pathology", "journal_abbreviation": "Am J Clin Pathol", "volume": "76", "issue": "1", "pages": "70-73", "doi": "10.1093/ajcp/76.1.70", "url": "https://pubmed.ncbi.nlm.nih.gov/7258153/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/7258153/", "https://europepmc.org/article/MED/7258153" ], "headline": "Alpha-gene dosage modifies HbS proportions", "finding": "A study of HbS carriers related chain-synthesis balance, cell indices and HbS proportions to inferred active alpha-gene number.", "model_relevance": "Supports an explicit alpha-chain dosage layer in globin interpretation.", "limitation": "The authors cautioned that overlapping biosynthesis values prevented reliable diagnosis in individual cases.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_6175208", "pmid": "6175208", "year": 1982, "title": "Identification and quantitation of embryonic and three types of fetal hemoglobin produced on induction of the human pluripotent leukemia cell line K-562 with hemin", "authors": [ "Fuhr JE", "Bamberger E", "Lozzio CB", "Lozzio BB", "Felice AE", "Altay G", "Webber BB", "Reese AL", "Mayson SM", "Huisman TH" ], "author_string": "Fuhr JE, Bamberger E, Lozzio CB, Lozzio BB, Felice AE, Altay G, Webber BB, Reese AL, Mayson SM, Huisman TH.", "journal": "American journal of hematology", "journal_abbreviation": "Am J Hematol", "volume": "12", "issue": "1", "pages": "1-12", "doi": "10.1002/ajh.2830120102", "url": "https://pubmed.ncbi.nlm.nih.gov/6175208/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6175208/", "https://europepmc.org/article/MED/6175208" ], "headline": "Embryonic and fetal globins in K562 cells", "finding": "Hemin-induced K562 cells produced embryonic Gower and Portland hemoglobins alongside HbF, with several gamma-chain types.", "model_relevance": "Embryonic and fetal products can coexist in an experimental erythroid-like state.", "limitation": "K562 is a leukemia cell line; its mixed globin output is not a normal fetal-liver or family HEP programme.", "timeline_type": "milestone", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_6291506", "pmid": "6291506", "year": 1982, "title": "Organization of alpha-chain genes among Hb G-Philadelphia heterozygotes in association with Hb S, beta-thalassemia, and alpha-thalassemia-2", "authors": [ "Felice AE", "Ozdonmez R", "Headlee ME", "Huisman TH" ], "author_string": "Felice AE, Ozdonmez R, Headlee ME, Huisman TH.", "journal": "Biochemical genetics", "journal_abbreviation": "Biochem Genet", "volume": "20", "issue": "7-8", "pages": "689-701", "doi": "10.1007/bf00483966", "url": "https://pubmed.ncbi.nlm.nih.gov/6291506/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6291506/", "https://europepmc.org/article/MED/6291506" ], "headline": "Mapping alpha-gene organization", "finding": "Felice and colleagues mapped alpha-chain gene organization in Hb G-Philadelphia carriers with other globin genotypes.", "model_relevance": "Moves from protein proportions toward the underlying gene arrangement.", "limitation": "PubMed supplies no abstract; detailed breakpoint or causal claims require the original article.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "ojwang_1983_kenya_deletion", "year": 1983, "title": "Gene deletion as the molecular basis for the Kenya-G gamma-HPFH condition", "authors": [ "Ojwang PJ", "Nakatsuji T", "Gardiner MB", "Reese AL", "Gilman JG", "Huisman TH" ], "journal": "Hemoglobin", "doi": "10.3109/03630268309048641", "pmid": "6200457", "url": "https://pubmed.ncbi.nlm.nih.gov/6200457/", "web_refs": [ "turn192search2" ], "finding": "Restriction mapping supported an approximately 22.5-kb deletion from the 3\u2032 A\u03b3 region through delta to the 5\u2032 beta region, consistent with the Hb Kenya fusion model.", "model_relevance": "A naturally rearranged locus changes the gamma\u2013delta intergenic interval and adult-gene configuration; intact-locus assumptions cannot simply be carried over.", "limitation": "Historical mapping is insufficient for exact modern assembly coordinates, and the paper did not assay BGLT3 or identify the allele of origin of retained G\u03b3 expression.", "huisman_author": true, "felice_author": false, "borg_author": false, "schroeder_author": false, "volume": "7", "issue": "2", "pages": "115-123", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6200457/", "https://europepmc.org/article/MED/6200457" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [ "T. H. J. Huisman" ], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": true }, { "id": "history_pmid_6526651", "pmid": "6526651", "year": 1984, "title": "Alternate organization of alpha G-Philadelphia globin genes among U.S. black and Italian Caucasian heterozygotes", "authors": [ "Sciarratta GV", "Sansone G", "Ivaldi G", "Felice AE", "Huisman TH" ], "author_string": "Sciarratta GV, Sansone G, Ivaldi G, Felice AE, Huisman TH.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "8", "issue": "6", "pages": "537-547", "doi": "10.3109/03630268408991740", "url": "https://pubmed.ncbi.nlm.nih.gov/6526651/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6526651/", "https://europepmc.org/article/MED/6526651" ], "headline": "Alpha-gene organization changes variant dosage", "finding": "Mapping showed different cis arrangements of the Hb G-Philadelphia gene in different families and related organization to variant abundance.", "model_relevance": "Illustrates how locus structure changes quantitative protein output.", "limitation": "Ethnic labels in the original title are historical; the result concerns gene organization, not biological racial categories.", "timeline_type": "supporting", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, Non-U.S. Gov't", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "padanilam_felice_1984_hbb", "year": 1984, "title": "Partial deletion of the 5' beta-globin gene region causes beta zero-thalassemia in members of an American black family", "authors": [ "Padanilam BJ", "Felice AE", "Huisman TH" ], "journal": "Blood", "pmid": "6089938", "url": "https://pubmed.ncbi.nlm.nih.gov/6089938/", "web_refs": [ "turn191search2", "turn191search47" ], "finding": "A roughly 1.35-kb deletion spanning the 5\u2032 HBB region into intron two was found in an HbS/beta-zero patient and a relative with beta-thalassemia trait.", "model_relevance": "Adult beta output can fail through a local cis lesion; distinguish beta loss from directly demonstrated gamma activation.", "limitation": "This lesion alone is not direct proof of LCR competition or of a specific gamma-repressor pathway.", "usa_context": "Medical College of Georgia and Veterans Administration Hemoglobin Research Laboratory, Augusta.", "huisman_author": true, "felice_author": true, "volume": "64", "issue": "4", "pages": "941-944", "doi": "10.1182/blood.v64.4.941.941", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6089938/", "https://europepmc.org/article/MED/6089938" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "schroeder_author": false, "joseph_borg_author": false, "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_6713099", "pmid": "6713099", "year": 1984, "title": "The rare alpha-thalassemia-1 of blacks is a zeta alpha-thalassemia-1 associated with deletion of all alpha- and zeta-globin genes", "authors": [ "Felice AE", "Cleek MP", "McKie K", "McKie V", "Huisman TH" ], "author_string": "Felice AE, Cleek MP, McKie K, McKie V, Huisman TH.", "journal": "Blood", "journal_abbreviation": "Blood", "volume": "63", "issue": "5", "pages": "1253-1257", "doi": "10.1182/blood.v63.5.1253.1253", "url": "https://pubmed.ncbi.nlm.nih.gov/6713099/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/6713099/", "https://europepmc.org/article/MED/6713099" ], "headline": "A deletion spans the alpha and zeta genes", "finding": "Restriction mapping identified an alpha-thalassemia deletion extending across the alpha- and zeta-globin gene complex.", "model_relevance": "Felice\u2019s work directly encompassed the embryonic alpha-like locus as well as adult globins.", "limitation": "A multigene deletion cannot identify one shared regulator of embryonic and adult expression.", "timeline_type": "milestone", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article", "Case Reports" ], "publication_type": "Case report", "publication_type_basis": "PubMed/Europe PMC case-report indexing; may include original molecular characterization.", "borg_author": false, "seminal": false }, { "id": "henthorn_felice_1985_deletion", "year": 1985, "title": "(A gamma delta beta)0-Thalassaemia in Blacks is due to a deletion of 34 kbp of DNA", "authors": [ "Henthorn PS", "Smithies O", "Nakatsuji T", "Felice AE", "Gardiner MB", "Reese AL", "Huisman TH" ], "journal": "British journal of haematology", "doi": "10.1111/j.1365-2141.1985.tb02999.x", "pmid": "2578804", "url": "https://pubmed.ncbi.nlm.nih.gov/2578804/", "web_refs": [ "turn198search2", "turn192search43" ], "finding": "Ten families previously assigned G\u03b3-delta-beta-zero thalassemia or HPFH labels shared a roughly 34-kb deletion removing part of A\u03b3 and all of pseudobeta, delta, and beta; the junction was cloned.", "model_relevance": "Track actual deleted elements, not phenotype labels alone; this rearrangement directly involves the intergenic and pseudogene-containing region.", "limitation": "Removing several elements together cannot identify which single deleted sequence is responsible for persistent gamma expression.", "usa_context": "University of Wisconsin\u2013Madison; Medical College of Georgia and Veterans Administration, Augusta.", "huisman_author": true, "felice_author": true, "borg_author": false, "schroeder_author": false, "volume": "59", "issue": "2", "pages": "343-356", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/2578804/", "https://europepmc.org/article/MED/2578804" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": true }, { "id": "gilman_1985_xmni", "year": 1985, "title": "DNA sequence variation associated with elevated fetal G gamma globin production", "authors": [ "Gilman JG", "Huisman TH" ], "journal": "Blood", "pmid": "2412616", "url": "https://pubmed.ncbi.nlm.nih.gov/2412616/", "web_refs": [ "turn194search0", "turn206search0" ], "finding": "The HBG2 \u2212158 T variant creating an XmnI site was associated with higher G\u03b3 contribution in sickle-cell and beta-thalassemia families.", "model_relevance": "Promoter/haplotype background can alter HBG2 contribution even when a family shares a major trans-acting mutation.", "limitation": "Association is not an isolated causal edit, and G\u03b3 percentage within HbF is not HbF percentage of total hemoglobin.", "huisman_author": true, "felice_author": false, "volume": "66", "issue": "4", "pages": "783-787", "doi": "10.1182/blood.v66.4.783.783", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/2412616/", "https://europepmc.org/article/MED/2412616" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "schroeder_author": false, "joseph_borg_author": false, "tags": [ "T. H. J. Huisman" ], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_3015767", "pmid": "3015767", "year": 1986, "title": "Different zeta globin gene deletions among black Americans", "authors": [ "Felice AE", "Cleek MP", "Marino EM", "McKie KM", "McKie VC", "Chang BK", "Huisman TH" ], "author_string": "Felice AE, Cleek MP, Marino EM, McKie KM, McKie VC, Chang BK, Huisman TH.", "journal": "Human genetics", "journal_abbreviation": "Hum Genet", "volume": "73", "issue": "3", "pages": "221-224", "doi": "10.1007/bf00401231", "url": "https://pubmed.ncbi.nlm.nih.gov/3015767/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/3015767/", "https://europepmc.org/article/MED/3015767" ], "headline": "Zeta-region deletions and adult alpha output", "finding": "The study characterized several zeta-region deletion configurations in Georgia and found carrier hematology consistent with preserved adult alpha-gene function.", "model_relevance": "Provides an early natural contrast between embryonic-region deletion and adult alpha output.", "limitation": "Carrier observations are not a complete assay of embryonic zeta regulation or homozygous consequences.", "timeline_type": "milestone", "tags": [ "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "lanclos_1987_hybrids", "year": 1987, "title": "Characterization of chromosomes with hybrid genes for Hb Lepore-Washington, Hb Lepore-Baltimore, Hb P-Nilotic, and Hb Kenya", "authors": [ "Lanclos KD", "Patterson J", "Efremov GD", "Wong SC", "Villegas A", "Ojwang PJ", "Wilson JB", "Kutlar F", "Huisman TH" ], "journal": "Human genetics", "doi": "10.1007/bf00284711", "pmid": "2442092", "url": "https://pubmed.ncbi.nlm.nih.gov/2442092/", "web_refs": [ "turn194search2" ], "finding": "Hybrid-gene chromosomes showed distinctive haplotypes, and HbF/G\u03b3 patterns in Hb Lepore-Washington were associated with the surrounding haplotype background.", "model_relevance": "Show rearrangement class and cis background as separate inputs; do not give all Hb Lepore carriers one fixed predicted HbF value.", "limitation": "Restriction-haplotype associations do not directly measure modern enhancer competition or three-dimensional contacts.", "huisman_author": true, "felice_author": false, "borg_author": false, "schroeder_author": false, "volume": "77", "issue": "1", "pages": "40-45", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/2442092/", "https://europepmc.org/article/MED/2442092" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [ "T. H. J. Huisman" ], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": false }, { "id": "history_pmid_2441597", "pmid": "2441597", "year": 1987, "title": "Effects of alpha-thalassemia-2 on the developmental changes of hematological values in children with sickle cell disease from Georgia", "authors": [ "Felice AE", "McKie KM", "Cleek MP", "Marino EM", "Kutlar A", "McKie VC" ], "author_string": "Felice AE, McKie KM, Cleek MP, Marino EM, Kutlar A, McKie VC.", "journal": "American journal of hematology", "journal_abbreviation": "Am J Hematol", "volume": "25", "issue": "4", "pages": "389-400", "doi": "10.1002/ajh.2830250405", "url": "https://pubmed.ncbi.nlm.nih.gov/2441597/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/2441597/", "https://europepmc.org/article/MED/2441597" ], "headline": "Development, alpha dosage and HbF interact", "finding": "Longitudinal data from children with sickle-cell disease connected alpha-gene number, age and HbF with evolving red-cell and anemia phenotypes.", "model_relevance": "A developmental model must account for protein composition and cell physiology as well as transcription.", "limitation": "The study was in sickle-cell disease; it does not identify the high-HbF modifier in the Maltese KLF1 family.", "timeline_type": "milestone", "tags": [ "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, U.S. Gov't, Non-P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_3690667", "pmid": "3690667", "year": 1987, "title": "Position-independent, high-level expression of the human beta-globin gene in transgenic mice", "authors": [ "Grosveld F", "van Assendelft GB", "Greaves DR", "Kollias G" ], "author_string": "Grosveld F, van Assendelft GB, Greaves DR, Kollias G.", "journal": "Cell", "journal_abbreviation": "Cell", "volume": "51", "issue": "6", "pages": "975-985", "doi": "10.1016/0092-8674(87)90584-8", "url": "https://pubmed.ncbi.nlm.nih.gov/3690667/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/3690667/", "https://europepmc.org/article/MED/3690667" ], "headline": "The locus control region principle", "finding": "A human beta-globin minilocus in transgenic mice achieved high erythroid expression largely independent of integration position and proportional to copy number.", "model_relevance": "Long-range regulatory DNA becomes essential to the locus model.", "limitation": "Transgene behavior does not specify every native human enhancer\u2013promoter interaction.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Grosveld F", "affiliation": "Laboratory of Gene Structure and Expression, National Institute for Medical Research, London, England." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "yang_1988_promoters", "year": 1988, "title": "Identification of base substitutions in the promoter regions of the A gamma- and G gamma-globin genes in A gamma- (or G gamma-) beta+-HPFH heterozygotes using the DNA-amplification-synthetic oligonucleotide procedure", "authors": [ "Yang KG", "Stoming TA", "Fei YJ", "Liang S", "Wong SC", "Masala B", "Huang RB", "Wei ZP", "Huisman TH" ], "journal": "Blood", "doi": "10.1182/blood.v71.5.1414.1414", "pmid": "2451950", "url": "https://pubmed.ncbi.nlm.nih.gov/2451950/", "web_refs": [ "turn202search0", "turn202search1" ], "finding": "Family screening identified HBG1 promoter variants at \u2212198, \u2212196, and \u2212117 and an HBG2 \u2212202 substitution among nondeletional HPFH families.", "model_relevance": "Local gamma-promoter variants provide a route to persistent HbF without requiring a KLF1 mutation or a large deletion.", "limitation": "Identifying variants in families does not establish the full modern transcription-factor binding mechanism at each base.", "usa_context": "Medical College of Georgia, Augusta.", "huisman_author": true, "felice_author": false, "borg_author": false, "schroeder_author": false, "volume": "71", "issue": "5", "pages": "1414-1417", "timeline_type": "milestone", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/2451950/", "https://europepmc.org/article/MED/2451950" ], "verified_at": "2026-10-23", "verification_note": "Existing atlas interpretation retained; bibliographic metadata rechecked against Europe PMC MED record.", "joseph_borg_author": false, "tags": [ "T. H. J. Huisman" ], "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "seminal": false }, { "id": "history_pmid_1709134", "pmid": "1709134", "year": 1991, "title": "The linkage of Hb Valletta [alpha 2 beta 287(f3)Thr----Pro] and Hb F-Malta-I [alpha 2G gamma 2117(G19)His----Arg] in the Maltese population", "authors": [ "Kutlar F", "Felice AE", "Grech JL", "Bannister WH", "Kutlar A", "Wilson JB", "Webber BB", "Hu HY", "Huisman TH" ], "author_string": "Kutlar F, Felice AE, Grech JL, Bannister WH, Kutlar A, Wilson JB, Webber BB, Hu HY, Huisman TH.", "journal": "Human genetics", "journal_abbreviation": "Hum Genet", "volume": "86", "issue": "6", "pages": "591-594", "doi": "10.1007/bf00201546", "url": "https://pubmed.ncbi.nlm.nih.gov/1709134/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/1709134/", "https://europepmc.org/article/MED/1709134" ], "headline": "A Maltese chromosome links fetal and adult markers", "finding": "Hb Valletta was identified and found tightly linked to Hb F-Malta-I in the sampled Maltese families and newborns.", "model_relevance": "The linked variants provide natural markers across the gamma\u2013beta interval.", "limitation": "Tight linkage in these samples does not establish an absolute absence of recombination or a causal switching element.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice", "T. H. J. Huisman" ], "felice_author": true, "huisman_author": true, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Kutlar F", "affiliation": "Department of Cell and Molecular Biology, Medical College of Georgia, Augusta 30912-2100." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, U.S. Gov't, P.H.S.", "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_7682653", "pmid": "7682653", "year": 1993, "title": "A novel, erythroid cell-specific murine transcription factor that binds to the CACCC element and is related to the Kr\u00fcppel family of nuclear proteins", "authors": [ "Miller IJ", "Bieker JJ" ], "author_string": "Miller IJ, Bieker JJ.", "journal": "Molecular and cellular biology", "journal_abbreviation": "Mol Cell Biol", "volume": "13", "issue": "5", "pages": "2776-2786", "doi": "10.1128/mcb.13.5.2776-2786.1993", "url": "https://pubmed.ncbi.nlm.nih.gov/7682653/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/7682653/", "https://europepmc.org/article/MED/7682653" ], "headline": "EKLF is identified", "finding": "Miller and Bieker isolated the erythroid factor EKLF, now KLF1, and characterized its CACCC-element binding.", "model_relevance": "Introduces a central adult beta-globin activator later linked to the Maltese family.", "limitation": "The discovery paper does not establish every downstream KLF1 target.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Miller IJ", "affiliation": "Department of Biochemistry, Mount Sinai School of Medicine, New York, New York 10029." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Comparative Study", "Research Support, U.S. Gov't, P.H.S.", "Research Support, Non-U.S. Gov't", "research-article", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_8518184", "pmid": "8518184", "year": 1993, "title": "The beta + IVS, I-NT no. 6 (T --> C) thalassaemia in heterozygotes with an associated Hb Valletta or Hb S heterozygosity in homozygotes from Malta", "authors": [ "Scerri CA", "Abela W", "Galdies R", "Pizzuto M", "Grech JL", "Felice AE" ], "author_string": "Scerri CA, Abela W, Galdies R, Pizzuto M, Grech JL, Felice AE.", "journal": "British journal of haematology", "journal_abbreviation": "Br J Haematol", "volume": "83", "issue": "4", "pages": "669-671", "doi": "10.1111/j.1365-2141.1993.tb04710.x", "url": "https://pubmed.ncbi.nlm.nih.gov/8518184/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/8518184/", "https://europepmc.org/article/MED/8518184" ], "headline": "Quantifying the Maltese IVS-I-6 allele", "finding": "Molecular screening established the prominence of HBB IVS-I-6 T>C in the sampled Maltese thalassemia families and used compound carriers to assess residual beta output.", "model_relevance": "Connects Maltese population genetics with quantitative adult globin expression.", "limitation": "The mutation frequency is from a selected sample, not a current population-wide estimate.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Scerri CA", "affiliation": "Department of Health, Valletta, Malta." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_7753194", "pmid": "7753194", "year": 1995, "title": "Defective haematopoiesis in fetal liver resulting from inactivation of the EKLF gene", "authors": [ "Nuez B", "Michalovich D", "Bygrave A", "Ploemacher R", "Grosveld F" ], "author_string": "Nuez B, Michalovich D, Bygrave A, Ploemacher R, Grosveld F.", "journal": "Nature", "journal_abbreviation": "Nature", "volume": "375", "issue": "6529", "pages": "316-318", "doi": "10.1038/375316a0", "url": "https://pubmed.ncbi.nlm.nih.gov/7753194/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/7753194/", "https://europepmc.org/article/MED/7753194" ], "headline": "Klf1 loss disrupts definitive erythropoiesis", "finding": "Targeted Eklf inactivation in mice disrupted fetal-liver erythropoiesis, establishing an essential developmental role.", "model_relevance": "Provides genetic evidence for the importance of KLF1 in adult-type erythroid development.", "limitation": "A complete mouse knockout is not equivalent to heterozygous human K288X.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Nuez B", "affiliation": "MGC Department of Cell Biology and Genetics, Erasmus University, Rotterdam, The Netherlands." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_7753195", "pmid": "7753195", "year": 1995, "title": "Lethal beta-thalassaemia in mice lacking the erythroid CACCC-transcription factor EKLF", "authors": [ "Perkins AC", "Sharpe AH", "Orkin SH" ], "author_string": "Perkins AC, Sharpe AH, Orkin SH.", "journal": "Nature", "journal_abbreviation": "Nature", "volume": "375", "issue": "6529", "pages": "318-322", "doi": "10.1038/375318a0", "url": "https://pubmed.ncbi.nlm.nih.gov/7753195/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/7753195/", "https://europepmc.org/article/MED/7753195" ], "headline": "Klf1 knockout causes beta-globin deficiency", "finding": "Perkins, Sharpe and Orkin found lethal fetal anemia with beta-globin deficiency after deleting Eklf in mice.", "model_relevance": "Links KLF1 requirement directly to adult beta output.", "limitation": "Native mice do not reproduce the human fetal gamma phase, and knockout dosage differs from the family.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Perkins AC", "affiliation": "Division of Hematology/oncology, Children's Hospital, Boston, Massachusetts, USA." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_9028826", "pmid": "9028826", "year": 1997, "title": "Hb Setif [alpha 94(G1)Asp-->Tyr] in Malta", "authors": [ "Borg I", "Valentino M", "Fiorini A", "Felice AE" ], "author_string": "Borg I, Valentino M, Fiorini A, Felice AE.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "21", "issue": "1", "pages": "91-96", "doi": "10.3109/03630269708997513", "url": "https://pubmed.ncbi.nlm.nih.gov/9028826/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/9028826/", "https://europepmc.org/article/MED/9028826" ], "headline": "Hb Setif in Malta", "finding": "A Maltese family report identified the alpha-chain variant Hb Setif and extended the local hemoglobin-variant catalogue.", "model_relevance": "Shows continuity of molecular characterization after the Augusta period.", "limitation": "The first author is Isabella Borg, not Joseph Borg.", "timeline_type": "supporting", "tags": [ "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Borg I", "affiliation": "Department of Pathology, University of Malta, Msida, Malta." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article", "Case Reports" ], "publication_type": "Case report", "publication_type_basis": "PubMed/Europe PMC case-report indexing; may include original molecular characterization.", "borg_author": false, "seminal": false }, { "id": "history_pmid_10335982", "pmid": "10335982", "year": 1999, "title": "Characterization and locus assignment of two alpha-globin variants present in the Maltese population: Hb St. Luke's [alpha95(G2)Pro-->Arg] and Hb Setif [alpha94(G1)Asp-->Tyr]", "authors": [ "Bezzina Wettinger S", "Galdies R", "Scerri C", "Felice AE" ], "author_string": "Bezzina Wettinger S, Galdies R, Scerri C, Felice AE.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "23", "issue": "2", "pages": "145-157", "doi": "10.3109/03630269908996159", "url": "https://pubmed.ncbi.nlm.nih.gov/10335982/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/10335982/", "https://europepmc.org/article/MED/10335982" ], "headline": "Assigning Maltese alpha variants to their genes", "finding": "Protein analysis and sequencing assigned Hb St. Luke\u2019s to an alpha1 gene and Hb Setif to an alpha2 gene.", "model_relevance": "Connects locus identity with measured output of natural alpha-chain markers.", "limitation": "Variant protein abundance is influenced by stability and assembly; it is not a direct transcript-count comparison.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Bezzina Wettinger S", "affiliation": "Department of Pathology, University of Malta, Msida." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_10490134", "pmid": "10490134", "year": 1999, "title": "Comparative in vivo expression of beta(+)-thalassemia alleles", "authors": [ "Marwan MM", "Scerri CA", "Zarroag SO", "Cao A", "Kyrri A", "Kalogirou E", "Kleanthous M", "Ioannou P", "Angastiniotis M", "Felice AE" ], "author_string": "Marwan MM, Scerri CA, Zarroag SO, Cao A, Kyrri A, Kalogirou E, Kleanthous M, Ioannou P, Angastiniotis M, Felice AE.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "23", "issue": "3", "pages": "221-229", "doi": "10.3109/03630269909005702", "url": "https://pubmed.ncbi.nlm.nih.gov/10490134/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/10490134/", "https://europepmc.org/article/MED/10490134" ], "headline": "Residual beta output becomes an allele-level measurement", "finding": "Compound heterozygotes carrying a stable variant enabled quantitative comparison of residual expression from several beta-plus-thalassemia alleles.", "model_relevance": "A model of globin balance needs quantitative residual adult output, not a binary beta on/off state.", "limitation": "Some alleles were represented by very small numbers of subjects.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Marwan MM", "affiliation": "Department of Pathology, University of Malta, Msida." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Comparative Study", "Research Support, Non-U.S. Gov't", "Multicenter Study", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_12504019", "pmid": "12504019", "year": 2002, "title": "Looping and interaction between hypersensitive sites in the active beta-globin locus", "authors": [ "Tolhuis B", "Palstra RJ", "Splinter E", "Grosveld F", "de Laat W" ], "author_string": "Tolhuis B, Palstra RJ, Splinter E, Grosveld F, de Laat W.", "journal": "Molecular cell", "journal_abbreviation": "Mol Cell", "volume": "10", "issue": "6", "pages": "1453-1465", "doi": "10.1016/s1097-2765(02)00781-5", "url": "https://pubmed.ncbi.nlm.nih.gov/12504019/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/12504019/", "https://europepmc.org/article/MED/12504019" ], "headline": "The locus is organized in three dimensions", "finding": "Chromosome-conformation analysis showed the active murine beta-globin genes near distant LCR hypersensitive sites.", "model_relevance": "A spatial interaction layer complements linear promoter and enhancer maps.", "limitation": "Population contact measurements do not establish a fixed loop in every cell.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Tolhuis B", "affiliation": "Department of Cell Biology and Genetics, Faculty of Medicine, Erasmus University, Rotterdam, P.O. Box 1738, 3000DR, Rotterdam, The Netherlands." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_12779278", "pmid": "12779278", "year": 2003, "title": "Hb St. Luke's [alpha95(G2)Pro --> Arg (alpha1)]", "authors": [ "Felice AE" ], "author_string": "Felice AE.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "27", "issue": "2", "pages": "137", "doi": "10.1081/hem-120021550", "url": "https://pubmed.ncbi.nlm.nih.gov/12779278/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/12779278/", "https://europepmc.org/article/MED/12779278" ], "headline": "Refining Hb St. Luke\u2019s nomenclature", "finding": "Felice published a short comment on the alpha1-associated Hb St. Luke\u2019s variant.", "model_relevance": "Maintains a traceable record of locus assignment and variant nomenclature.", "limitation": "This is a comment/letter, not a new mechanistic discovery.", "timeline_type": "context", "tags": [ "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Letter", "Comment" ], "publication_type": "Commentary", "publication_type_basis": "PubMed/Europe PMC types: Letter and Comment.", "borg_author": false, "seminal": false }, { "id": "history_pmid_14517543", "pmid": "14517543", "year": 2003, "title": "The beta-globin nuclear compartment in development and erythroid differentiation", "authors": [ "Palstra RJ", "Tolhuis B", "Splinter E", "Nijmeijer R", "Grosveld F", "de Laat W" ], "author_string": "Palstra RJ, Tolhuis B, Splinter E, Nijmeijer R, Grosveld F, de Laat W.", "journal": "Nature genetics", "journal_abbreviation": "Nat Genet", "volume": "35", "issue": "2", "pages": "190-194", "doi": "10.1038/ng1244", "url": "https://pubmed.ncbi.nlm.nih.gov/14517543/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/14517543/", "https://europepmc.org/article/MED/14517543" ], "headline": "Chromatin organization changes with development", "finding": "Palstra and colleagues examined the beta-globin active chromatin hub across development and erythroid differentiation.", "model_relevance": "Supports modeling regulatory contacts as stage-dependent.", "limitation": "Chromatin contacts alone do not establish all causal steps of switching.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Palstra RJ", "affiliation": "Department of Cell Biology and Genetics, ErasmusMC, PO Box 1738, 3000 DR Rotterdam, The Netherlands." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_15489291", "pmid": "15489291", "year": 2004, "title": "The active spatial organization of the beta-globin locus requires the transcription factor EKLF", "authors": [ "Drissen R", "Palstra RJ", "Gillemans N", "Splinter E", "Grosveld F", "Philipsen S", "de Laat W" ], "author_string": "Drissen R, Palstra RJ, Gillemans N, Splinter E, Grosveld F, Philipsen S, de Laat W.", "journal": "Genes & development", "journal_abbreviation": "Genes Dev", "volume": "18", "issue": "20", "pages": "2485-2490", "doi": "10.1101/gad.317004", "url": "https://pubmed.ncbi.nlm.nih.gov/15489291/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/15489291/", "https://europepmc.org/article/MED/15489291" ], "headline": "KLF1 helps organize the active locus", "finding": "Experimental analysis found that formation of the active beta-globin chromatin hub depended on EKLF.", "model_relevance": "Connects a transcription factor to long-range locus organization.", "limitation": "The mouse result does not measure chromatin contacts in individual Maltese carriers.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Drissen R", "affiliation": "Erasmus University Medical Center, Department of Cell Biology, The Netherlands." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Comparative Study", "Research Support, Non-U.S. Gov't", "research-article", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "pmid-17486512", "pmid": "17486512", "year": 2007, "title": "A review of cis-trans interplay between DNA sequences 5' to the (G)gamma- and beta-globin genes among Hb F-Malta-I heterozygotes/homozygotes and beta-thalassemia homozygotes/compound heterozygotes, and the effects of hydroxyurea on the Hb F/F-erythrocyte; the need for large multicenter trials", "authors": [ "Alex E Felice", "Joseph Borg", "Monica Pizzuto", "Wilhelmina Cassar", "Ruth Galdies", "Stephanie Bezzina Wettinger", "Svetlana Pulis", "Gary J Hunter", "Mary R Caruana", "Mario Farrugia", "Christian A Scerri" ], "author_string": "Felice AE, Borg J, Pizzuto M, Cassar W, Galdies R, Wettinger SB, Pulis S, Hunter GJ, Caruana MR, Farrugia M, Scerri CA.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "31", "issue": "2", "pages": "279-88", "doi": "10.1080/03630260701297261", "url": "https://doi.org/10.1080/03630260701297261", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/17486512/", "https://europepmc.org/article/MED/17486512" ], "headline": "Cis\u2013trans interplay and hydroxyurea response", "finding": "Connects Maltese haemoglobin variants, cis\u2013trans interactions and hydroxyurea response in the earlier Felice\u2013Borg research programme.", "model_relevance": "A direct pre-2009 bridge from Felice\u2019s programme into Joseph Borg\u2019s globin research.", "limitation": "Review and synthesis of existing literature; not an independent experimental test of the family model.", "timeline_type": "review", "tags": [ "Joseph Borg", "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": true, "verified_affiliations": [ { "author": "Felice AE", "affiliation": "Laboratory of Molecular Genetics, and Thalassemia and Molecular Genetics Clinic, Division of Pathology, St. Luke's Hospital, Department of Health, G'Mangia. Malta. alex.felice@um.edu.mt" } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article", "Research Support, Non-U.S. Gov't" ], "publication_type": "Review", "publication_type_basis": "Title and abstract identify a synthesis/review; classification does not rely solely on generic Journal Article indexing.", "borg_author": true, "seminal": false, "first_publication_year": 2007, "issue_date": "2007", "online_date": null, "author_line": "Alex E Felice; Joseph Borg; Monica Pizzuto; Wilhelmina Cassar; Ruth Galdies; Stephanie Bezzina Wettinger; Svetlana Pulis; Gary J Hunter; Mary R Caruana; Mario Farrugia; Christian A Scerri", "joseph_role": "Co-author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/17486512/" ], "topics": [ "Globins & erythropoiesis", "Pharmacogenomics & therapeutics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Connects Maltese haemoglobin variants, cis\u2013trans interactions and hydroxyurea response in the earlier Felice\u2013Borg research programme.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/17486512/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/17486512/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [], "status": "published", "checked": "2026-10-23", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin", "genetics" ] }, { "id": "history_pmid_17365007", "pmid": "17365007", "year": 2007, "title": "Developmental effect of the XmnI site on Ggamma-globin gene expression among newborn Hb F-Malta-I [Ggamma117(G19)His-->Arg, CAT-->CGT] heterozygotes and adult beta+ -Thalassemia homozygotes", "authors": [ "Pulis S", "Scerri CA", "Wismayer PS", "Galdies R", "Wettinger SB", "Felice AE" ], "author_string": "Pulis S, Scerri CA, Wismayer PS, Galdies R, Wettinger SB, Felice AE.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "31", "issue": "1", "pages": "71-82", "doi": "10.1080/03630260601057187", "url": "https://pubmed.ncbi.nlm.nih.gov/17365007/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/17365007/", "https://europepmc.org/article/MED/17365007" ], "headline": "XmnI effects depend on developmental context", "finding": "Maltese Hb F-Malta-I newborns and adults with beta-thalassemia showed different relationships between the XmnI-linked background and gamma output.", "model_relevance": "A cis marker\u2019s association can depend on developmental and hematopoietic context.", "limitation": "Association with a marker does not isolate the causal nucleotide or mechanism.", "timeline_type": "milestone", "tags": [ "Malta", "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Pulis S", "affiliation": "Laboratory of Molecular Genetics, Department of Physiology and Biochemistry, University of Malta, Msida, Malta. svetlanapulis@hotmail.com" } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": false }, { "id": "history_pmid_17592125", "pmid": "17592125", "year": 2007, "title": "Intergenic variants of HBS1L-MYB are responsible for a major quantitative trait locus on chromosome 6q23 influencing fetal hemoglobin levels in adults", "authors": [ "Thein SL", "Menzel S", "Peng X", "Best S", "Jiang J", "Close J", "Silver N", "Gerovasilli A", "Ping C", "Yamaguchi M", "Wahlberg K", "Ulug P", "Spector TD", "Garner C", "Matsuda F", "Farrall M", "Lathrop M" ], "author_string": "Thein SL, Menzel S, Peng X, Best S, Jiang J, Close J, Silver N, Gerovasilli A, Ping C, Yamaguchi M, Wahlberg K, Ulug P, Spector TD, Garner C, Matsuda F, Farrall M, Lathrop M.", "journal": "Proceedings of the National Academy of Sciences of the United States of America", "journal_abbreviation": "Proc Natl Acad Sci U S A", "volume": "104", "issue": "27", "pages": "11346-11351", "doi": "10.1073/pnas.0611393104", "url": "https://pubmed.ncbi.nlm.nih.gov/17592125/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/17592125/", "https://europepmc.org/article/MED/17592125" ], "headline": "A trans-associated modifier interval", "finding": "High-resolution human association mapping identified HBS1L\u2013MYB-region variants linked to F-cell variation.", "model_relevance": "A genetic modifier layer extends beyond the beta-globin locus.", "limitation": "The 2007 mapping study does not alone resolve the entire enhancer-to-MYB mechanism.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Thein SL", "affiliation": "King's College London School of Medicine, Division of Gene and Cell Based Therapy, King's College Hospital, London SE5 9PJ, United Kingdom. sl.thein@kcl.ac.uk" } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_19056937", "pmid": "19056937", "year": 2008, "title": "Human fetal hemoglobin expression is regulated by the developmental stage-specific repressor BCL11A", "authors": [ "Sankaran VG", "Menne TF", "Xu J", "Akie TE", "Lettre G", "Van Handel B", "Mikkola HK", "Hirschhorn JN", "Cantor AB", "Orkin SH" ], "author_string": "Sankaran VG, Menne TF, Xu J, Akie TE, Lettre G, Van Handel B, Mikkola HK, Hirschhorn JN, Cantor AB, Orkin SH.", "journal": "Science (New York, N.Y.)", "journal_abbreviation": "Science", "volume": "322", "issue": "5909", "pages": "1839-1842", "doi": "10.1126/science.1165409", "url": "https://pubmed.ncbi.nlm.nih.gov/19056937/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/19056937/", "https://europepmc.org/article/MED/19056937" ], "headline": "BCL11A is functionally linked to HbF repression", "finding": "BCL11A knockdown in primary adult erythroid cells induced fetal hemoglobin, linking a human genetic modifier to repression.", "model_relevance": "Supplies the repressor arm that the Maltese KLF1 discovery subsequently connected to KLF1.", "limitation": "A major repressor is not a complete explanation of all developmental and family variation.", "timeline_type": "milestone", "tags": [], "felice_author": false, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Sankaran VG", "affiliation": "Division of Hematology/Oncology, Children's Hospital Boston, Harvard Stem Cell Institute, Harvard Medical School, Boston, MA 02115, USA." } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article", "Research Support, N.I.H., Extramural" ], "publication_type": "Original study", "publication_type_basis": "Original experimental, observational, structural or genetic study described by the title/abstract or publisher page; journal format Letter alone is not treated as commentary.", "borg_author": false, "seminal": true }, { "id": "history_pmid_19657830", "pmid": "19657830", "year": 2009, "title": "An electronic infrastructure for research and treatment of the thalassemias and other hemoglobinopathies: the Euro-mediterranean ITHANET project", "authors": [ "Lederer CW", "Basak AN", "Aydinok Y", "Christou S", "El-Beshlawy A", "Eleftheriou A", "Fattoum S", "Felice AE", "Fibach E", "Galanello R", "Gambari R", "Gavrila L", "Giordano PC", "Grosveld F", "Hassapopoulou H", "Hladka E", "Kanavakis E", "Locatelli F", "Old J", "Patrinos GP", "Romeo G", "Taher A", "Traeger-Synodinos J", "Vassiliou P", "Villegas A", "Voskaridou E", "Wajcman H", "Zafeiropoulos A", "Kleanthous M" ], "author_string": "Lederer CW, Basak AN, Aydinok Y, Christou S, El-Beshlawy A, Eleftheriou A, Fattoum S, Felice AE, Fibach E, Galanello R, Gambari R, Gavrila L, Giordano PC, Grosveld F, Hassapopoulou H, Hladka E, Kanavakis E, Locatelli F, Old J, Patrinos GP, Romeo G, Taher A, Traeger-Synodinos J, Vassiliou P, Villegas A, Voskaridou E, Wajcman H, Zafeiropoulos A, Kleanthous M.", "journal": "Hemoglobin", "journal_abbreviation": "Hemoglobin", "volume": "33", "issue": "3", "pages": "163-176", "doi": "10.1080/03630260903089177", "url": "https://pubmed.ncbi.nlm.nih.gov/19657830/", "source_urls": [ "https://pubmed.ncbi.nlm.nih.gov/19657830/", "https://europepmc.org/article/MED/19657830" ], "headline": "Shared infrastructure for hemoglobinopathy research", "finding": "The ITHANET consortium described a shared portal for mutations, protocols and international research coordination, with Felice as a coauthor.", "model_relevance": "Connects local disease expertise to reusable international knowledge resources.", "limitation": "This is an infrastructure contribution, not a newly identified switching mechanism.", "timeline_type": "resource", "tags": [ "Alex E. Felice" ], "felice_author": true, "huisman_author": false, "schroeder_author": false, "joseph_borg_author": false, "verified_affiliations": [ { "author": "Lederer CW", "affiliation": "Molecular Genetics Thalassaemia Department, The Cyprus Institute of Neurology and Genetics, Nicosia, Cyprus. lederer@cing.ac.cy" } ], "verification_note": "Bibliographic metadata retrieved from Europe PMC MED/PubMed record; significance paraphrased from indexed abstract, publisher page or narrowly from title when no abstract is available.", "verified_at": "2026-10-23", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "publication_type": "Resource", "publication_type_basis": "Paper describes the ITHANET research infrastructure and portal.", "borg_author": false, "seminal": false }, { "id": "pmid-19631200", "title": "Genetic recombination as a major cause of mutagenesis in the human globin gene clusters", "year": 2009, "first_publication_year": 2009, "issue_date": "2009-12", "online_date": "2009-07-22", "journal": "Clinical Biochemistry", "journal_abbreviation": "Clin Biochem", "volume": "42", "issue": "18", "pages": "1839-50", "doi": "10.1016/j.clinbiochem.2009.07.014", "pmid": "19631200", "url": "https://doi.org/10.1016/j.clinbiochem.2009.07.014", "authors": [ "Joseph Borg", "Marianthi Georgitsi", "Vassiliki Aleporou-Marinou", "Panagoula Kollia", "George P Patrinos" ], "author_line": "Joseph Borg; Marianthi Georgitsi; Vassiliki Aleporou-Marinou; Panagoula Kollia; George P Patrinos", "joseph_role": "First-listed author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/19631200/" ], "publication_type": "Review", "indexed_publication_types": [ "Journal Article", "Research Support, Non-U.S. Gov't", "Review" ], "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reviews how recombination generates deletions, duplications and hybrid genes in the human globin clusters, connecting structural variation with haemoglobin disease.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/19631200/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/19631200/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Erasmus MC, Faculty of Medicine and Health Sciences, MGC-Department of Cell Biology and Genetics, Rotterdam, The Netherlands." ], "status": "published", "checked": "2026-10-23", "finding": "Reviews how recombination generates deletions, duplications and hybrid genes in the human globin clusters, connecting structural variation with haemoglobin disease.", "authorship_note": "Joseph Borg: First-listed author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Review and synthesis of existing literature; not an independent experimental test of the family model." }, { "id": "pmid-20676099", "year": 2010, "title": "Haploinsufficiency for the erythroid transcription factor KLF1 causes hereditary persistence of fetal hemoglobin", "authors": [ "Joseph Borg", "Petros Papadopoulos", "Marianthi Georgitsi", "Laura Guti\u00e9rrez", "Godfrey Grech", "Pavlos Fanis", "Marios Phylactides", "Annemieke J M H Verkerk", "Peter J van der Spek", "Christian A Scerri", "Wilhelmina Cassar", "Ruth Galdies", "Wilfred van Ijcken", "Zeliha Ozg\u00fcr", "Nynke Gillemans", "Jun Hou", "Marisa Bugeja", "Frank G Grosveld", "Marieke von Lindern", "Alex E Felice", "George P Patrinos", "Sjaak Philipsen" ], "journal": "Nature Genetics", "doi": "10.1038/ng.630", "pmid": "20676099", "url": "https://doi.org/10.1038/ng.630", "pmc_url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC2930131/", "web_refs": [ "turn206search1" ], "finding": "The Maltese family study identified KLF1 p.K288X haploinsufficiency and linked reduced KLF1 activity to reduced BCL11A expression and persistent fetal haemoglobin.", "model_relevance": "Establish the shared trans-acting KLF1 route and distinguish it from the cis routes identified in the older family studies.", "limitation": "This does not by itself identify every modifier responsible for high versus moderate HbF among carriers, and does not establish BGLT3 as that modifier.", "huisman_author": false, "felice_author": true, "inclusion_reason": "Continuity from the Augusta scientific programme to the Maltese KLF1 family.", "borg_author": true, "schroeder_author": false, "first_publication_year": 2010, "issue_date": "2010-09", "online_date": "2010-08-01", "journal_abbreviation": "Nat Genet", "volume": "42", "issue": "9", "pages": "801-5", "author_line": "Joseph Borg; Petros Papadopoulos; Marianthi Georgitsi; Laura Guti\u00e9rrez; Godfrey Grech; Pavlos Fanis; Marios Phylactides; Annemieke J M H Verkerk; Peter J van der Spek; Christian A Scerri; Wilhelmina Cassar; Ruth Galdies; Wilfred van Ijcken; Zeliha Ozg\u00fcr; Nynke Gillemans; Jun Hou; Marisa Bugeja; Frank G Grosveld; Marieke von Lindern; Alex E Felice; George P Patrinos; Sjaak Philipsen", "joseph_role": "Co-first author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/20676099/", "https://www.nature.com/articles/ng.630" ], "publication_type": "Research letter", "indexed_publication_types": [ "Case Reports", "Journal Article", "Research Support, N.I.H., Extramural", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis" ], "is_globin": true, "timeline_recommended": true, "featured": true, "significance": "The Maltese family study identified KLF1 p.K288X haploinsufficiency and linked reduced KLF1 activity to reduced BCL11A expression and persistent fetal haemoglobin.