Table 7.

Autoinflammatory disorders

DiseaseGenetic defectInheritanceOMIMT cellsB cellsFunctional defectAssociated features
1. Type 1 interferonopathies 
AD STING-associated vasculopathy, infantile-onset (SAVI) TMEM173** (STING) AD 612374 Not assessed Not assessed STING activates both the NF-κB and IRF3 transcription pathways to induce the expression of IFN Skin vasculopathy, inflammatory lung disease, systemic autoinflammation and ICC, FCL 
AR STING-associated vasculopathy, infantile-onset (SAVI) AR GOF 615934 Not assessed Not assessed STING activates both the NF-κB and IRF3 transcription pathways to induce the expression of IFN FTT, early-onset rash, fever, dyspnea, interstitial lung disease/pneumonitis, polyarthritis, autoAbs, increased inflammatory markers, IFN gene signature. Phenocopy of SAVI due to AD GOF TMEM173 
ADA2 deficiency ADA2 AR 607575 Not assessed Not assessed ADAs deactivate extracellular adenosine and terminate signaling through adenosine receptors Polyarteritis nodosa, childhood-onset, early-onset recurrent ischemic stroke and fever; some patients develop hypogammaglobulinemia 
TREX1 deficiency, Aicardi–Goutières syndrome 1 (AGS1) TREX1 AR 606609 Not assessed Not assessed Intracellular accumulation of abnormal ssDNA species leading to increased type I IFN production Classical AGS, SLE, FCL 
AD 
RNASEH2B deficiency, AGS2 RNASEH2B AR 610326 Not assessed Not assessed Intracellular accumulation of abnormal RNA-DNA hybrid species leading to increased type I IFN production Classical AGS, SP 
RNASEH2C deficiency, AGS3 RNASEH2C AR 610330 Not assessed Not assessed Classical AGS 
RNASEH2A deficiency, AGS4 RNASEH2A AR 606034 Not assessed Not assessed Classical AGS 
SAMHD1 deficiency, AGS5 SAMHD1 AR 606754 Not assessed Not assessed Controls dNTPs in the cytosol, failure of which leads to increased type I IFN production Classical AGS, FCL 
ADAR1 deficiency, AGS6 ADAR1 AR 615010 Not assessed Not assessed Catalyzes the deamination of adenosine to inosine in dsRNA substrates, failure of which leads to increased type I IFN production Classical AGS, BSN, SP 
AD (G1007R) NA 
Aicardi–Goutières syndrome 7 (AGS7) IFIH1 AD GOF 615846 Not assessed Not assessed IFIH1 gene encodes a cytoplasmic viral RNA receptor that activates type I interferon signaling through the MAVS adaptor molecule Classical AGS, SLE, SP, SMS 
DNase II deficiency DNASE2 AR 619858 Not assessed Not assessed DNase II degrades and eliminates DNA. Loss of DNase II activity induces type I interferon signaling AGS 
LSM11 deficiency LSM11 AR 619486 Not assessed Not assessed Increased IFN signaling in fibroblasts AGS, type 1 IFN-opathy 
RNU7-1 deficiency RNU7-1 AR 619487 Not assessed Not assessed Increased IFN signaling in fibroblasts AGS, type 1 IFN-opathy 
ARF1 deficiency ARF1 AD 103180 Not assessed Not assessed Increased type I IFN signaling in cell lines and patient cells AGS, type 1 IFN-opathy 
Pediatric SLE due to DNASE1L3 deficiency DNASE1L3 AR 614420   DNASE1L3 is an endonuclease that degrades extracellular DNA. DNASE1L3 deficiency decreases clearance of apoptotic cells Very early-onset SLE, reduced complement levels, autoantibodies (dsDNA, ANCA), lupus nephritis, hypocomplementemic urticarial vasculitis syndrome 
Spondyloenchondrodysplasia with immune dysregulation (SPENCD) ACP5 AR 171640 Not assessed Not assessed Upregulation of IFN through mechanism possibly relating to pDCS Short stature, SP, ICC, SLE, thrombocytopenia and autoimmune hemolytic anemia, possibly recurrent bacterial and viral infections 
USP18 deficiency USP18 AR 607057 Not assessed Not assessed Defective negative regulation of ISG15 leading to increased IFN TORCH-like syndrome, autoinflammation, and mycobacterial disease 
OAS1 GOF OAS1 AD GOF 618042  Low Increased interferon from recognition of RNA Pulmonary alveolar proteinosis, skin rash 
CDC42 deficiency CDC42 AD 616737 Normal/decreased Normal/decreased ↑serum levels of IL-1, IL-18, IFN-α↑, ferritin, sCD25, CRP, etc. Mutation affects actin function, ↓ NK cell cytotoxicity Neonatal onset: pancytopenia, fever, rash, hepatosplenomegaly, multisystemic inflammation, myelofibrosis/proliferation, HLH, enterocolitis; recurrent GIT/URT infections; neurodevelopmental delay, FTT 
STAT2 loss of negative regulation STAT2 AR 616636 Increased Normal Patient cells hypersensitive to IFN-α, GOF for induction of the late (not early) response to type 1 IFNs due to impaired interaction of mutant STAT2 with USP18, a negative regulator of type 1 IFN responses Severe fatal early-onset autoinflammation, ↑serum IFN-α, IL-6, TNF-α, phenocopy of USP18 deficiency 
ATAD3A deficiency ATAD3A AD/AR 617183 Not assessed Not assessed Elevated ISG expression, increased serum type 1 IFNs Predominantly neurological defects (development delay, spasticity) 