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/20676099/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/20676099/", "https://www.nature.com/articles/ng.630" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Laboratory of Molecular Genetics, Department of Physiology and Biochemistry, University of Malta, Msida, Malta." ], "status": "published", "checked": "2026-10-23", "authorship_note": "Joseph Borg: Co-first author.", "website_topics": [ "globin" ], "seminal": true }, { "id": "pmid-21531944", "title": "Erythroid phenotypes associated with KLF1 mutations", "year": 2011, "first_publication_year": 2011, "issue_date": "2011-05", "online_date": null, "journal": "Haematologica", "journal_abbreviation": "Haematologica", "volume": "96", "issue": "5", "pages": "635-8", "doi": "10.3324/haematol.2011.043265", "pmid": "21531944", "url": "https://doi.org/10.3324/haematol.2011.043265", "authors": [ "Joseph Borg", "George P Patrinos", "Alex E Felice", "Sjaak Philipsen" ], "author_line": "Joseph Borg; George P Patrinos; Alex E Felice; Sjaak Philipsen", "joseph_role": "First-listed author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/21531944/" ], "publication_type": "Editorial", "indexed_publication_types": [ "Comment", "Editorial", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Synthesizes the emerging spectrum of KLF1-associated erythroid phenotypes and the relationship between adult globin activation and fetal globin repression.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/21531944/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/21531944/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [], "status": "published", "checked": "2026-10-23", "finding": "Synthesizes the emerging spectrum of KLF1-associated erythroid phenotypes and the relationship between adult globin activation and fetal globin repression.", "authorship_note": "Joseph Borg: First-listed author.", "website_topics": [ "globin" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "pmid-21423179", "title": "Systematic documentation and analysis of human genetic variation in hemoglobinopathies using the microattribution approach", "year": 2011, "first_publication_year": 2011, "issue_date": "2011-03-20", "online_date": "2011-03-20", "journal": "Nature Genetics", "journal_abbreviation": "Nat Genet", "volume": "43", "issue": "4", "pages": "295-301", "doi": "10.1038/ng.785", "pmid": "21423179", "url": "https://doi.org/10.1038/ng.785", "authors": [ "Belinda Giardine", "Joseph Borg", "Douglas R Higgs", "Kenneth R Peterson", "Sjaak Philipsen", "Donna Maglott", "Belinda K Singleton", "David J Anstee", "A Nazli Basak", "Barnaby Clark", "Flavia C Costa", "Paula Faustino", "Halyna Fedosyuk", "Alex E Felice", "Alain Francina", "Renzo Galanello", "Monica V E Gallivan", "Marianthi Georgitsi", "Richard J Gibbons", "Piero C Giordano", "Cornelis L Harteveld", "James D Hoyer", "Martin Jarvis", "Philippe Joly", "Emmanuel Kanavakis", "Panagoula Kollia", "Stephan Menzel", "Webb Miller", "Kamran Moradkhani", "John Old", "Adamantia Papachatzopoulou", "Manoussos N Papadakis", "Petros Papadopoulos", "Sonja Pavlovic", "Lucia Perseu", "Milena Radmilovic", "Cathy Riemer", "Stefania Satta", "Iris Schrijver", "Maja Stojiljkovic", "Swee Lay Thein", "Jan Traeger-Synodinos", "Ray Tully", "Takahito Wada", "John S Waye", "Claudia Wiemann", "Branka Zukic", "David H K Chui", "Henri Wajcman", "Ross C Hardison", "George P Patrinos" ], "author_line": "Belinda Giardine; Joseph Borg; Douglas R Higgs; Kenneth R Peterson; Sjaak Philipsen; Donna Maglott; Belinda K Singleton; David J Anstee; A Nazli Basak; Barnaby Clark; Flavia C Costa; Paula Faustino; Halyna Fedosyuk; Alex E Felice; Alain Francina; Renzo Galanello; Monica V E Gallivan; Marianthi Georgitsi; Richard J Gibbons; Piero C Giordano; Cornelis L Harteveld; James D Hoyer; Martin Jarvis; Philippe Joly; Emmanuel Kanavakis; Panagoula Kollia; Stephan Menzel; Webb Miller; Kamran Moradkhani; John Old; Adamantia Papachatzopoulou; Manoussos N Papadakis; Petros Papadopoulos; Sonja Pavlovic; Lucia Perseu; Milena Radmilovic; Cathy Riemer; Stefania Satta; Iris Schrijver; Maja Stojiljkovic; Swee Lay Thein; Jan Traeger-Synodinos; Ray Tully; Takahito Wada; John S Waye; Claudia Wiemann; Branka Zukic; David H K Chui; Henri Wajcman; Ross C Hardison; George P Patrinos", "joseph_role": "Co-first author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/21423179/", "https://pmc.ncbi.nlm.nih.gov/articles/PMC3878152/", "https://www.nature.com/articles/ng.785" ], "publication_type": "Research article", "indexed_publication_types": [ "Journal Article", "Research Support, N.I.H., Extramural", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": true, "significance": "Established microattribution across interlinked haemoglobin-variant databases, encouraging systematic sharing and recognition of individual variant observations.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/21423179/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/21423179/", "https://pmc.ncbi.nlm.nih.gov/articles/PMC3878152/", "https://www.nature.com/articles/ng.785" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration. Publisher explicitly states Belinda Giardine and Joseph Borg contributed equally.", "joseph_affiliations": [], "status": "published", "checked": "2026-10-23", "finding": "Established microattribution across interlinked haemoglobin-variant databases, encouraging systematic sharing and recognition of individual variant observations.", "authorship_note": "Joseph Borg: Co-first author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": true, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "pmid-23057549", "title": "KLF10 gene expression is associated with high fetal hemoglobin levels and with response to hydroxyurea treatment in \u03b2-hemoglobinopathy patients", "year": 2012, "first_publication_year": 2012, "issue_date": "2012-10", "online_date": null, "journal": "Pharmacogenomics", "journal_abbreviation": "Pharmacogenomics", "volume": "13", "issue": "13", "pages": "1487-500", "doi": "10.2217/pgs.12.125", "pmid": "23057549", "url": "https://doi.org/10.2217/pgs.12.125", "authors": [ "Joseph Borg", "Marios Phylactides", "Marina Bartsakoulia", "Christina Tafrali", "Carsten Lederer", "Alexander E Felice", "Adamantia Papachatzopoulou", "Alexandra Kourakli", "Eleana F Stavrou", "Soteroula Christou", "Jun Hou", "Sophia Karkabouna", "Christina Lappa-Manakou", "Zeliha Ozgur", "Wilfred van Ijcken", "Marieke von Lindern", "Frank G Grosveld", "Marianthi Georgitsi", "Marina Kleanthous", "Sjaak Philipsen", "George P Patrinos" ], "author_line": "Joseph Borg; Marios Phylactides; Marina Bartsakoulia; Christina Tafrali; Carsten Lederer; Alexander E Felice; Adamantia Papachatzopoulou; Alexandra Kourakli; Eleana F Stavrou; Soteroula Christou; Jun Hou; Sophia Karkabouna; Christina Lappa-Manakou; Zeliha Ozgur; Wilfred van Ijcken; Marieke von Lindern; Frank G Grosveld; Marianthi Georgitsi; Marina Kleanthous; Sjaak Philipsen; George P Patrinos", "joseph_role": "First-listed author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/23057549/" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis", "Pharmacogenomics & therapeutics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Identified KLF10 as a candidate pharmacogenomic marker associated with HbF levels and hydroxyurea response in Maltese and Hellenic cohorts. Association does not establish a universal causal switch.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/23057549/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/23057549/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Erasmus University Medical Center, Department of Cell Biology, Rotterdam, The Netherlands." ], "status": "published", "checked": "2026-10-23", "finding": "Identified KLF10 as a candidate pharmacogenomic marker associated with HbF levels and hydroxyurea response in Maltese and Hellenic cohorts. Association does not establish a universal causal switch.", "authorship_note": "Joseph Borg: First-listed author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "pmid-23556445", "title": "Genomic variation in the MAP3K5 gene is associated with \u03b2-thalassemia disease severity and hydroxyurea treatment efficacy", "year": 2013, "first_publication_year": 2013, "issue_date": "2013-04", "online_date": null, "journal": "Pharmacogenomics", "journal_abbreviation": "Pharmacogenomics", "volume": "14", "issue": "5", "pages": "469-83", "doi": "10.2217/pgs.13.31", "pmid": "23556445", "url": "https://doi.org/10.2217/pgs.13.31", "authors": [ "Christina Tafrali", "Arsinoi Paizi", "Joseph Borg", "Milena Radmilovic", "Marina Bartsakoulia", "Emily Giannopoulou", "Olga Giannakopoulou", "Maja Stojiljkovic-Petrovic", "Branka Zukic", "Konstantinos Poulas", "Eleana F Stavrou", "Polyxeni Lambropoulou", "Alexandra Kourakli", "Alexander E Felice", "Adamantia Papachatzopoulou", "Sjaak Philipsen", "Sonja Pavlovic", "Marianthi Georgitsi", "George P Patrinos" ], "author_line": "Christina Tafrali; Arsinoi Paizi; Joseph Borg; Milena Radmilovic; Marina Bartsakoulia; Emily Giannopoulou; Olga Giannakopoulou; Maja Stojiljkovic-Petrovic; Branka Zukic; Konstantinos Poulas; Eleana F Stavrou; Polyxeni Lambropoulou; Alexandra Kourakli; Alexander E Felice; Adamantia Papachatzopoulou; Sjaak Philipsen; Sonja Pavlovic; Marianthi Georgitsi; George P Patrinos", "joseph_role": "Co-author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/23556445/" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis", "Pharmacogenomics & therapeutics", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Associated MAP3K5 variation and expression with beta-thalassaemia severity and hydroxyurea response, extending HbF research beyond KLF1 alone.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/23556445/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/23556445/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [], "status": "published", "checked": "2026-10-23", "finding": "Associated MAP3K5 variation and expression with beta-thalassaemia severity and hydroxyurea response, extending HbF research beyond KLF1 alone.", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "xjenza-2014-klfs", "title": "Control of globin gene expression by Kruppel-like Factors", "year": 2014, "first_publication_year": 2014, "journal": "Xjenza Online", "authors": [ "Laura Grech", "Joseph Borg" ], "author_line": "Laura Grech; Joseph Borg", "publication_type": "Review", "url": "https://www.um.edu.mt/library/oar/handle/123456789/2033", "source_url": "https://www.um.edu.mt/library/oar/handle/123456789/2033", "evidence_urls": [ "https://www.um.edu.mt/library/oar/handle/123456789/2033", "https://www.um.edu.mt/library/oar/bitstream/123456789/2033/1/Control%20of%20globin%20gene%20expression%20by%20Kruppel-like%20Factors.pdf" ], "doi": "10.7423/XJENZA.2014.1.10", "pmid": null, "topics": [ "Globins & erythropoiesis" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reviews KLF1, KLF2, KLF3, KLF8 and KLF10 across developmental globin control, connecting the Maltese KLF1 result with a broader regulatory network.", "joseph_role": "Last-listed and corresponding author", "verification": "Actual paper PDF confirms Laura Grech first, Joseph Borg second and corresponding. Repository page range conflicts with PDF; PDF pagination used.", "status": "published", "checked": "2026-10-23", "volume": "2", "issue": "1", "pages": "66\u201377", "finding": "Reviews KLF1, KLF2, KLF3, KLF8 and KLF10 across developmental globin control, connecting the Maltese KLF1 result with a broader regulatory network.", "authorship_note": "Joseph Borg: Last-listed and corresponding author.", "website_topics": [ "globin" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Review and synthesis of existing literature; not an independent experimental test of the family model." }, { "id": "pmid-25155936", "title": "Individualizing fetal hemoglobin augmenting therapy for \u03b2-type hemoglobinopathies patients", "year": 2014, "first_publication_year": 2014, "issue_date": "2014-07", "online_date": null, "journal": "Pharmacogenomics", "journal_abbreviation": "Pharmacogenomics", "volume": "15", "issue": "10", "pages": "1355-64", "doi": "10.2217/pgs.14.101", "pmid": "25155936", "url": "https://doi.org/10.2217/pgs.14.101", "authors": [ "Aikaterini Gravia", "Vasiliki Chondrou", "Argyro Sgourou", "Ioanna Papantoni", "Joseph Borg", "Theodora Katsila", "Adamantia Papachatzopoulou", "George P Patrinos" ], "author_line": "Aikaterini Gravia; Vasiliki Chondrou; Argyro Sgourou; Ioanna Papantoni; Joseph Borg; Theodora Katsila; Adamantia Papachatzopoulou; George P Patrinos", "joseph_role": "Co-author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/25155936/" ], "publication_type": "Review", "indexed_publication_types": [ "Journal Article", "Research Support, Non-U.S. Gov't", "Review" ], "topics": [ "Globins & erythropoiesis", "Pharmacogenomics & therapeutics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reviews how genetic differences may guide individualisation of HbF-augmenting treatment in beta-type haemoglobinopathies.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/25155936/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/25155936/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [], "status": "published", "checked": "2026-10-23", "finding": "Reviews how genetic differences may guide individualisation of HbF-augmenting treatment in beta-type haemoglobinopathies.", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Review and synthesis of existing literature; not an independent experimental test of the family model." }, { "id": "lcr_loop", "from": "LCR contact promoted by tethered LDB1", "to": "HBG transcription", "effect": "activation", "level": "causal engineered looping", "finding": "Forced LCR-gamma contact activates gamma and reduces beta expression in primary adult human erythroblasts; related embryonic-gene experiments were performed in mice.", "limit": "An artificial looping perturbation establishes capacity, not a measured contact trajectory for each Maltese donor.", "source": "Deng et al. Cell 2014", "doi": "10.1016/j.cell.2014.05.050", "url": "https://pubmed.ncbi.nlm.nih.gov/25126789/", "ref": "turn204view2", "input_section": "edges", "title": "Reactivation of developmentally silenced globin genes by forced chromatin looping", "authors": [ "Deng W", "Rupon JW", "Krivega I", "Breda L", "Motta I", "Jahn KS", "Reik A", "Gregory PD", "Rivella S", "Dean A", "Blobel GA" ], "authors_full": [ "Wulan Deng", "Jeremy W Rupon", "Ivan Krivega", "Laura Breda", "Irene Motta", "Kristen S Jahn", "Andreas Reik", "Philip D Gregory", "Stefano Rivella", "Ann Dean", "Gerd A Blobel" ], "year": 2014, "journal": "Cell", "journal_abbreviation": "Cell", "volume": "158", "issue": "4", "pages": "849-860", "pmid": "25126789", "pmcid": "PMC4134511", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "research-article", "Journal Article", "Research Support, N.I.H., Extramural" ], "epmc_first_publication_date": "2014-08-01", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/25126789/", "https://europepmc.org/article/MED/25126789", "https://api.crossref.org/works/10.1016/j.cell.2014.05.050" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2014, 8 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Wulan Deng; Jeremy W Rupon; Ivan Krivega; Laura Breda; Irene Motta; Kristen S Jahn; Andreas Reik; Philip D Gregory; Stefano Rivella; Ann Dean; Gerd A Blobel", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "An artificial looping perturbation establishes capacity, not a measured contact trajectory for each Maltese donor.", "mechanistic": true, "seminal": true }, { "id": "pmid-25232502", "title": "Transfusion-independent \u03b2(0)-thalassemia after bone marrow transplantation failure: proposed involvement of high parental HbF and an epigenetic mechanism", "year": 2014, "first_publication_year": 2014, "issue_date": "2014", "online_date": "2014-09-05", "journal": "American Journal of Blood Research", "journal_abbreviation": "Am J Blood Res", "volume": "4", "issue": "1", "pages": "27-32", "doi": null, "pmid": "25232502", "url": "https://pubmed.ncbi.nlm.nih.gov/25232502/", "authors": [ "Katia Paciaroni", "Guido Lucarelli", "Fabrizio Martelli", "Anna Rita Migliaccio", "Marieke von Lindern", "Joseph Borg", "Nynke Gillemans", "Thamar B van Dijk", "Sjaak Philipsen" ], "author_line": "Katia Paciaroni; Guido Lucarelli; Fabrizio Martelli; Anna Rita Migliaccio; Marieke von Lindern; Joseph Borg; Nynke Gillemans; Thamar B van Dijk; Sjaak Philipsen", "joseph_role": "Co-author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/25232502/" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article" ], "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Described transfusion independence after failed marrow transplantation and proposed roles for inherited high HbF and epigenetic effects; a clinical observation, not proof of a general mechanism.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/25232502/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/25232502/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Department of Applied Biomedical Science, Faculty of Health Sciences, University of Malta Msida, MSD2090, Malta ; Department of Cell Biology, Erasmus MC P.O. box 2040, 3000 CA Rotterdam, The Netherlands." ], "status": "published", "checked": "2026-10-23", "finding": "Described transfusion independence after failed marrow transplantation and proposed roles for inherited high HbF and epigenetic effects; a clinical observation, not proof of a general mechanism.", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "pmid-24137000", "title": "Updates of the HbVar database of human hemoglobin variants and thalassemia mutations", "year": 2014, "first_publication_year": 2013, "issue_date": "2014-01", "online_date": "2013-10-16", "journal": "Nucleic Acids Research", "journal_abbreviation": "Nucleic Acids Res", "volume": "42", "issue": "Database issue", "pages": "D1063-9", "doi": "10.1093/nar/gkt911", "pmid": "24137000", "url": "https://doi.org/10.1093/nar/gkt911", "authors": [ "Belinda Giardine", "Joseph Borg", "Emmanouil Viennas", "Cristiana Pavlidis", "Kamran Moradkhani", "Philippe Joly", "Marina Bartsakoulia", "Cathy Riemer", "Webb Miller", "Giannis Tzimas", "Henri Wajcman", "Ross C Hardison", "George P Patrinos" ], "author_line": "Belinda Giardine; Joseph Borg; Emmanouil Viennas; Cristiana Pavlidis; Kamran Moradkhani; Philippe Joly; Marina Bartsakoulia; Cathy Riemer; Webb Miller; Giannis Tzimas; Henri Wajcman; Ross C Hardison; George P Patrinos", "joseph_role": "Co-author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/24137000/" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article", "Research Support, N.I.H., Extramural", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Expanded HbVar as a curated resource for haemoglobin variants and thalassaemia mutations, supporting interpretation of globin genotype\u2013phenotype relationships.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/24137000/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/24137000/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [], "status": "published", "checked": "2026-10-23", "finding": "Expanded HbVar as a curated resource for haemoglobin variants and thalassaemia mutations, supporting interpretation of globin genotype\u2013phenotype relationships.", "authorship_note": "Joseph Borg: Co-author.", "date_note": "First published online 2013-10-16; journal issue 2014-01. Listed once under the issue year.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "pmid-25480500", "title": "A genetic score for the prediction of beta-thalassemia severity", "year": 2015, "first_publication_year": 2014, "issue_date": "2015-04", "online_date": "2014-12-05", "journal": "Haematologica", "journal_abbreviation": "Haematologica", "volume": "100", "issue": "4", "pages": "452-7", "doi": "10.3324/haematol.2014.113886", "pmid": "25480500", "url": "https://doi.org/10.3324/haematol.2014.113886", "authors": [ "Fabrice Danjou", "Marcella Francavilla", "Franco Anni", "Stefania Satta", "Franca-Rosa Demartis", "Lucia Perseu", "Matteo Manca", "Maria Carla Sollaino", "Laura Manunza", "Elisabetta Mereu", "Giuseppe Marceddu", "Serge Pissard", "Philippe Joly", "Isabelle Thuret", "Raffaella Origa", "Joseph Borg", "Gian Luca Forni", "Antonio Piga", "Maria Eliana Lai", "Catherine Badens", "Paolo Moi", "Renzo Galanello" ], "author_line": "Fabrice Danjou; Marcella Francavilla; Franco Anni; Stefania Satta; Franca-Rosa Demartis; Lucia Perseu; Matteo Manca; Maria Carla Sollaino; Laura Manunza; Elisabetta Mereu; Giuseppe Marceddu; Serge Pissard; Philippe Joly; Isabelle Thuret; Raffaella Origa; Joseph Borg; Gian Luca Forni; Antonio Piga; Maria Eliana Lai; Catherine Badens; Paolo Moi; Renzo Galanello", "joseph_role": "Co-author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/25480500/" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article", "Multicenter Study" ], "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Combined globin and HbF-modifier variants into a validated genetic severity score for beta-thalassaemia, connecting molecular variation with transfusion-related clinical outcomes.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/25480500/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/25480500/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Department of Applied Biomedical Science, Faculty of Health Sciences and Laboratory of Molecular Genetics, Department of Physiology and Biochemistry, University of Malta, Msida, Malta." ], "status": "published", "checked": "2026-10-23", "finding": "Combined globin and HbF-modifier variants into a validated genetic severity score for beta-thalassaemia, connecting molecular variation with transfusion-related clinical outcomes.", "authorship_note": "Joseph Borg: Co-author.", "date_note": "First published online 2014-12-05; journal issue 2015-04. Listed once under the issue year.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "source": "Masuda et al. Science 2016", "doi": "10.1126/science.aad3312", "url": "https://pubmed.ncbi.nlm.nih.gov/26816381/", "finding": "LRF and BCL11A independently repress fetal hemoglobin; avoids treating LRF as merely a downstream BCL11A component.", "ref": "turn195search2", "input_section": "optional_sources", "title": "Transcription factors LRF and BCL11A independently repress expression of fetal hemoglobin", "authors": [ "Masuda T", "Wang X", "Maeda M", "Canver MC", "Sher F", "Funnell AP", "Fisher C", "Suciu M", "Martyn GE", "Norton LJ", "Zhu C", "Kurita R", "Nakamura Y", "Xu J", "Higgs DR", "Crossley M", "Bauer DE", "Orkin SH", "Kharchenko PV", "Maeda T" ], "authors_full": [ "Takeshi Masuda", "Xin Wang", "Manami Maeda", "Matthew C Canver", "Falak Sher", "Alister P W Funnell", "Chris Fisher", "Maria Suciu", "Gabriella E Martyn", "Laura J Norton", "Catherine Zhu", "Ryo Kurita", "Yukio Nakamura", "Jian Xu", "Douglas R Higgs", "Merlin Crossley", "Daniel E Bauer", "Stuart H Orkin", "Peter V Kharchenko", "Takahiro Maeda" ], "year": 2016, "journal": "Science", "journal_abbreviation": "Science", "volume": "351", "issue": "6270", "pages": "285-289", "pmid": "26816381", "pmcid": "PMC4778394", "indexed_publication_types": [ "research-article", "Journal Article" ], "epmc_first_publication_date": "2016-01-01", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/26816381/", "https://europepmc.org/article/MED/26816381", "https://api.crossref.org/works/10.1126/science.aad3312" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2016, 1, 15 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Takeshi Masuda; Xin Wang; Manami Maeda; Matthew C Canver; Falak Sher; Alister P W Funnell; Chris Fisher; Maria Suciu; Gabriella E Martyn; Laura J Norton; Catherine Zhu; Ryo Kurita; Yukio Nakamura; Jian Xu; Douglas R Higgs; Merlin Crossley; Daniel E Bauer; Stuart H Orkin; Peter V Kharchenko; Takahiro Maeda", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "id": "modern_optional_26816381", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Scope is limited to the reported experimental systems.", "mechanistic": true, "seminal": true }, { "id": "hbbp1_dna", "from": "HBBP1 DNA region", "to": "Adult repressive locus architecture", "effect": "supports repression", "level": "developmental contact comparison and deletion", "finding": "Removing the HBBP1 region shifts adult erythroid chromatin contacts toward a fetal-like configuration, increases LCR-gamma contact and raises gamma transcription.", "limit": "DNA deletion and RNA depletion are different perturbations; this evidence cannot assign a repressive sign to HBBP1 RNA abundance.", "source": "Huang et al. Genes & Development 2017", "doi": "10.1101/gad.303461.117", "url": "https://pubmed.ncbi.nlm.nih.gov/28916711/", "ref": "turn193search4", "input_section": "edges", "title": "Comparative analysis of three-dimensional chromosomal architecture identifies a novel fetal hemoglobin regulatory element", "authors": [ "Huang P", "Keller CA", "Giardine B", "Grevet JD", "Davies JOJ", "Hughes JR", "Kurita R", "Nakamura Y", "Hardison RC", "Blobel GA" ], "authors_full": [ "Peng Huang", "Cheryl A Keller", "Belinda Giardine", "Jeremy D Grevet", "James O J Davies", "Jim R Hughes", "Ryo Kurita", "Yukio Nakamura", "Ross C Hardison", "Gerd A Blobel" ], "year": 2017, "journal": "Genes & Development", "journal_abbreviation": "Genes Dev", "volume": "31", "issue": "16", "pages": "1704-1713", "pmid": "28916711", "pmcid": "PMC5647940", "indexed_publication_types": [ "Comparative Study", "research-article", "Journal Article" ], "epmc_first_publication_date": "2017-08-01", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/28916711/", "https://europepmc.org/article/MED/28916711", "https://api.crossref.org/works/10.1101/gad.303461.117" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2017, 8, 15 ], "crossref_online_date_parts": [ 2017, 9, 15 ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Peng Huang; Cheryl A Keller; Belinda Giardine; Jeremy D Grevet; James O J Davies; Jim R Hughes; Ryo Kurita; Yukio Nakamura; Ross C Hardison; Gerd A Blobel", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "date_note": "Crossref issue date 15 August 2017 and online date 15 September 2017 differ; retain year 2017 without claiming issue date as first publication.", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "DNA deletion and RNA depletion are different perturbations; this evidence cannot assign a repressive sign to HBBP1 RNA abundance.", "mechanistic": true, "seminal": false }, { "id": "klf1_lrf", "from": "KLF1", "to": "ZBTB7A/LRF", "effect": "activation", "level": "direct binding and perturbation", "finding": "KLF1 directly activates ZBTB7A through its proximal promoter in erythroid cells.", "limit": "LRF RNA abundance is not equivalent to its protein concentration, DNA occupancy or repressor-complex activity.", "source": "Norton et al. Blood Advances 2017", "doi": "10.1182/bloodadvances.2016002303", "url": "https://pubmed.ncbi.nlm.nih.gov/29296711/", "ref": "turn189search0", "input_section": "edges", "title": "KLF1 directly activates expression of the novel fetal globin repressor ZBTB7A/LRF in erythroid cells", "authors": [ "Norton LJ", "Funnell APW", "Burdach J", "Wienert B", "Kurita R", "Nakamura Y", "Philipsen S", "Pearson RCM", "Quinlan KGR", "Crossley M" ], "authors_full": [ "Laura J Norton", "Alister P W Funnell", "Jon Burdach", "Beeke Wienert", "Ryo Kurita", "Yukio Nakamura", "Sjaak Philipsen", "Richard C M Pearson", "Kate G R Quinlan", "Merlin Crossley" ], "year": 2017, "journal": "Blood Advances", "journal_abbreviation": "Blood Adv", "volume": "1", "issue": "11", "pages": "685-692", "pmid": "29296711", "pmcid": "PMC5727813", "indexed_publication_types": [ "research-article", "Journal Article" ], "epmc_first_publication_date": "2017-04-25", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/29296711/", "https://europepmc.org/article/MED/29296711", "https://api.crossref.org/works/10.1182/bloodadvances.2016002303" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2017, 4, 25 ], "crossref_online_date_parts": [ 2017, 4, 25 ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Laura J Norton; Alister P W Funnell; Jon Burdach; Beeke Wienert; Ryo Kurita; Yukio Nakamura; Sjaak Philipsen; Richard C M Pearson; Kate G R Quinlan; Merlin Crossley", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "LRF RNA abundance is not equivalent to its protein concentration, DNA occupancy or repressor-complex activity.", "mechanistic": true, "seminal": false }, { "id": "bcl_promoter", "from": "BCL11A", "to": "HBG promoters", "effect": "repression", "level": "direct binding, protein-domain and promoter-editing evidence", "finding": "BCL11A binds the distal TGACCA element near -115 and directly represses gamma promoters; perturbing the site relieves repression.", "limit": "Motif presence or ATAC accessibility alone does not demonstrate BCL11A occupancy in a family sample.", "source": "Liu et al. Cell 2018", "doi": "10.1016/j.cell.2018.03.016", "url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC5889339/", "ref": "turn193search2", "input_section": "edges", "title": "Direct Promoter Repression by BCL11A Controls the Fetal to Adult Hemoglobin Switch", "authors": [ "Liu N", "Hargreaves VV", "Zhu Q", "Kurland JV", "Hong J", "Kim W", "Sher F", "Macias-Trevino C", "Rogers JM", "Kurita R", "Nakamura Y", "Yuan GC", "Bauer DE", "Xu J", "Bulyk ML", "Orkin SH" ], "authors_full": [ "Nan Liu", "Victoria V Hargreaves", "Qian Zhu", "Jesse V Kurland", "Jiyoung Hong", "Woojin Kim", "Falak Sher", "Claudio Macias-Trevino", "Julia M Rogers", "Ryo Kurita", "Yukio Nakamura", "Guo-Cheng Yuan", "Daniel E Bauer", "Jian Xu", "Martha L Bulyk", "Stuart H Orkin" ], "year": 2018, "journal": "Cell", "journal_abbreviation": "Cell", "volume": "173", "issue": "2", "pages": "430-442.e17", "pmid": "29606353", "pmcid": "PMC5889339", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "research-article", "Journal Article", "Research Support, N.I.H., Extramural" ], "epmc_first_publication_date": "2018-03-29", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/29606353/", "https://europepmc.org/article/MED/29606353", "https://api.crossref.org/works/10.1016/j.cell.2018.03.016" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2018, 4 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Nan Liu; Victoria V Hargreaves; Qian Zhu; Jesse V Kurland; Jiyoung Hong; Woojin Kim; Falak Sher; Claudio Macias-Trevino; Julia M Rogers; Ryo Kurita; Yukio Nakamura; Guo-Cheng Yuan; Daniel E Bauer; Jian Xu; Martha L Bulyk; Stuart H Orkin", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Motif presence or ATAC accessibility alone does not demonstrate BCL11A occupancy in a family sample.", "mechanistic": true, "seminal": true }, { "id": "bglt3", "from": "BGLT3 locus transcription and RNA", "to": "HBG transcription", "effect": "activation", "level": "deletion, RNA depletion, overexpression and looping assays", "finding": "Both transcription through BGLT3 and its RNA support gamma expression. Locus transcription supports local looping; the RNA interacts with Mediator. Overexpression increased gamma in K562.", "limit": "RNA alone did not rescue deletion or transcription blockade. Epsilon was not reduced by BGLT3 deletion. Do not label BGLT3 a universal embryonic-globin switch.", "source": "Ivaldi et al. Blood 2018", "doi": "10.1182/blood-2018-07-862003", "url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC6213316/", "ref": "turn183search0", "input_section": "edges", "title": "Fetal \u03b3-globin genes are regulated by the BGLT3 long noncoding RNA locus", "authors": [ "Ivaldi MS", "Diaz LF", "Chakalova L", "Lee J", "Krivega I", "Dean A" ], "authors_full": [ "Maria Soledad Ivaldi", "Luis Francisco Diaz", "Lyubomira Chakalova", "Jongjoo Lee", "Ivan