RELA haploinsufficiency RELA AD 618287 Normal/increased Normal  Chronic mucocutaneous ulceration
Impaired NF-κB activation; reduced production of inflammatory cytokines 
RELA interferonopathya RELA AD DN 618287   Leukocyte TLR7-dependent type I/III IFN production Patients with RELA DN mutations shared clinical phenotypes with RELA haploinsufficiency, presenting chronic mucocutaneous ulcerations and autoimmune hematological disorders such as immune thrombocytopenia (ITP) and neutropenia. However, patients with RELA DN mutations additionally presented periodic fever, IBD juvenile idiopathic arthritis (JIA), and skin involvement 
DiseaseGenetic defectInheritanceOMIMT cellsB cellsFunctional defectAssociated features
1. Type 1 interferonopathies 
AD STING-associated vasculopathy, infantile-onset (SAVI) TMEM173** (STING) AD 612374 Not assessed Not assessed STING activates both the NF-κB and IRF3 transcription pathways to induce the expression of IFN Skin vasculopathy, inflammatory lung disease, systemic autoinflammation and ICC, FCL 
AR STING-associated vasculopathy, infantile-onset (SAVI) AR GOF 615934 Not assessed Not assessed STING activates both the NF-κB and IRF3 transcription pathways to induce the expression of IFN FTT, early-onset rash, fever, dyspnea, interstitial lung disease/pneumonitis, polyarthritis, autoAbs, increased inflammatory markers, IFN gene signature. Phenocopy of SAVI due to AD GOF TMEM173 
ADA2 deficiency ADA2 AR 607575 Not assessed Not assessed ADAs deactivate extracellular adenosine and terminate signaling through adenosine receptors Polyarteritis nodosa, childhood-onset, early-onset recurrent ischemic stroke and fever; some patients develop hypogammaglobulinemia 
TREX1 deficiency, Aicardi–Goutières syndrome 1 (AGS1) TREX1 AR 606609 Not assessed Not assessed Intracellular accumulation of abnormal ssDNA species leading to increased type I IFN production Classical AGS, SLE, FCL 
AD 
RNASEH2B deficiency, AGS2 RNASEH2B AR 610326 Not assessed Not assessed Intracellular accumulation of abnormal RNA-DNA hybrid species leading to increased type I IFN production Classical AGS, SP 
RNASEH2C deficiency, AGS3 RNASEH2C AR 610330 Not assessed Not assessed Classical AGS 
RNASEH2A deficiency, AGS4 RNASEH2A AR 606034 Not assessed Not assessed Classical AGS 
SAMHD1 deficiency, AGS5 SAMHD1 AR 606754 Not assessed Not assessed Controls dNTPs in the cytosol, failure of which leads to increased type I IFN production Classical AGS, FCL 
ADAR1 deficiency, AGS6 ADAR1 AR 615010 Not assessed Not assessed Catalyzes the deamination of adenosine to inosine in dsRNA substrates, failure of which leads to increased type I IFN production Classical AGS, BSN, SP 
AD (G1007R) NA 
Aicardi–Goutières syndrome 7 (AGS7) IFIH1 AD GOF 615846 Not assessed Not assessed IFIH1 gene encodes a cytoplasmic viral RNA receptor that activates type I interferon signaling through the MAVS adaptor molecule Classical AGS, SLE, SP, SMS 
DNase II deficiency DNASE2 AR 619858 Not assessed Not assessed DNase II degrades and eliminates DNA. Loss of DNase II activity induces type I interferon signaling AGS 
LSM11 deficiency LSM11 AR 619486 Not assessed Not assessed Increased IFN signaling in fibroblasts AGS, type 1 IFN-opathy 
RNU7-1 deficiency RNU7-1 AR 619487 Not assessed Not assessed Increased IFN signaling in fibroblasts AGS, type 1 IFN-opathy 
ARF1 deficiency ARF1 AD 103180 Not assessed Not assessed Increased type I IFN signaling in cell lines and patient cells AGS, type 1 IFN-opathy 
Pediatric SLE due to DNASE1L3 deficiency DNASE1L3 AR 614420   DNASE1L3 is an endonuclease that degrades extracellular DNA. DNASE1L3 deficiency decreases clearance of apoptotic cells Very early-onset SLE, reduced complement levels, autoantibodies (dsDNA, ANCA), lupus nephritis, hypocomplementemic urticarial vasculitis syndrome 
Spondyloenchondrodysplasia with immune dysregulation (SPENCD) ACP5 AR 171640 Not assessed Not assessed Upregulation of IFN through mechanism possibly relating to pDCS Short stature, SP, ICC, SLE, thrombocytopenia and autoimmune hemolytic anemia, possibly recurrent bacterial and viral infections 
USP18 deficiency USP18 AR 607057 Not assessed Not assessed Defective negative regulation of ISG15 leading to increased IFN TORCH-like syndrome, autoinflammation, and mycobacterial disease 
OAS1 GOF OAS1 AD GOF 618042  Low Increased interferon from recognition of RNA Pulmonary alveolar proteinosis, skin rash 
CDC42 deficiency CDC42 AD 616737 Normal/decreased Normal/decreased ↑serum levels of IL-1, IL-18, IFN-α↑, ferritin, sCD25, CRP, etc. Mutation affects actin function, ↓ NK cell cytotoxicity Neonatal onset: pancytopenia, fever, rash, hepatosplenomegaly, multisystemic inflammation, myelofibrosis/proliferation, HLH, enterocolitis; recurrent GIT/URT infections; neurodevelopmental delay, FTT 