Krivega", "Ann Dean" ], "year": 2018, "journal": "Blood", "journal_abbreviation": "Blood", "volume": "132", "issue": "18", "pages": "1963-1973", "pmid": "30150205", "pmcid": "PMC6213316", "indexed_publication_types": [ "Research Support, N.I.H., Intramural", "research-article", "Journal Article" ], "epmc_first_publication_date": "2018-08-27", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/30150205/", "https://europepmc.org/article/MED/30150205", "https://api.crossref.org/works/10.1182/blood-2018-07-862003" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2018, 11, 1 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Maria Soledad Ivaldi; Luis Francisco Diaz; Lyubomira Chakalova; Jongjoo Lee; Ivan Krivega; Ann Dean", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "RNA alone did not rescue deletion or transcription blockade. Epsilon was not reduced by BGLT3 deletion. Do not label BGLT3 a universal embryonic-globin switch.", "mechanistic": true, "seminal": false }, { "id": "lrf_promoter", "from": "ZBTB7A/LRF", "to": "HBG promoters", "effect": "repression", "level": "direct binding and natural-variant recreation", "finding": "LRF binds near -200; naturally occurring HPFH promoter mutations can prevent its binding and increase gamma expression. BCL11A-associated HPFH mutations cluster near -115.", "limit": "Not every promoter HPFH allele works by removing a repressor; some create activator sites.", "source": "Martyn et al. Nature Genetics 2018", "doi": "10.1038/s41588-018-0085-0", "url": "https://pubmed.ncbi.nlm.nih.gov/29610478/", "ref": "turn207search2", "input_section": "edges", "title": "Natural regulatory mutations elevate the fetal globin gene via disruption of BCL11A or ZBTB7A binding", "authors": [ "Martyn GE", "Wienert B", "Yang L", "Shah M", "Norton LJ", "Burdach J", "Kurita R", "Nakamura Y", "Pearson RCM", "Funnell APW", "Quinlan KGR", "Crossley M" ], "authors_full": [ "Gabriella E Martyn", "Beeke Wienert", "Lu Yang", "Manan Shah", "Laura J Norton", "Jon Burdach", "Ryo Kurita", "Yukio Nakamura", "Richard C M Pearson", "Alister P W Funnell", "Kate G R Quinlan", "Merlin Crossley" ], "year": 2018, "journal": "Nature Genetics", "journal_abbreviation": "Nat Genet", "volume": "50", "issue": "4", "pages": "498-503", "pmid": "29610478", "pmcid": null, "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "epmc_first_publication_date": "2018-04-02", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/29610478/", "https://europepmc.org/article/MED/29610478", "https://api.crossref.org/works/10.1038/s41588-018-0085-0" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2018, 4 ], "crossref_online_date_parts": [ 2018, 4, 2 ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Gabriella E Martyn; Beeke Wienert; Lu Yang; Manan Shah; Laura J Norton; Jon Burdach; Ryo Kurita; Yukio Nakamura; Richard C M Pearson; Alister P W Funnell; Kate G R Quinlan; Merlin Crossley", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Not every promoter HPFH allele works by removing a repressor; some create activator sites.", "mechanistic": true, "seminal": true }, { "id": "mmsc-2018-globin-rearrangements", "title": "Novel beta globin gene cluster rearrangements and deletions in the Maltese islands", "year": 2018, "first_publication_year": 2018, "journal": "10th Malta Medical School Conference: conference abstract book", "authors": [ "Alexander Camilleri", "Ruth Galdies", "Wilhelmina Cassar", "Laura Grech", "Joseph J. Borg", "Jeremy Cutajar", "Christian A. Scerri", "Alex E. Felice" ], "author_line": "Alexander Camilleri; Ruth Galdies; Wilhelmina Cassar; Laura Grech; Joseph J. Borg; Jeremy Cutajar; Christian A. Scerri; Alex E. Felice", "publication_type": "Conference abstract", "url": "https://www.um.edu.mt/library/oar/handle/123456789/99427", "source_url": "https://www.um.edu.mt/library/oar/handle/123456789/99427", "evidence_urls": [ "https://www.um.edu.mt/library/oar/handle/123456789/99427" ], "doi": null, "pmid": null, "topics": [ "Globins & erythropoiesis" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reports Maltese beta-globin-cluster rearrangements and deletions, providing a structural-variation strand alongside the transcription-factor studies.", "joseph_role": "Co-author", "verification": "University repository citation includes Jeremy Cutajar, while its separate author-index field omits him. Citation author order retained; conference abstract explicitly labelled.", "status": "published", "checked": "2026-10-23", "finding": "Reports Maltese beta-globin-cluster rearrangements and deletions, providing a structural-variation strand alongside the transcription-factor studies.", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Conference abstract: preliminary reported findings, with less methodological detail than a full research article. It is not treated as an independently replicated mechanism." }, { "source": "Martyn et al. Blood 2019", "doi": "10.1182/blood-2018-07-863951", "url": "https://pubmed.ncbi.nlm.nih.gov/30617196/", "finding": "The -113A>G HPFH allele creates an activating GATA1 site without disrupting BCL11A binding; natural HPFH can add activation as well as remove repression.", "ref": "turn207search0", "input_section": "optional_sources", "title": "A natural regulatory mutation in the proximal promoter elevates fetal globin expression by creating a de novo GATA1 site", "authors": [ "Martyn GE", "Wienert B", "Kurita R", "Nakamura Y", "Quinlan KGR", "Crossley M" ], "authors_full": [ "Gabriella E Martyn", "Beeke Wienert", "Ryo Kurita", "Yukio Nakamura", "Kate G R Quinlan", "Merlin Crossley" ], "year": 2019, "journal": "Blood", "journal_abbreviation": "Blood", "volume": "133", "issue": "8", "pages": "852-856", "pmid": "30617196", "pmcid": null, "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "epmc_first_publication_date": "2019-01-07", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/30617196/", "https://europepmc.org/article/MED/30617196", "https://api.crossref.org/works/10.1182/blood-2018-07-863951" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2019, 2, 21 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Gabriella E Martyn; Beeke Wienert; Ryo Kurita; Yukio Nakamura; Kate G R Quinlan; Merlin Crossley", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "id": "modern_optional_30617196", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Scope is limited to the reported experimental systems.", "mechanistic": true, "seminal": false }, { "id": "synapse-2019-thalassaemia", "title": "Diagnosis of beta thalassaemia trait", "year": 2019, "first_publication_year": 2019, "journal": "The Synapse", "authors": [ "Joseph Borg", "Laura Grech" ], "author_line": "Joseph Borg; Laura Grech", "publication_type": "Clinical educational article", "url": "https://www.um.edu.mt/library/oar/handle/123456789/70964", "source_url": "https://www.um.edu.mt/library/oar/handle/123456789/70964", "evidence_urls": [ "https://www.um.edu.mt/library/oar/handle/123456789/70964" ], "doi": null, "pmid": null, "topics": [ "Globins & erythropoiesis" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Explains the laboratory diagnosis of beta-thalassaemia trait and its relevance to haemoglobin assessment in clinical practice.", "joseph_role": "First-listed author", "verification": "Verified against publisher or University of Malta repository record.", "status": "published", "checked": "2026-10-23", "volume": "18", "issue": "5", "pages": "11\u201313", "finding": "Explains the laboratory diagnosis of beta-thalassaemia trait and its relevance to haemoglobin assessment in clinical practice.", "authorship_note": "Joseph Borg: First-listed author.", "website_topics": [ "globin" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Clinical educational contribution; not a mechanistic discovery about globin switching." }, { "id": "doi-blood-2020-143116", "title": "A Functional Single Nucleotide Polymorphism in the A\u03b3 Globin Gene Promoter Affects Globin Chain Synthesis", "year": 2020, "first_publication_year": 2020, "journal": "Blood", "authors": [ "Elena Spiteri", "Laura Grech", "Alexander E. Felice", "Joseph Borg" ], "author_line": "Elena Spiteri; Laura Grech; Alexander E. Felice; Joseph Borg", "publication_type": "Conference abstract", "url": "https://doi.org/10.1182/blood-2020-143116", "source_url": "https://www.um.edu.mt/library/oar/handle/123456789/99067", "evidence_urls": [ "https://www.um.edu.mt/library/oar/handle/123456789/99067", "https://api.crossref.org/works/10.1182/blood-2020-143116" ], "doi": "10.1182/blood-2020-143116", "pmid": null, "topics": [ "Globins & erythropoiesis" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reports an association between the HBG1 promoter variant rs368698783, within the LYAR-binding region, and HbF in Maltese cohorts. This is conference-level evidence.", "joseph_role": "Co-author", "verification": "Verified against publisher or University of Malta repository record.", "status": "published", "checked": "2026-10-23", "pages": "20-20", "volume": "136", "issue": "Supplement 1", "online_date": "2020-11-05", "finding": "Reports an association between the HBG1 promoter variant rs368698783, within the LYAR-binding region, and HbF in Maltese cohorts. This is conference-level evidence.", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Conference abstract: preliminary reported findings, with less methodological detail than a full research article. It is not treated as an independently replicated mechanism." }, { "id": "doi-blood-2020-143018", "title": "Genetic Heterogeneity of KLF1, a Master Regulator of Erythropoiesis, Revealed an Autosomal Recessive \u03a8\u03b2-Thalassemia and a Very Strong Promoter In Vivo", "year": 2020, "first_publication_year": 2020, "journal": "Blood", "authors": [ "Laura Grech", "Joseph Borg", "Ruth Galdies", "Carmen Attard", "Christian A. Scerri", "Sjaak Philipsen", "Alexander E. Felice" ], "author_line": "Laura Grech; Joseph Borg; Ruth Galdies; Carmen Attard; Christian A. Scerri; Sjaak Philipsen; Alexander E. Felice", "publication_type": "Conference abstract", "url": "https://doi.org/10.1182/blood-2020-143018", "source_url": "https://www.um.edu.mt/library/oar/handle/123456789/99065", "evidence_urls": [ "https://www.um.edu.mt/library/oar/handle/123456789/99065", "https://api.crossref.org/works/10.1182/blood-2020-143018" ], "doi": "10.1182/blood-2020-143018", "pmid": null, "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reports KLF1 promoter/coding haplotypes and variable phenotypes across additional Maltese families. This is a conference abstract, so its mechanistic proposals remain qualified.", "joseph_role": "Co-author", "verification": "Verified against publisher or University of Malta repository record.", "status": "published", "checked": "2026-10-23", "pages": "7-7", "volume": "136", "issue": "Supplement 1", "online_date": "2020-11-05", "finding": "Reports KLF1 promoter/coding haplotypes and variable phenotypes across additional Maltese families. This is a conference abstract, so its mechanistic proposals remain qualified.", "authorship_note": "Joseph Borg: Co-author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Conference abstract: preliminary reported findings, with less methodological detail than a full research article. It is not treated as an independently replicated mechanism." }, { "id": "pmid-33877003", "title": "Drug safety in thalassemia: lessons from the present and directions for the future", "year": 2021, "first_publication_year": 2021, "issue_date": "2021-08", "online_date": "2021-05-04", "journal": "Expert Opinion on Drug Safety", "journal_abbreviation": "Expert Opin Drug Saf", "volume": "20", "issue": "8", "pages": "937-947", "doi": "10.1080/14740338.2021.1919081", "pmid": "33877003", "url": "https://doi.org/10.1080/14740338.2021.1919081", "authors": [ "Laura Grech", "Janet Sultana", "Karen Borg", "Joseph Borg" ], "author_line": "Laura Grech; Janet Sultana; Karen Borg; Joseph Borg", "joseph_role": "Last-listed author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/33877003/" ], "publication_type": "Review", "indexed_publication_types": [ "Journal Article", "Review" ], "topics": [ "Globins & erythropoiesis", "Pharmacogenomics & therapeutics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reviews treatment safety in thalassaemia and the implications of emerging therapies for clinical management.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/33877003/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/33877003/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Department of Applied Biomedical Science, Faculty of Health Sciences, University of Malta, Imsida, Malta." ], "status": "published", "checked": "2026-10-23", "finding": "Reviews treatment safety in thalassaemia and the implications of emerging therapies for clinical management.", "authorship_note": "Joseph Borg: Last-listed author.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Review and synthesis of existing literature; not an independent experimental test of the family model." }, { "id": "hbbp1_rna", "from": "HBBP1 RNA / ELK1 pathway", "to": "HBG expression", "effect": "activation in tested models", "level": "cell-model overexpression, interaction and promoter assays", "finding": "HBBP1 overexpression increased gamma RNA and protein in HUDEP-2; the study implicates ELK1, which also binds and activates the gamma promoter.", "limit": "Keep distinct from HBBP1 DNA's architectural role. This does not establish the sign of changing endogenous HBBP1 RNA in every developmental context.", "source": "Ma et al. Biochemical and Biophysical Research Communications 2021", "doi": "10.1016/j.bbrc.2021.03.051", "url": "https://pubmed.ncbi.nlm.nih.gov/33744764/", "ref": "turn205search7", "input_section": "edges", "title": "Long noncoding RNA HBBP1 enhances \u03b3-globin expression through the ETS transcription factor ELK1", "authors": [ "Ma SP", "Xi HR", "Gao XX", "Yang JM", "Kurita R", "Nakamura Y", "Song XM", "Chen HY", "Lu DR" ], "authors_full": [ "Shuang-Ping Ma", "Hai-Rui Xi", "Xu-Xia Gao", "Jing-Min Yang", "Ryo Kurita", "Yukio Nakamura", "Xian-Min Song", "Hong-Yan Chen", "Da-Ru Lu" ], "year": 2021, "journal": "Biochemical and Biophysical Research Communications", "journal_abbreviation": "Biochem Biophys Res Commun", "volume": "552", "issue": null, "pages": "157-163", "pmid": "33744764", "pmcid": null, "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "Journal Article" ], "epmc_first_publication_date": "2021-03-19", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/33744764/", "https://europepmc.org/article/MED/33744764" ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Shuang-Ping Ma; Hai-Rui Xi; Xu-Xia Gao; Jing-Min Yang; Ryo Kurita; Yukio Nakamura; Xian-Min Song; Hong-Yan Chen; Da-Ru Lu", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Keep distinct from HBBP1 DNA's architectural role. This does not establish the sign of changing endogenous HBBP1 RNA in every developmental context.", "mechanistic": true, "seminal": false }, { "id": "hif_bglt3", "from": "HIF1alpha/HIF1beta", "to": "BGLT3 regulatory DNA", "effect": "activation", "level": "binding, base-edited motifs, primary-cell expression and contact assays", "finding": "HIF1 binds two BGLT3 hypoxia-response motifs, recruits activators, opens chromatin and promotes LCR-gamma interactions. Disrupting the motifs impairs gamma induction.", "limit": "HIF1A RNA is not a proxy for stabilized HIF1alpha protein. This pathway did not generate a uniform embryonic program: HBZ decreased in the VHL-disruption study.", "source": "Feng et al. Nature 2022", "doi": "10.1038/s41586-022-05312-w", "url": "https://pubmed.ncbi.nlm.nih.gov/36224385/", "ref": "turn188search0", "input_section": "edges", "title": "Activation of \u03b3-globin expression by hypoxia-inducible factor 1\u03b1", "authors": [ "Feng R", "Mayuranathan T", "Huang P", "Doerfler PA", "Li Y", "Yao Y", "Zhang J", "Palmer LE", "Mayberry K", "Christakopoulos GE", "Xu P", "Li C", "Cheng Y", "Blobel GA", "Simon MC", "Weiss MJ" ], "authors_full": [ "Ruopeng Feng", "Thiyagaraj Mayuranathan", "Peng Huang", "Phillip A