STAT2 loss of negative regulation STAT2 AR 616636 Increased Normal Patient cells hypersensitive to IFN-α, GOF for induction of the late (not early) response to type 1 IFNs due to impaired interaction of mutant STAT2 with USP18, a negative regulator of type 1 IFN responses Severe fatal early-onset autoinflammation, ↑serum IFN-α, IL-6, TNF-α, phenocopy of USP18 deficiency 
ATAD3A deficiency ATAD3A AD/AR 617183 Not assessed Not assessed Elevated ISG expression, increased serum type 1 IFNs Predominantly neurological defects (development delay, spasticity) 
RELA haploinsufficiency RELA AD 618287 Normal/increased Normal  Chronic mucocutaneous ulceration
Impaired NF-κB activation; reduced production of inflammatory cytokines 
RELA interferonopathya RELA AD DN 618287   Leukocyte TLR7-dependent type I/III IFN production Patients with RELA DN mutations shared clinical phenotypes with RELA haploinsufficiency, presenting chronic mucocutaneous ulcerations and autoimmune hematological disorders such as immune thrombocytopenia (ITP) and neutropenia. However, patients with RELA DN mutations additionally presented periodic fever, IBD juvenile idiopathic arthritis (JIA), and skin involvement 
DiseaseGenetic defectInheritanceOMIMAffected cellsFunctional defectAssociated features
2. Defects affecting the inflammasome 
Familial Mediterranean fever (FMF) MEFV** AR LOF 249100 Mature granulocytes, cytokine-activated monocytes Increased pyrin inflammasome-mediated induction of IL-1β Recurrent fever, serositis, and inflammation responsive to colchicine. Predisposes to vasculitis and inflammatory bowel disease, SAA amyloidosis 
AD 134610 Mature granulocytes, cytokine-activated monocytes Usually, M694del variant. Other missense variants in the B-Box and CC domains cause constitutive pyrin activation 
Pyogenic sterile arthritis, pyoderma gangrenosum, acne (PAPA) syndrome, hyperzincemia, and hypercalprotectinemia PSTPIP1 AD 604416 PMNs, monocytes Activation of the pyrin inflammasome; high production of IL-1 and IL-18 cytokines; interferon signature Destructive arthritis, inflammatory skin rash, myositis 
Mevalonate kinase deficiency (hyper-IgD syndrome/HIDS) MVK AR 260920 Somatic and hematopoietic Defect in production of isoprenoids, which are synthesized via mevalonate pathway and play a role in regulation of many signaling pathways Periodic fever and leukocytosis with usually high IgD levels 
PMVK deficiency PMVK AR NA Leukocytes Similar to MVK deficiency, increased IL-1β Recurrent fever episodes, arthritis, and cytopenia 
Muckle–Wells syndrome NLRP3** AD GOF 191900 PMNs Monocytes Activation of cryopyrin inflammasome results in increased production of IL-1/IL-18 cytokines and cell death via pyroptosis Urticaria, SNHL, SAA amyloidosis 
Familial cold autoinflammatory syndrome 1 AD GOF 120100 PMNs, monocytes Nonpruritic urticaria, arthritis, chills, fever, and leukocytosis after cold exposure 
Neonatal-onset multisystem inflammatory disease (NOMID)/chronic infantile neurological cutaneous and articular syndrome (CINCA) AD GOF 607115 PMNs, chondrocytes Neonatal-onset rash, chronic meningitis, and arthropathy with fever and inflammation 
Keratitis fugax hereditaria associated with c.61G>C
NLRP3 
AD GOF 606416  Episodic conjunctival injection, ocular pain, photophobia, foreign body sensation, and excessive tearing during acute attacks. Corneal opacities during attacks 
Familial cold autoinflammatory syndrome 2 NLRP12 AD GOF 611762 PMNs, monocytes Nonpruritic urticaria, arthritis, chills, fever, and leukocytosis after cold exposure 
NLRC4-MAS (macrophage-activating syndrome) NLRC4 AD GOF 616050 PMNs, monocytes, macrophages, intestinal epithelial cells GOF mutation in NLRC4 results in elevated secretion of IL-1β and IL-18, as well as macrophage activation Severe enterocolitis and macrophage activation syndrome 
Familial cold autoinflammatory syndrome 4 616115 
APLAID or autoinflammation, antibody deficiency, and immune dysregulation Missense variants
PLCG2
Small intragenic deletions 
AD GOF/LOF 614878 B cells, NK, mast cells Mutations affect the autoinhibitory domains and activate NF-κB and MAPK pathways Cold urticaria hypogammaglobulinemia, impaired humoral immunity, autoantibodies, autoinflammation, granulomas 
PLAID or familial cold autoinflammatory syndrome 3 614468 
Autoinflammation with arthritis and dyskeratosis (AIADK; NLRP1 deficiency) NLRP1 AR 617388 Keratinocytes and leukocytes Systemic elevation of IL-18, IL-1β, caspase 1, suggesting activation of NLRP1 inflammasome Dyskeratosis, autoimmunity, and arthritis 
NLRP1 GOF NLRP1 AD GOF 615225 Keratinocytes Spontaneous production of IL-1β and IL-18 cytokines in keratinocytes Palmoplantar carcinoma, corneal scarring; recurrent respiratory papillomatosis 
Autoinflammation with episodic fever and lymphadenopathy/cleavage-resistant RIPK1-induced autoinflammatory syndrome/CRIA RIPK1 AD 618852 Leukocytes and fibroblasts TNF-induced cell death via apoptosis and necroptosis Long-lasting fever episodes, lymphadenopathy, splenohepatomegaly, ulcers, arthralgia, GI features 
Chronic recurrent multifocal osteomyelitis and congenital dyserythropoietic anemia (Majeed syndrome) LPIN2 AR 609628 Neutrophils, bone marrow cells Dysregulation in cholesterol synthesis impairs the negative regulation of NLRP3 in macrophages resulting in high production of IL-1 Chronic recurrent multifocal osteomyelitis, transfusion-dependent anemia, cutaneous inflammatory disorders 
3. Non-inflammasome–related conditions 
TNF receptor–associated periodic syndrome (TRAPS) TNFRSF1A** AD 142680 PMNs, monocytes Mutations in the extracellular domain of 55-kD TNF receptor cause protein misfolding and intracellular receptor retention resulting in upregulation of ER stress Recurrent fever, serositis, rash, and ocular or joint inflammation 
Blau syndrome NOD2** AD 186580 Monocytes, intestinal epithelial cells Mutations in nucleotide binding site of CARD15 result in constitutive activation of NOD2 noadosome nd upregulation of NF-κB signaling Uveitis, granulomatous synovitis, camptodactyly, rash, and cranial neuropathies; 30% develop Crohn’s colitis 
ADAM17 deficiency ADAM17 AR 614328 Leukocytes and epithelial cells Defective TNF-α production Early-onset diarrhea and skin lesions 
DIRA (deficiency of the interleukin-1 receptor antagonist) IL1RN AR 612852 PMNs, monocytes Mutations in the IL-1 receptor antagonist allow unopposed action of IL-1α and IL-1β Neonatal onset of sterile multifocal osteomyelitis, periostitis, and pustulosis 
Loss of IL-1R1 sensitivity to IL-Ra (LIRSA/CRMO3) (1 patient) IL-1R1 AD 259680 T cells and B cells Activated myeloid cells. Loss of IL-1R1 biding to endogenous IL-Ra Arthritis, osteolytic/sclerotic bone lesions, poor growth, no rash, no fever 
DITRA (deficiency of IL-36 receptor antagonist) IL36RN AR 614204 Keratinocytes, leukocytes Mutations in the IL-36 receptor antagonist allow unopposed action of IL-1α and IL-1β Pustular psoriasis 
Histiocytosis-lymphadenopathy plus syndrome/H syndrome (ENT3) SLC29A3 AR 602782 Leukocytes, histiocytes Defect in nucleoside transport functions of hENT3 leads to histiocytic infiltration of numerous organs Hyperpigmentation hypertrichosis, hepatosplenomegaly, heart anomalies, hearing loss, hypogonadism, low height, and occasionally hyperglycemia/diabetes mellitus 
CAMPS (CARD14-mediated psoriasis) CARD14 AD 602723 Mainly in keratinocytes Mutations in CARD14 activate the NF-κB pathway and production of IL-8 Psoriasis 
Cherubism SH3BP2 AD 118400 Stromal cells, bone cells Hyperactivated macrophages and osteoclasts, increased NF-κB signaling Bone degeneration in jaws 
PRAAS-CANDLE (chronic atypical neutrophilic dermatitis with lipodystrophy) PSMB8* AR and AD 256040 Keratinocytes, B-cell adipose cells Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature Contractures, panniculitis, ICC, fevers 
PSMG2 AR 609702 Lymphocytes Panniculitis, lipodystrophy, autoimmune hemolytic anemia 
PSMB10 AR 619175 Lymphocytes Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature Periorbital and hand–foot annular rash (neutrophilic dermatosis), microcytic anemia, long slender fingers, hepatomegaly and splenomegaly 
PSMB9 AR or digenic or DN 617591 Lymphocytes Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature  
PSMB4 AR or digenic 617591 Lymphocytes Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature Panniculitis, lipodystrophy, autoimmune hemolytic anemia 
PRAID POMP AD 618048 Lymphocytes Increased accumulation of ubiquitinated proteins and ER stress with increased IFN signaling mediated by increased PKR signaling CANDLE (chronic atypical neutrophilic dermatitis with lipodystrophy)/ interstitial lung disease in one patient, liver disease in one patient. Recurrent and opportunistic infections. Low CD8 T cells, skewing toward naïve T cells. Low B cells and positive autoantibodies 
PSMB9 deficiency (G156D) PSMB9 AD (DN LOF) 617591 Leukocytes (mild pancytopenia) Decreased protein expression and reduced proteasome activities. Elevated levels of inflammatory cytokines (IL-6, IL-18, IP-10, IFN-α), liver enzymes in blood and CSF (IFN-α), hyperactivation of IFN-α, pSTAT1 Severe autoinflammatory phenotype (neonatal-onset fever, skin rash, myositis, severe pulmonary hypertension, basal ganglia calcification), periodic inflammatory exacerbation; immunodeficiency. Partial phenocopy of PRAAS 
Autoinflammation with neurodevelopmental disease PSMD12 AR 617516 CNS, lymphocytes ↑ peripheral blood type I IFN gene signature has been reported for some patients Intellectual disability, developmental delay, urticarial skin rash, elevated interferon signature 