Doerfler", "Yichao Li", "Yu Yao", "Jingjing Zhang", "Lance E Palmer", "Kalin Mayberry", "Georgios E Christakopoulos", "Peng Xu", "Chunliang Li", "Yong Cheng", "Gerd A Blobel", "M Celeste Simon", "Mitchell J Weiss" ], "year": 2022, "journal": "Nature", "journal_abbreviation": "Nature", "volume": "610", "issue": "7933", "pages": "783-790", "pmid": "36224385", "pmcid": "PMC9773321", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "research-article", "Journal Article", "Research Support, N.I.H., Extramural" ], "epmc_first_publication_date": "2022-10-12", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/36224385/", "https://europepmc.org/article/MED/36224385", "https://api.crossref.org/works/10.1038/s41586-022-05312-w" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2022, 10, 12 ], "crossref_online_date_parts": [ 2022, 10, 12 ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Ruopeng Feng; Thiyagaraj Mayuranathan; Peng Huang; Phillip A Doerfler; Yichao Li; Yu Yao; Jingjing Zhang; Lance E Palmer; Kalin Mayberry; Georgios E Christakopoulos; Peng Xu; Chunliang Li; Yong Cheng; Gerd A Blobel; M Celeste Simon; Mitchell J Weiss", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "HIF1A RNA is not a proxy for stabilized HIF1alpha protein. This pathway did not generate a uniform embryonic program: HBZ decreased in the VHL-disruption study.", "mechanistic": true, "seminal": false }, { "id": "competition", "from": "Adult HBB promoter activity", "to": "LCR availability for HBG", "effect": "competition", "level": "causal promoter mutagenesis and contact assays", "finding": "Disabling the HBB promoter increases HBG expression and redirects LCR contacts. Removing coding sequence while retaining the promoter has a much weaker effect.", "limit": "Competition is a major component; enhancer rearrangement, local silencing and deletion geometry also matter. Do not equate low HBB RNA with promoter deletion.", "source": "Topfer et al. Blood 2022", "doi": "10.1182/blood.2021014205", "url": "https://pubmed.ncbi.nlm.nih.gov/35090172/", "ref": "turn193search0", "geo": "GSE194064", "input_section": "edges", "title": "Disrupting the adult globin promoter alleviates promoter competition and reactivates fetal globin gene expression", "authors": [ "Topfer SK", "Feng R", "Huang P", "Ly LC", "Martyn GE", "Blobel GA", "Weiss MJ", "Quinlan KGR", "Crossley M" ], "authors_full": [ "Sarah K Topfer", "Ruopeng Feng", "Peng Huang", "Lana C Ly", "Gabriella E Martyn", "Gerd A Blobel", "Mitchell J Weiss", "Kate G R Quinlan", "Merlin Crossley" ], "year": 2022, "journal": "Blood", "journal_abbreviation": "Blood", "volume": "139", "issue": "14", "pages": "2107-2118", "pmid": "35090172", "pmcid": "PMC8990374", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "research-article", "Journal Article", "Research Support, N.I.H., Extramural" ], "epmc_first_publication_date": "2022-04-01", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/35090172/", "https://europepmc.org/article/MED/35090172", "https://api.crossref.org/works/10.1182/blood.2021014205" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2022, 4, 7 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Sarah K Topfer; Ruopeng Feng; Peng Huang; Lana C Ly; Gabriella E Martyn; Gerd A Blobel; Mitchell J Weiss; Kate G R Quinlan; Merlin Crossley", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Competition is a major component; enhancer rearrangement, local silencing and deletion geometry also matter. Do not equate low HBB RNA with promoter deletion.", "mechanistic": true, "seminal": false }, { "id": "pmid-35013432", "title": "Epigenomic analysis of KLF1 haploinsufficiency in primary human erythroblasts", "year": 2022, "first_publication_year": 2022, "issue_date": "2022-01-10", "online_date": "2022-01-10", "journal": "Scientific Reports", "journal_abbreviation": "Sci Rep", "volume": "12", "issue": "1", "pages": "336", "doi": "10.1038/s41598-021-04126-6", "pmid": "35013432", "url": "https://doi.org/10.1038/s41598-021-04126-6", "authors": [ "Steven Heshusius", "Laura Grech", "Nynke Gillemans", "Rutger W W Brouwer", "Xander T den Dekker", "Wilfred F J van IJcken", "Benjamin Nota", "Alex E Felice", "Thamar B van Dijk", "Marieke von Lindern", "Joseph Borg", "Emile van den Akker", "Sjaak Philipsen" ], "author_line": "Steven Heshusius; Laura Grech; Nynke Gillemans; Rutger W W Brouwer; Xander T den Dekker; Wilfred F J van IJcken; Benjamin Nota; Alex E Felice; Thamar B van Dijk; Marieke von Lindern; Joseph Borg; Emile van den Akker; Sjaak Philipsen", "joseph_role": "Joint senior author (equal contribution)", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/35013432/", "https://www.nature.com/articles/s41598-021-04126-6" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article", "Research Support, Non-U.S. Gov't" ], "topics": [ "Globins & erythropoiesis", "Molecular genetics" ], "is_globin": true, "timeline_recommended": true, "featured": true, "significance": "Combined RNA expression, chromatin accessibility and promoter assays in Maltese KLF1 p.K288X carriers. Variation in expression of the intact KLF1 allele was proposed to explain part of the HbF variability.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/35013432/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/35013432/", "https://www.nature.com/articles/s41598-021-04126-6" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Department of Applied Biomedical Science, Faculty of Health Sciences, Mater Dei Hospital, Msida, MSD2090, Malta." ], "status": "published", "checked": "2026-10-23", "finding": "Combined RNA expression, chromatin accessibility and promoter assays in Maltese KLF1 p.K288X carriers. Variation in expression of the intact KLF1 allele was proposed to explain part of the HbF variability.", "authorship_note": "Joseph Borg: Joint senior author (equal contribution).", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "pmid-34004015", "title": "Novel therapies in \u03b2-thalassaemia", "year": 2022, "first_publication_year": 2021, "issue_date": "2022-06", "online_date": "2021-06-04", "journal": "British Journal of Clinical Pharmacology", "journal_abbreviation": "Br J Clin Pharmacol", "volume": "88", "issue": "6", "pages": "2509-2524", "doi": "10.1111/bcp.14918", "pmid": "34004015", "url": "https://doi.org/10.1111/bcp.14918", "authors": [ "Laura Grech", "Karen Borg", "Joseph Borg" ], "author_line": "Laura Grech; Karen Borg; Joseph Borg", "joseph_role": "Last-listed author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/34004015/" ], "publication_type": "Review", "indexed_publication_types": [ "Journal Article", "Review" ], "topics": [ "Globins & erythropoiesis", "Pharmacogenomics & therapeutics" ], "is_globin": true, "timeline_recommended": true, "featured": false, "significance": "Reviews strategies to improve globin-chain balance and ineffective erythropoiesis, including pharmacological, gene-addition and gene-editing approaches.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/34004015/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/34004015/" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Centre for Molecular Medicine and Biobanking, University of Malta, Malta.", "Applied Biomedical Science, Faculty of Health Sciences, University of Malta, Malta." ], "status": "published", "checked": "2026-10-23", "finding": "Reviews strategies to improve globin-chain balance and ineffective erythropoiesis, including pharmacological, gene-addition and gene-editing approaches.", "authorship_note": "Joseph Borg: Last-listed author.", "date_note": "First published online 2021-06-04; journal issue 2022-06. Listed once under the issue year.", "website_topics": [ "globin", "genetics" ], "borg_author": true, "felice_author": false, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Review and synthesis of existing literature; not an independent experimental test of the family model." }, { "id": "bglt3_repression", "from": "BCL11A/LRF, G9a and CHD4", "to": "BGLT3", "effect": "repression", "level": "human erythroid knockout, knockdown, drug and chromatin experiments", "finding": "G9a-dependent chromatin repression and CHD4 recruitment connect BCL11A/LRF with BGLT3 repression. BGLT3 knockdown attenuated gamma induction after repressor knockout. BGLT3 overexpression increased gamma in HUDEP-2.", "limit": "The authors' universal wording refers to tested systems and inducers. RK-701 mouse embryonic-globin induction is not direct human BGLT3 gene-therapy evidence.", "source": "Takase et al. Nature Communications 2023", "doi": "10.1038/s41467-022-35404-0", "url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC9837035/", "ref": "turn185search0", "archive": "DRA014673", "input_section": "edges", "title": "A specific G9a inhibitor unveils BGLT3 lncRNA as a universal mediator of chemically induced fetal globin gene expression", "authors": [ "Takase S", "Hiroyama T", "Shirai F", "Maemoto Y", "Nakata A", "Arata M", "Matsuoka S", "Sonoda T", "Niwa H", "Sato S", "Umehara T", "Shirouzu M", "Nishigaya Y", "Sumiya T", "Hashimoto N", "Namie R", "Usui M", "Ohishi T", "Ohba SI", "Kawada M", "Hayashi Y", "Harada H", "Yamaguchi T", "Shinkai Y", "Nakamura Y", "Yoshida M", "Ito A" ], "authors_full": [ "Shohei Takase", "Takashi Hiroyama", "Fumiyuki Shirai", "Yuki Maemoto", "Akiko Nakata", "Mayumi Arata", "Seiji Matsuoka", "Takeshi Sonoda", "Hideaki Niwa", "Shin Sato", "Takashi Umehara", "Mikako Shirouzu", "Yosuke Nishigaya", "Tatsunobu Sumiya", "Noriaki Hashimoto", "Ryosuke Namie", "Masaya Usui", "Tomokazu Ohishi", "Shun-Ichi Ohba", "Manabu Kawada", "Yoshihiro Hayashi", "Hironori Harada", "Tokio Yamaguchi", "Yoichi Shinkai", "Yukio Nakamura", "Minoru Yoshida", "Akihiro Ito" ], "year": 2023, "journal": "Nature Communications", "journal_abbreviation": "Nat Commun", "volume": "14", "issue": "1", "pages": "23", "pmid": "36635268", "pmcid": "PMC9837035", "indexed_publication_types": [ "Research Support, Non-U.S. Gov't", "research-article", "Journal Article" ], "epmc_first_publication_date": "2023-01-12", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/36635268/", "https://europepmc.org/article/MED/36635268", "https://api.crossref.org/works/10.1038/s41467-022-35404-0" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2023, 1, 12 ], "crossref_online_date_parts": [ 2023, 1, 12 ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Shohei Takase; Takashi Hiroyama; Fumiyuki Shirai; Yuki Maemoto; Akiko Nakata; Mayumi Arata; Seiji Matsuoka; Takeshi Sonoda; Hideaki Niwa; Shin Sato; Takashi Umehara; Mikako Shirouzu; Yosuke Nishigaya; Tatsunobu Sumiya; Noriaki Hashimoto; Ryosuke Namie; Masaya Usui; Tomokazu Ohishi; Shun-Ichi Ohba; Manabu Kawada; Yoshihiro Hayashi; Hironori Harada; Tokio Yamaguchi; Yoichi Shinkai; Yukio Nakamura; Minoru Yoshida; Akihiro Ito", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "publication_date": "2023-01-12", "date_note": "Published 12 January 2023; the 2022 component of the DOI is not the publication year.", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "The authors' universal wording refers to tested systems and inducers. RK-701 mouse embryonic-globin induction is not direct human BGLT3 gene-therapy evidence.", "mechanistic": true, "seminal": false }, { "id": "pmid-38862545", "title": "Spatiotemporal expression and control of haemoglobin in space", "year": 2024, "first_publication_year": 2024, "issue_date": "2024-06-11", "online_date": "2024-06-11", "journal": "Nature Communications", "journal_abbreviation": "Nat Commun", "volume": "15", "issue": "1", "pages": "4927", "doi": "10.1038/s41467-024-49289-8", "pmid": "38862545", "url": "https://doi.org/10.1038/s41467-024-49289-8", "authors": [ "Josef Borg", "Conor Loy", "JangKeun Kim", "Alfred Buhagiar", "Christopher Chin", "Namita Damle", "Iwijn De Vlaminck", "Alex Felice", "Tammy Liu", "Irina Matei", "Cem Meydan", "Masafumi Muratani", "Omary Mzava", "Eliah Overbey", "Krista A Ryon", "Scott M Smith", "Braden T Tierney", "Guy Trudel", "Sara R Zwart", "Afshin Beheshti", "Christopher E Mason", "Joseph Borg" ], "author_line": "Josef Borg; Conor Loy; JangKeun Kim; Alfred Buhagiar; Christopher Chin; Namita Damle; Iwijn De Vlaminck; Alex Felice; Tammy Liu; Irina Matei; Cem Meydan; Masafumi Muratani; Omary Mzava; Eliah Overbey; Krista A Ryon; Scott M Smith; Braden T Tierney; Guy Trudel; Sara R Zwart; Afshin Beheshti; Christopher E Mason; Joseph Borg", "joseph_role": "Last-listed and co-corresponding author", "role_evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/38862545/", "https://www.nature.com/articles/s41467-024-49289-8" ], "publication_type": "Journal article", "indexed_publication_types": [ "Journal Article", "Research Support, N.I.H., Extramural" ], "topics": [ "Globins & erythropoiesis", "Space bioscience" ], "is_globin": true, "timeline_recommended": true, "featured": true, "significance": "Integrated datasets from NASA GeneLab, the NASA Twins Study, JAXA and Inspiration4 to examine globin expression across spaceflight contexts. The study establishes context-dependent responses rather than a closed universal switching mechanism.", "source_url": "https://pubmed.ncbi.nlm.nih.gov/38862545/", "evidence_urls": [ "https://pubmed.ncbi.nlm.nih.gov/38862545/", "https://www.nature.com/articles/s41467-024-49289-8" ], "verification": "Bibliographic metadata retrieved from NCBI PubMed XML. Correct researcher checked by author-specific Malta biomedical affiliation, ORCID or the corroborated globin collaboration.", "joseph_affiliations": [ "Faculty of Health Sciences, University of Malta, Msida, MSD2080, Malta. joseph.j.borg@um.edu.mt." ], "status": "published", "checked": "2026-10-23", "finding": "Integrated datasets from NASA GeneLab, the NASA Twins Study, JAXA and Inspiration4 to examine globin expression across spaceflight contexts. The study establishes context-dependent responses rather than a closed universal switching mechanism.", "authorship_note": "Joseph Borg: Last-listed and co-corresponding author.", "website_topics": [ "globin", "space" ], "borg_author": true, "felice_author": true, "huisman_author": false, "schroeder_author": false, "seminal": false, "model_relevance": "A contribution to the continuing Maltese programme in globin biology and haemoglobinopathies.", "limitation": "Interpretation is restricted to the study design and specimens reported in this publication; it does not establish the unresolved modifier in the Maltese family." }, { "id": "distance", "from": "Enhancer proximity", "to": "HBG/HBE/HBZ activation", "effect": "activation after engineered rearrangement", "level": "causal deletion/inversion in adult erythroid cells and ex-vivo differentiated HSPCs", "finding": "Bringing strong enhancers closer through intervening-sequence deletion or inversion can override developmental silencing at beta-like and alpha-like globin loci.", "limit": "This is not evidence of such rearrangements in the Maltese family and does not identify a single shared transcription factor.", "source": "Felder et al. Blood 2025", "doi": "10.1182/blood.2024028128", "url": "https://pubmed.ncbi.nlm.nih.gov/40456162/", "ref": "turn197search4", "geo": "GSE274030; GSE274029", "input_section": "edges", "title": "Reactivation of developmentally silenced globin genes through forced linear recruitment of remote enhancers", "authors": [ "Felder AK", "Tjalsma SJD", "Verhagen HJMP", "Majied R", "Verstegen MJAM", "Verheul TCJ", "van Haren J", "Mohnani R", "Gremmen R", "Krijger PHL", "Philipsen S", "van den Akker E", "de Laat W" ], "authors_full": [ "Anna-Karina Felder", "Sjoerd J D Tjalsma", "Han J M P Verhagen", "Rezin Majied", "Marjon J A M Verstegen", "Thijs C J Verheul", "Jeffrey van Haren", "Rebecca Mohnani", "Richard Gremmen", "Peter H L Krijger", "Sjaak Philipsen", "Emile van den Akker", "Wouter de Laat" ], "year": 2025, "journal": "Blood", "journal_abbreviation": "Blood", "volume": "146", "issue": "6", "pages": "732-744", "pmid": "40456162", "pmcid": "PMC12824704", "indexed_publication_types": [ "research-article", "Journal Article" ], "epmc_first_publication_date": "2025-08-01", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/40456162/", "https://europepmc.org/article/MED/40456162" ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Anna-Karina Felder; Sjoerd J D Tjalsma; Han J M P Verhagen; Rezin Majied; Marjon J A M Verstegen; Thijs C J Verheul; Jeffrey van Haren; Rebecca Mohnani; Richard Gremmen; Peter H L Krijger; Sjaak Philipsen; Emile van den Akker; Wouter de Laat", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "publication_date": "2025-08-07", "date_note": "Blood issue date verified on PubMed; year 2025 despite 2024 in DOI.", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "This is not evidence of such rearrangements in the Maltese family and does not identify a single shared transcription factor.", "mechanistic": true, "seminal": false }, { "id": "igf2bp1", "from": "IGF2BP1", "to": "HIC2/BCL11A and HBG RNA translation", "effect": "supports fetal hemoglobin", "level": "human erythroid perturbation and RNA-binding/translation assays", "finding": "IGF2BP1 supports HIC2, a BCL11A repressor, and directly supports HBG1/2 translation through stop-codon-proximal m6A-containing RNA elements.", "limit": "A new 2026 mechanistic layer; family mediation is untested. This specifically cautions against equating RNA fractions with hemoglobin protein fractions.", "source": "Coyne et al. Blood 2026", "doi": "10.1182/blood.2025031840", "url": "https://pubmed.ncbi.nlm.nih.gov/42341077/", "ref": "turn205search1", "geo": "GSE303945", "publication_date": "2026-09-24", "input_section": "edges", "title": "Direct and indirect regulation of fetal globin transcript by the RNA-binding protein IGF2BP1", "authors": [ "Coyne S", "Hwang GH", "Wu T", "Hossain M", "Vinjamur D", "Starrs M", "Zeng J", "Andresen F", "Gutierrez A", "Shimamura A", "Bauer DE" ], "authors_full": [ "Steven Coyne", "Gue-Ho Hwang", "Ting Wu", "Mir Hossain", "Divya Vinjamur", "Marlena Starrs", "Jing Zeng", "Felicia Andresen", "Ashley Gutierrez", "Akiko Shimamura", "Daniel E Bauer" ], "year": 2026, "journal": "Blood", "journal_abbreviation": "Blood", "volume": "148", "issue": "13", "pages": "1737-1749", "pmid": "42341077", "pmcid": "PMC13642966", "indexed_publication_types": [ "research-article", "Journal Article" ], "epmc_first_publication_date": "2026-09-01", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/42341077/", "https://europepmc.org/article/MED/42341077", "https://api.crossref.org/works/10.1182/blood.2025031840" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2026, 9, 24 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Steven Coyne; Gue-Ho Hwang; Ting Wu; Mir Hossain; Divya Vinjamur; Marlena Starrs; Jing Zeng; Felicia Andresen; Ashley Gutierrez; Akiko Shimamura; Daniel E Bauer", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "date_note": "Blood issue date confirmed by PubMed, publisher and Crossref; earlier online appearance in June 2026. Eligible before 23 October 2026.", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "A new 2026 mechanistic layer; family mediation is untested. This specifically cautions against equating RNA fractions with hemoglobin protein fractions.", "mechanistic": true, "seminal": false }, { "id": "lrf_reversible", "from": "LRF protein loss and restoration", "to": "HBG repression state", "effect": "reversible", "level": "controlled protein degradation and restoration in HUDEP-2", "finding": "Gamma activation after LRF depletion can reverse when LRF returns, including RNA and chromatin changes; LRF-NuRD association persists without BCL11A.", "limit": "Demonstrated in immature erythroid models; not a direct measurement of family T0-to-T48 mechanism.", "source": "Miyazaki et al. Blood Red Cells & Iron 2026", "doi": "10.1016/j.brci.2026.100087", "url": "https://www.sciencedirect.com/science/article/pii/S3050598426000429", "ref": "turn197search0", "geo": "GSE309998; GSE309999", "input_section": "edges", "title": "Reversible regulation of \u03b3-globin expression by LRF through a BCL11A-independent mechanism", "authors": [ "Manaka Miyazaki", "Kiyoko Setoguchi", "Takeshi Masuda", "Takuya Morikawa", "Stuti Mehta", "Shinya Matsumoto", "Yuya Kunisaki", "Stuart H. Orkin", "Yuichiro Semba", "Takahiro Maeda" ], "authors_full": [ "Manaka Miyazaki", "Kiyoko Setoguchi", "Takeshi Masuda", "Takuya Morikawa", "Stuti Mehta", "Shinya Matsumoto", "Yuya Kunisaki", "Stuart H. Orkin", "Yuichiro Semba", "Takahiro Maeda" ], "year": 2026, "journal": "Blood Red Cells & Iron", "volume": "2", "issue": "3", "pages": "100087", "pmid": null, "pmcid": null, "indexed_publication_types": [ "journal-article" ], "publication_type": "Original study", "publication_format": "Short communication", "verification_sources": [ "https://api.crossref.org/works/10.1016/j.brci.2026.100087", "https://www.sciencedirect.com/science/article/pii/S3050598426000429", "https://www.sciencedirect.com/journal/blood-red-cells-and-iron/vol/2/issue/3", "https://hyoka.ofc.kyushu-u.ac.jp/html/100019616_en.html?k=Y" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2026, 9 ], "crossref_online_date_parts": [], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Manaka Miyazaki; Kiyoko Setoguchi; Takeshi Masuda; Takuya Morikawa; Stuti Mehta; Shinya Matsumoto; Yuya Kunisaki; Stuart H. Orkin; Yuichiro Semba; Takahiro Maeda", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "publication_date": "2026-09", "date_precision": "month", "date_note": "Crossref and publisher issue list: volume 2, issue 3, September 2026. Institutional profiles list April or July 2026, likely earlier online/reporting dates; exact first-online day is not asserted. All are before 23 October 2026.", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Demonstrated in immature erythroid models; not a direct measurement of family T0-to-T48 mechanism.", "mechanistic": true, "seminal": false }, { "source": "Rahaman et al. Life Science Alliance 2026", "doi": "10.26508/lsa.202603620", "url": "https://pubmed.ncbi.nlm.nih.gov/42373543/", "finding": "UCA1/miR-148b modulates BCL11A and gamma in HUDEP-2 and CD34-derived cells; optional additional RNA regulatory layer, not a Maltese-family finding.", "ref": "turn197search3", "input_section": "optional_sources", "title": "UCA1 lncRNA regulates \u03b3-globin expression by modulating the miR-148b/BCL11A axis", "authors": [ "Rahaman M", "Bhattacharya S", "Mukherjee M", "Bhowmick C", "Shukla PC", "Dolai TK", "Chakravorty N" ], "authors_full": [ "Motiur Rahaman", "Shatarupa Bhattacharya", "Mandrita Mukherjee", "Chiranjib Bhowmick", "Praphulla Chandra Shukla", "Tuphan Kanti Dolai", "Nishant Chakravorty" ], "year": 2026, "journal": "Life Science Alliance", "journal_abbreviation": "Life Sci Alliance", "volume": "9", "issue": "9", "pages": "e202603620", "pmid": "42373543", "pmcid": "PMC13315483", "indexed_publication_types": [ "research-article", "Journal Article" ], "epmc_first_publication_date": "2026-06-29", "publication_type": "Original study", "verification_sources": [ "https://pubmed.ncbi.nlm.nih.gov/42373543/", "https://europepmc.org/article/MED/42373543", "https://api.crossref.org/works/10.26508/lsa.202603620" ], "crossref_type": "journal-article", "crossref_published_date_parts": [ 2026, 6, 29 ], "crossref_online_date_parts": [ 2026, 6, 29 ], "verified_at": "2026-10-23", "published_by_cutoff": true, "verification_note": "DOI matched to exact bibliographic metadata; original finding, limit and evidence-level wording preserved unchanged. Europe PMC synthetic first-of-month dates are retained only as raw metadata, not asserted as exact publication days.", "author_string": "Motiur Rahaman; Shatarupa Bhattacharya; Mandrita Mukherjee; Chiranjib Bhowmick; Praphulla Chandra Shukla; Tuphan Kanti Dolai; Nishant Chakravorty", "joseph_borg_author": false, "felice_author": false, "huisman_author": false, "schroeder_author": false, "publication_date": "2026-06-29", "issue_date": "2026-09", "date_note": "Online publication 29 June 2026; print/issue September 2026, as stated by PubMed and Crossref.", "id": "modern_optional_42373543", "model_relevance": "Experimental evidence for the regulatory network explored in the atlas.", "limitation": "Scope is limited to the reported experimental systems.", "mechanistic": true, "seminal": false } ], "additional_context": [ { "year": 1970, "title": "Changes in the Nature of Human Fetal Hemoglobin During the First Year of Life", "authors": [ "Titus H J Huisman", "Walter A Schroeder", "Audrey K Brown" ], "journal": "Pediatr Res. 1970;4:466\u2013467", "doi": "10.1203/00006450-197009000-00130", "url": "https://www.nature.com/articles/pr1970174", "web_refs": [ "turn212search0" ], "note": "Short published abstract preceding the 1971 article; seven infants showed G\u03b3:A\u03b3 near 3:1 at birth and 2:3 by around five months. Ratios concern gamma-chain composition, not total HbF." }, { "year": 1990, "title": "The Brazilian type of nondeletional A gamma-fetal hemoglobin has a C----G substitution at nucleotide \u2212195 of the A gamma-globin gene", "authors": [ "F F Costa", "M A Zago", "G Cheng", "J F Nechtman", "T A Stoming", "T H Huisman" ], "journal": "Blood. 1990;76(9):1896\u20131897", "pmid": "2224140", "url": "https://pubmed.ncbi.nlm.nih.gov/2224140/", "web_refs": [ "turn202search2" ], "note": "Additional primary promoter-variant report; correspondence with no PubMed abstract. Use 1990, not the incorrect 1988 date found in some secondary citations." }, { "year": 2023, "title": "The Beautiful World of Human Haemoglobin (revisited)", "authors": [ "Alex E Felice" ], "journal": "Xjenza Online. 2023;11 Special Issue:102\u2013119", "url": "https://www.um.edu.mt/library/oar/bitstream/123456789/123710/1/The_Beautiful_World_of_Human_Haemoglobin_2023.pdf", "web_refs": [ "turn202view1" ], "type": "First-person historical review; not primary experimental evidence", "note": "Felice describes his Medical College of Georgia/Augusta work during 1976\u20131989, his 1981 PhD under Huisman, and his return to Malta in 1989. Good provenance for an Augusta-to-Malta historical panel." } ], "reviewed": "2026-10-23", "editorial_notes": [ "Use measured contributions rather than describing every included paper as seminal. Milestones, supporting studies, review and resource entries are separately typed.", "The 1977 Nature paper lists Felice at Royal University of Malta, Huisman at Medical College of Georgia and Schroeder at Caltech. Do not replace these publication-level affiliations with an assumed Augusta affiliation for Felice.", "The 1972 four-gamma-gene paper is historically relevant but its proposed normal four-nonallelic-gene model is superseded. Preserve that distinction visibly if included.", "Joseph Borg coauthored the 2007 cis\u2013trans/HU review; his verified globin bibliography begins earlier than 2009. The 1997 Hb Setif paper is by Isabella Borg, not Joseph Borg.", "The 1982 K562 paper does not demonstrate a common upstream switch factor for the Maltese family; cell line, culture, native development and patient blood are distinct contexts.", "The 1971 G\u03b3:A\u03b3 ratios describe the composition of HbF, not percentage HbF among all hemoglobins.", "The 2003 Hb St. Luke's entry is a comment/letter and the 2009 ITHANET paper is infrastructure; neither is a new switching mechanism.", "Do not describe Joseph Borg as a Nobel laureate or imply an award he has not received.", "The 2010 KLF1 paper and 2011 microattribution paper are the two verified Nature Genetics contributions. They answer different scientific questions.", "The verified EMBO publication is EMBO Reports, not EMBO Journal.", "Blood supplement publications in this catalogue are conference abstracts, not full-length peer-reviewed research articles.", "The 2024 haemoglobin-space paper has Josef Borg as a co-first author and Joseph Borg as a last-listed/co-corresponding author. Keep these two researchers distinct.", "A visual continuum can group research into eras without inventing annual milestones or overstating causality." ], "date_policy": "year follows indexed issue/publication year; online_date and first_publication_year retain online-first dates. Do not duplicate the 2025/2026 TGFBI paper or the 2021/2022 thalassaemia review." }, "provenance": { "family": "Accepted FamF v9 source package and 23 October graded-pattern audit. Seven donors \u00d7 T0/T48; repeated processing views are sensitivity analyses, not biological replicates.", "rna": "hg19; NH-weighted first-strand primary view. Locus values are log2(count/geometric mean of erythroid reference counts). Ref: heme3 = ALAS2/FECH/AHSP; membrane6 = SLC4A1/GYPA/EPB42/SPTA1/SPTB/ANK1; combined9 = both.", "regulators": "Inherited accepted log2 RNA estimates, with optional subtraction of erythroid reference-panel mean log2 expression. These are not protein activities. ZFP57 zeros are inherited transformed estimates, not proof of absence.", "atac": "GRCh38 promoter windows \u00b1250/\u00b11000 bp; inherited log2(normalized fragment signal + 0.5) values. ATAC measures accessibility, not factor occupancy or looping. Broad-window estimates are sensitive to filtering/window choice.", "coordinates": "GRCh38 zero-based half-open window intervals. RNA and ATAC joined by verified gene names; no na\u00efve cross-build coordinate join. Locus illustration is transcriptional order, reverse genomic orientation, not to scale.", "time": "Family T0/T48 are culture time points, not fetal/adult stages. Intermediate frames interpolate log2 values only; endpoints alone are observed.", "development": "Two independent public RNA panels are compared within study and matched culture day. No cross-study quantitative developmental curve is fitted.", "protein": "Nikita Camilleri mature circulating RBC data: 22 donors, 11 carriers and 11 controls, family-adjusted association across 1,600 quantified proteins. No sample-level join to all four HEP donors. Intensities are not molar globin fractions; imputation/peptide specificity limits apply.", "genetics": "Filtered WGS and SNP exports constrain candidates but do not uniquely identify a modifier. M8 source DNA is not verified; conditional phase is not direct molecular phase.", "animation": "Normal-development curve heights, contact weights and particle speeds are illustrative. Family expression values are measured summaries, but drawn contacts are literature-informed hypotheses, not patient contact measurements.", "identifiability": "FamF high group has one donor (M2) versus M4/M5. M8 is an external cultured-HEP comparator. No causal coefficients, treatment predictions or new family significance tests are inferred.", "hashes": { "ATAC_Atlas.csv": "2e7a39e15e8ed00ff18b98379736219fc1c62bbe93057cf442555a685a1dee07", "ATAC_Module_Counts.csv": "5a1d7875ebb74eb5eb22e3018ba5e161faae84121c838ef7ef36aa1fc590d14c", "ATAC_Region_Values.csv": "f6777c8bdfc7f07dd13f399843bedb99d827e69ed940593584a77c9f0e98b538", "Array_M2_vs_Other_Carriers.csv": "e0cd370280a4e088bef7ca504f4d16a6eecc8f00d6e8c12c4cf3777c1683e457", "Canonical_Repressor_Anchor_Support.csv": "e41d2b2ad14f88d380e1724801ca7c70227f54de84e81eaa60532768f3d56f95", "Family_Summary.json": "94f436dcbfda5ba880a2f31cc56a30d6bf0b689f11fcba169cc431943bdf5c56", "Genetics_qPCR_Summary.json": "0ba88fb97f33f3a9fbc53db8671f832d4a7d73b79a084bbd4355bacfeb924772", "KLF1_Allele_Proxy.csv": "850a42d0032d586f469b6c2566620e4aa6f73bd7517905e8e86176ad531db06d", "Locus_RNA.csv": "236c4267a9b971b03c348a41a861169370d25a244711b755ab852d35bdcb638b", "Proteomics_family_adjusted_effects.csv": "ea332c5f48d7419db3c0b643b9afa253e0186e7aa0a46e0b4bfeac94b8eba0e7", "Published_Phenotypes.csv": "a1fe5433ebd0bbd6c3c0fafb22480719b1fd50e8c236b6c18322680571895a6b", "Regulator_RNA_Values.csv": "0fd12ff9619f81ea0929a36f97e751cdfd06b1cae7b1432f22ed8d0fe9c7eeae", "Selected_RNA_donor_values.csv": "0018004b4b902b0444bbba287e10b4a0c8c1dad84ef31dc0e16081d450ab09a3", "development_panel.json": "ed1265287bd60b3cb7ac32f766387efa598ac26a1efae46f593fcfaed769aa02", "history.json": "76c93f3ce6b2e2008526f6d5758886923bbac03e9ed79222e15dcc6c116f73ad", "modern_evidence.json": "94dbc66b045bf224b7813e8e5ee69aea6c3b687bf534ef708bc66852d6bf6c09", "nurd_evidence.json": "4e3b07f733840eac60313d563fd0320407ab1a59fdefb9f0c50225342667a5f9" } } }