COPA syndrome COPA AD 601924 PMN and tissue-specific cells Defective intracellular transport via the coat protein complex I (COPI). Exacerbated STING-mediated type I interferon response Autoimmune inflammatory arthritis and interstitial lung disease with Th17 dysregulation and autoantibody production 
Otulipenia/ORAS OTULIN AR/AD 615712 Leukocytes, fibroblasts Increase LUBAC induction of NF-κB and interferon activation leading to high proinflammatory cytokine levels. Increase in TNF-induced cell death Fever, diarrhea, skin abscesses, panniculitis 
Dominant negative OTULIN-related autoinflammatory syndrome (3 patients) OTULIN AD 615712 Lymphocytes and fibroblasts Decreased catalytic activity, accumulation of linear ubiquitin chains, increased TNF-induced cell death, Spontaneous systemic inflammation 
OTULIN haploinsufficiency OTULIN AD 615712 Epithelial cells Increased activity of caveolin-1 stabilizes ADAM10 receptor for S. aureus toxin Susceptibility to S. aureus infections in epithelial cells 
Haploinsufficiency of A20/HA20 TNFAIP3 AD 616744 Lymphocytes Defective inhibition of NF-κB signaling pathway Arthralgia, mucosal ulcers, ocular inflammation 
AP1S3 deficiency AP1S3 AR 615781 Keratinocytes Disrupted TLR3 translocation Pustular psoriasis 
ALPI deficiency ALPI AR 171740 Intestinal epithelial cells Deficient inhibition of LPS in intestine Inflammatory bowel disease 
TRIM22 TRIM22 AR 606559 Macrophages, intestinal epithelial cells Granulomatous colitis Inflammatory bowel disease 
T-cell lymphoma subcutaneous panniculitis-like (TIM3 deficiency) HAVCR2 AR 618398 Leukocytes Increased inflammasome activity due to defective checkpoint signaling Panniculitis, HLH, polyclonal cutaneous T-cell infiltrates or T-cell lymphoma 
C2orf69 deficiency (28 patients) C2orf69 AR 619423 Outer mitochondrial membrane of all cells C2orf69 regulates mitochondrial function; protein deficiency causes respiratory chain defects Early-onset severe autoinflammation disorder, often fatal. Global developmental delay, with recurrent seizures, muscle weakness due to glycogen deposits 
SYK GOF SYK AD GOF 619381 Lymphocytes, osteoclasts Increased SYK phosphorylation, enhanced NF-κB, JNK, and ERK signaling. Mutated T cells are hypersensitive to stimulation and produce various proinflammatory chemokines and cytokines (IL-17, IL-22, TNF, IFN-γ) Recurrent infections, multi-organ inflammation/inflammatory disease (gut, skin, CNS, lung, liver), B-cell lymphoma reported in 2 pts 
HCK GOF HCK AD GOF 620296 Lymphocytes Increased kinase activity of HCK mutant in vitro; ↑ production of inflammatory cytokines (IL-1β, IL-6, IL-8, TNF-α), ROS Cutaneous vasculitis, inflammatory leukocyte infiltration of the lungs (pulmonary fibrosis) and skin, anemia, hepatosplenomegaly 
NEMO exon 5 deletion IKBKG XL 301081 Leukocytes Mutant NEMO lacks exon 5 (NEMO-Dex5), fails to bind TBK1; NEMO-Dex5 stabilized IKKi, strong NF-κB, and interferon gene expression signatures Fever, skin rash, systemic autoinflammation, infections, CNS involvement, panniculitis, uveitis, hepatosplenomegaly, ectodermal dysplasia 
TBK1 deficiency TBK1 AR 620880 Leukocytes Autoinflammation driven by TNF-induced RIPK1-dependent cell death Chronic systemic autoinflammation (polyarthritis, vasculitis, rash); delayed neurocognitive development 
Retinal dystrophy, optic nerve edema, splenomegaly, anhidrosis, and headache (ROSAH) ALPK1 AD 614979 Lymphocytes Immune activation with increased NF-κB signaling, STAT1 phosphorylation, and interferon gene expression signature Retinal dystrophy, optic nerve edema, splenomegaly, anhidrosis, and migraine headache, fever, arthritis, colitis, dental abnormalities 
LYN GOF
Systemic autoinflammatory disease with vasculitis, SAIDV 
LYN AD GOF 620376 Endothelial cells and neutrophils Activated endothelial cells, constitutively active neutrophils Diffuse purpuric rash/atopic dermatitis, fever, hepatosplenomegaly, liver fibrosis/calcifications, arthritis, periorbital edema, respiratory insufficiency, colitis, poor growth 
SHARPIN deficiency SHARPIN AR 620795 Impaired development of germinal centers in secondary lymphoid organs, low CD20+cells, increased memory B cells Defect in LUBAC function, attenuated canonical NF-κB responses, increased TNF-induced cell death Arthritis, fever, colitis, amylopectinosis 
Disabling pansclerotic morphea of childhood STAT4 AD GOF 620443 Low CD4 T cells Unstimulated fibroblasts produce high levels of IL-6 Skin sclerosis, poor wound healing, joint contractures, mucosal ulcerations 
DiseaseGenetic defectInheritanceOMIMAffected cellsFunctional defectAssociated features
2. Defects affecting the inflammasome 
Familial Mediterranean fever (FMF) MEFV** AR LOF 249100 Mature granulocytes, cytokine-activated monocytes Increased pyrin inflammasome-mediated induction of IL-1β Recurrent fever, serositis, and inflammation responsive to colchicine. Predisposes to vasculitis and inflammatory bowel disease, SAA amyloidosis 
AD 134610 Mature granulocytes, cytokine-activated monocytes Usually, M694del variant. Other missense variants in the B-Box and CC domains cause constitutive pyrin activation 
Pyogenic sterile arthritis, pyoderma gangrenosum, acne (PAPA) syndrome, hyperzincemia, and hypercalprotectinemia PSTPIP1 AD 604416 PMNs, monocytes Activation of the pyrin inflammasome; high production of IL-1 and IL-18 cytokines; interferon signature Destructive arthritis, inflammatory skin rash, myositis 
Mevalonate kinase deficiency (hyper-IgD syndrome/HIDS) MVK AR 260920 Somatic and hematopoietic Defect in production of isoprenoids, which are synthesized via mevalonate pathway and play a role in regulation of many signaling pathways Periodic fever and leukocytosis with usually high IgD levels 
PMVK deficiency PMVK AR NA Leukocytes Similar to MVK deficiency, increased IL-1β Recurrent fever episodes, arthritis, and cytopenia 
Muckle–Wells syndrome NLRP3** AD GOF 191900 PMNs Monocytes Activation of cryopyrin inflammasome results in increased production of IL-1/IL-18 cytokines and cell death via pyroptosis Urticaria, SNHL, SAA amyloidosis 
Familial cold autoinflammatory syndrome 1 AD GOF 120100 PMNs, monocytes Nonpruritic urticaria, arthritis, chills, fever, and leukocytosis after cold exposure 
Neonatal-onset multisystem inflammatory disease (NOMID)/chronic infantile neurological cutaneous and articular syndrome (CINCA) AD GOF 607115 PMNs, chondrocytes Neonatal-onset rash, chronic meningitis, and arthropathy with fever and inflammation 
Keratitis fugax hereditaria associated with c.61G>C
NLRP3 
AD GOF 606416  Episodic conjunctival injection, ocular pain, photophobia, foreign body sensation, and excessive tearing during acute attacks. Corneal opacities during attacks 
Familial cold autoinflammatory syndrome 2 NLRP12 AD GOF 611762 PMNs, monocytes Nonpruritic urticaria, arthritis, chills, fever, and leukocytosis after cold exposure 
NLRC4-MAS (macrophage-activating syndrome) NLRC4 AD GOF 616050 PMNs, monocytes, macrophages, intestinal epithelial cells GOF mutation in NLRC4 results in elevated secretion of IL-1β and IL-18, as well as macrophage activation Severe enterocolitis and macrophage activation syndrome 
Familial cold autoinflammatory syndrome 4 616115 
APLAID or autoinflammation, antibody deficiency, and immune dysregulation Missense variants
PLCG2
Small intragenic deletions 
AD GOF/LOF 614878 B cells, NK, mast cells Mutations affect the autoinhibitory domains and activate NF-κB and MAPK pathways Cold urticaria hypogammaglobulinemia, impaired humoral immunity, autoantibodies, autoinflammation, granulomas 
PLAID or familial cold autoinflammatory syndrome 3 614468 
Autoinflammation with arthritis and dyskeratosis (AIADK; NLRP1 deficiency) NLRP1 AR 617388 Keratinocytes and leukocytes Systemic elevation of IL-18, IL-1β, caspase 1, suggesting activation of NLRP1 inflammasome Dyskeratosis, autoimmunity, and arthritis 
NLRP1 GOF NLRP1 AD GOF 615225 Keratinocytes Spontaneous production of IL-1β and IL-18 cytokines in keratinocytes Palmoplantar carcinoma, corneal scarring; recurrent respiratory papillomatosis 
Autoinflammation with episodic fever and lymphadenopathy/cleavage-resistant RIPK1-induced autoinflammatory syndrome/CRIA RIPK1 AD 618852 Leukocytes and fibroblasts TNF-induced cell death via apoptosis and necroptosis Long-lasting fever episodes, lymphadenopathy, splenohepatomegaly, ulcers, arthralgia, GI features 
Chronic recurrent multifocal osteomyelitis and congenital dyserythropoietic anemia (Majeed syndrome) LPIN2 AR 609628 Neutrophils, bone marrow cells Dysregulation in cholesterol synthesis impairs the negative regulation of NLRP3 in macrophages resulting in high production of IL-1 Chronic recurrent multifocal osteomyelitis, transfusion-dependent anemia, cutaneous inflammatory disorders 
3. Non-inflammasome–related conditions 
TNF receptor–associated periodic syndrome (TRAPS) TNFRSF1A** AD 142680 PMNs, monocytes Mutations in the extracellular domain of 55-kD TNF receptor cause protein misfolding and intracellular receptor retention resulting in upregulation of ER stress Recurrent fever, serositis, rash, and ocular or joint inflammation 
Blau syndrome NOD2** AD 186580 Monocytes, intestinal epithelial cells Mutations in nucleotide binding site of CARD15 result in constitutive activation of NOD2 noadosome nd upregulation of NF-κB signaling Uveitis, granulomatous synovitis, camptodactyly, rash, and cranial neuropathies; 30% develop Crohn’s colitis 
ADAM17 deficiency ADAM17 AR 614328 Leukocytes and epithelial cells Defective TNF-α production Early-onset diarrhea and skin lesions 
DIRA (deficiency of the interleukin-1 receptor antagonist) IL1RN AR 612852 PMNs, monocytes Mutations in the IL-1 receptor antagonist allow unopposed action of IL-1α and IL-1β Neonatal onset of sterile multifocal osteomyelitis, periostitis, and pustulosis 
Loss of IL-1R1 sensitivity to IL-Ra (LIRSA/CRMO3) (1 patient) IL-1R1 AD 259680 T cells and B cells Activated myeloid cells. Loss of IL-1R1 biding to endogenous IL-Ra Arthritis, osteolytic/sclerotic bone lesions, poor growth, no rash, no fever 
DITRA (deficiency of IL-36 receptor antagonist) IL36RN AR 614204 Keratinocytes, leukocytes Mutations in the IL-36 receptor antagonist allow unopposed action of IL-1α and IL-1β Pustular psoriasis 
Histiocytosis-lymphadenopathy plus syndrome/H syndrome (ENT3) SLC29A3 AR 602782 Leukocytes, histiocytes Defect in nucleoside transport functions of hENT3 leads to histiocytic infiltration of numerous organs Hyperpigmentation hypertrichosis, hepatosplenomegaly, heart anomalies, hearing loss, hypogonadism, low height, and occasionally hyperglycemia/diabetes mellitus 
CAMPS (CARD14-mediated psoriasis) CARD14 AD 602723 Mainly in keratinocytes Mutations in CARD14 activate the NF-κB pathway and production of IL-8 Psoriasis 
Cherubism SH3BP2 AD 118400 Stromal cells, bone cells Hyperactivated macrophages and osteoclasts, increased NF-κB signaling Bone degeneration in jaws 
PRAAS-CANDLE (chronic atypical neutrophilic dermatitis with lipodystrophy) PSMB8* AR and AD 256040 Keratinocytes, B-cell adipose cells Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature Contractures, panniculitis, ICC, fevers 
PSMG2 AR 609702 Lymphocytes Panniculitis, lipodystrophy, autoimmune hemolytic anemia 
PSMB10 AR 619175 Lymphocytes Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature Periorbital and hand–foot annular rash (neutrophilic dermatosis), microcytic anemia, long slender fingers, hepatomegaly and splenomegaly 
PSMB9 AR or digenic or DN 617591 Lymphocytes Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature  
PSMB4 AR or digenic 617591 Lymphocytes Proteasome dysfunction with accumulation of ubiquitinated proteins and ER stress. Increased interferon signature Panniculitis, lipodystrophy, autoimmune hemolytic anemia 
PRAID POMP AD 618048 Lymphocytes Increased accumulation of ubiquitinated proteins and ER stress with increased IFN signaling mediated by increased PKR signaling CANDLE (chronic atypical neutrophilic dermatitis with lipodystrophy)/ interstitial lung disease in one patient, liver disease in one patient. Recurrent and opportunistic infections. Low CD8 T cells, skewing toward naïve T cells. Low B cells and positive autoantibodies 
PSMB9 deficiency (G156D) PSMB9 AD (DN LOF) 617591 Leukocytes (mild pancytopenia) Decreased protein expression and reduced proteasome activities. Elevated levels of inflammatory cytokines (IL-6, IL-18, IP-10, IFN-α), liver enzymes in blood and CSF (IFN-α), hyperactivation of IFN-α, pSTAT1 Severe autoinflammatory phenotype (neonatal-onset fever, skin rash, myositis, severe pulmonary hypertension, basal ganglia calcification), periodic inflammatory exacerbation; immunodeficiency. Partial phenocopy of PRAAS 
Autoinflammation with neurodevelopmental disease PSMD12 AR 617516 CNS, lymphocytes ↑ peripheral blood type I IFN gene signature has been reported for some patients Intellectual disability, developmental delay, urticarial skin rash, elevated interferon signature 
COPA syndrome COPA AD 601924 PMN and tissue-specific cells Defective intracellular transport via the coat protein complex I (COPI). Exacerbated STING-mediated type I interferon response Autoimmune inflammatory arthritis and interstitial lung disease with Th17 dysregulation and autoantibody production 
Otulipenia/ORAS OTULIN AR/AD 615712 Leukocytes, fibroblasts Increase LUBAC induction of NF-κB and interferon activation leading to high proinflammatory cytokine levels. Increase in TNF-induced cell death Fever, diarrhea, skin abscesses, panniculitis 
Dominant negative OTULIN-related autoinflammatory syndrome (3 patients) OTULIN AD 615712 Lymphocytes and fibroblasts Decreased catalytic activity, accumulation of linear ubiquitin chains, increased TNF-induced cell death, Spontaneous systemic inflammation 
OTULIN haploinsufficiency OTULIN AD 615712 Epithelial cells Increased activity of caveolin-1 stabilizes ADAM10 receptor for S. aureus toxin Susceptibility to S. aureus infections in epithelial cells 
Haploinsufficiency of A20/HA20 TNFAIP3 AD 616744 Lymphocytes Defective inhibition of NF-κB signaling pathway Arthralgia, mucosal ulcers, ocular inflammation 
AP1S3 deficiency AP1S3 AR 615781 Keratinocytes Disrupted TLR3 translocation Pustular psoriasis 
ALPI deficiency ALPI AR 171740 Intestinal epithelial cells Deficient inhibition of LPS in intestine Inflammatory bowel disease 
TRIM22 TRIM22 AR 606559 Macrophages, intestinal epithelial cells Granulomatous colitis Inflammatory bowel disease 
T-cell lymphoma subcutaneous panniculitis-like (TIM3 deficiency) HAVCR2 AR 618398 Leukocytes Increased inflammasome activity due to defective checkpoint signaling Panniculitis, HLH, polyclonal cutaneous T-cell infiltrates or T-cell lymphoma 
C2orf69 deficiency (28 patients) C2orf69 AR 619423 Outer mitochondrial membrane of all cells C2orf69 regulates mitochondrial function; protein deficiency causes respiratory chain defects Early-onset severe autoinflammation disorder, often fatal. Global developmental delay, with recurrent seizures, muscle weakness due to glycogen deposits 
SYK GOF SYK AD GOF 619381 Lymphocytes, osteoclasts Increased SYK phosphorylation, enhanced NF-κB, JNK, and ERK signaling. Mutated T cells are hypersensitive to stimulation and produce various proinflammatory chemokines and cytokines (IL-17, IL-22, TNF, IFN-γ) Recurrent infections, multi-organ inflammation/inflammatory disease (gut, skin, CNS, lung, liver), B-cell lymphoma reported in 2 pts 
HCK GOF HCK AD GOF 620296 Lymphocytes Increased kinase activity of HCK mutant in vitro; ↑ production of inflammatory cytokines (IL-1β, IL-6, IL-8, TNF-α), ROS Cutaneous vasculitis, inflammatory leukocyte infiltration of the lungs (pulmonary fibrosis) and skin, anemia, hepatosplenomegaly 
NEMO exon 5 deletion IKBKG XL 301081 Leukocytes Mutant NEMO lacks exon 5 (NEMO-Dex5), fails to bind TBK1; NEMO-Dex5 stabilized IKKi, strong NF-κB, and interferon gene expression signatures Fever, skin rash, systemic autoinflammation, infections, CNS involvement, panniculitis, uveitis, hepatosplenomegaly, ectodermal dysplasia 
TBK1 deficiency TBK1 AR 620880 Leukocytes Autoinflammation driven by TNF-induced RIPK1-dependent cell death Chronic systemic autoinflammation (polyarthritis, vasculitis, rash); delayed neurocognitive development 
Retinal dystrophy, optic nerve edema, splenomegaly, anhidrosis, and headache (ROSAH) ALPK1 AD 614979 Lymphocytes Immune activation with increased NF-κB signaling, STAT1 phosphorylation, and interferon gene expression signature Retinal dystrophy, optic nerve edema, splenomegaly, anhidrosis, and migraine headache, fever, arthritis, colitis, dental abnormalities 
LYN GOF
Systemic autoinflammatory disease with vasculitis, SAIDV 
LYN AD GOF 620376 Endothelial cells and neutrophils Activated endothelial cells, constitutively active neutrophils Diffuse purpuric rash/atopic dermatitis, fever, hepatosplenomegaly, liver fibrosis/calcifications, arthritis, periorbital edema, respiratory insufficiency, colitis, poor growth 
SHARPIN deficiency SHARPIN AR 620795 Impaired development of germinal centers in secondary lymphoid organs, low CD20+cells, increased memory B cells Defect in LUBAC function, attenuated canonical NF-κB responses, increased TNF-induced cell death Arthritis, fever, colitis, amylopectinosis 
Disabling pansclerotic morphea of childhood STAT4 AD GOF 620443 Low CD4 T cells Unstimulated fibroblasts produce high levels of IL-6 Skin sclerosis, poor wound healing, joint contractures, mucosal ulcerations 

IFN, interferon; HSM, hepatosplenomegaly; CSF, cerebrospinal fluid; SLE, systemic lupus erythematosus; TORCH, toxoplasmosis, other, rubella, cytomegalovirus, and herpes infections; SNHL, sensorineural hearing loss; AGS, Aicardi-Goutières syndrome; BSN, bilateral striatal necrosis; FCL, familial chilblain lupus; ICC, intracranial calcification; IFN, interferon type I; pDCs, plasmacytoid dendritic cells; SP, spastic paraparesis; SMS, Singleton–Merten syndrome; ss, single-stranded; ADA, adenosine deaminase; CNS, central nervous system; IBD, inflammatory bowel disease; autoAbs, autoantibodies.

* variants in PSMB4, PSMB9, PSMA3, and POMP have been proposed to cause a similar CANDLE phenotype in compound heterozygous monogenic (PSMB4), digenic (PSMA3/PSMB8, PSMB9/PSMB4, PSMB4/PSMB8), and AD monogenic (POMP) models (119). Only G156D mutation in PSMB9 has been shown to cause an autoinflammatory phenotype with immunodeficiency in patients and mouse model (120).

Total number of disorders in Table 7: 69.

New IEIs: 11, STAT4 GOF, PMVK, ALPK1, LYN GOF, SHARPIN, LSM11, RNU71, ARF1, OTULIN (two new entries), and RELA (84, 85, 86, 87, 88, 89, 90, 91, 92, 93, 94).

** depicts that somatic mutations mimicking the germline disorder have been described for this gene.

a

RELA previously described as causing combined immunodeficiency a second entry included here as DN mutations are associated with an inflammatory phenotype with different mechanism of disease. OTULIN is repeated three times as different mechanisms of disease give rise to different phenotypes. NLRP1 is also repeated twice as AR and AD forms result in different phenotypes.

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