Table 4.
Comparison of ICOSL, ICOS, and NIK deficiency in mouse and human
FeaturesMiceHumans
ICOSLG deficiencyICOS deficiencyNIK (MAP3K14) deficiencyICOSLG deficiencyICOS deficiencyNIK (MAP3K14) deficiency
Infections    Recurrent respiratory tract infections Recurrent respiratory tract infections Recurrent respiratory tract infections 
    Intestinal infections (CampylobacterIntestinal infections (Campylobacter, Salmonella norovirus, adenovirus, CryptosporidiumIntestinal infections (Cryptosporidium; CMV) 
     Skin abscesses Osteomyelitis with BCG 
    Herpesviridae-related infections Herpesviridae-related infections; recurrent HSV (labialis, keratitis, genital), CMV, HHV6  
    HPV HPV: warts, cancer  
     Pneumocystis jiroveci pneumonia  
Other salient clinical features    Intermittent chronic diarrhea of unclear etiology (no evidence of infectious or inflammatory bowel disease) Autoimmunity: RA, IBD, interstitial pneumonitis, psoriasis Granulomatous hepatitis (possibly related to disseminated BCG) 
    Splenomegaly Splenomegaly  
    Dentigerous cyst Granulomatous skin diseases  
     Cytopenia; neutropenia; thrombocytopenia (2/15 patients each)  
     Cancer: HPV vulva; LGL-T; SCC  
Immunoglobulins    Hypogammaglobulinemia of IgG, IgA, IgM Hypogammaglobulinemia of IgG and IgA; some with low-normal IgM Hypogammaglobulinemia of IgG, IgA; IgM low/normal/elevated 
Baseline ↓ IgG1 ↓ IgG1 Normal levels of IgG, IgM    
 ↓ IgG2a, IgG2b, IgA ↓ IgE ↓ IgA    
   (Some mouse models had ↓ IgG2) Normal levels of IgG1, IgG2a    
   ↓ IgG2b, IgG3    
Response to vaccination Intact response to T cell–independent antigen (based on IgM and IgG3 response to trinitrophenol-Ficoll) Intact response to T cell–independent antigen (based on IgM and IgG3 response to trinitrophenol-Ficoll) Poor (nearly absent) response to ovalbumin immunization    
 ↓ T cell–dependent responses (based on ↓ IgG1, ↓ IgE) ↓ T cell–dependent responses (based on ↓ IgG1, ↓ IgG2a, ↓ IgE)     
Lymphocytes No effect on T or B cell development from thymus or bone marrow No effect on T or B cell development from thymus or bone marrow Thymus: no effect on T cells Circulating absolute T cell counts: decreased Circulating absolute T cell counts: normal Circulating absolute T cell counts: normal 
    Circulating CD4+ T cell subset: decreased Circulating CD4+ T cell subset: variable Circulating CD4+ T cell subset: normal 
 No effect on overall composition of mature B and T cell subsets in spleen No effect on overall composition of mature B and T cell subsets in spleen Spleen: no effect on T cells; decreased B220+ IgM+ cells Circulating CD8+ T cell subset: decreased Circulating CD8+ T cell subset: variable Circulating CD8+ T cell subset: normal 
    Memory T cell counts: decreased Memory T cell counts: variable Memory T cell counts: normal 
   Bone marrow: increased B220+ CD25+ and B220+ CD43 Circulating Tfh (CD4+ CXCR5+ CD45RA): low–normal Reduction in circulating Tfh (defined as CD4+ CXCR5+ CD45RAReduction in circulating Tfh (CD4+ CXCR5+ CD45RA
    T reg cells: normal T reg cells: normal T reg cells: normal 
    B cell counts: decreased in childhood; intermittently within reference range in adulthood B cell counts: normal when diagnosed in childhood; decreased when diagnosed in adulthood B cell counts: decreased 
    Circulating naive B cells: increased Circulating naive B cells: high (children); decreased (adults) Circulating naive B cells: high (children) 
    Circulating switched memory B cell cells: decreased Circulating switched memory B cell cells: decreased Circulating switched memory B cell cells: decreased 
      NK cell counts: decreased 
Lymphoid organs ↓ number and size of GC formation ↓ number and size of GC formation Absent lymph nodes; abnormal lymphoid architecture in thymus and spleen    
Myeloid cells Not reported Not reported ↓ CD11b+ monocytes in spleen Progressive neutropenia Neutropenia (n = 2) Not reported 
FeaturesMiceHumans
ICOSLG deficiencyICOS deficiencyNIK (MAP3K14) deficiencyICOSLG deficiencyICOS deficiencyNIK (MAP3K14) deficiency
Infections    Recurrent respiratory tract infections Recurrent respiratory tract infections Recurrent respiratory tract infections 
    Intestinal infections (CampylobacterIntestinal infections (Campylobacter, Salmonella norovirus, adenovirus, CryptosporidiumIntestinal infections (Cryptosporidium; CMV) 
     Skin abscesses Osteomyelitis with BCG 
    Herpesviridae-related infections Herpesviridae-related infections; recurrent HSV (labialis, keratitis, genital), CMV, HHV6  
    HPV HPV: warts, cancer  
     Pneumocystis jiroveci pneumonia  
Other salient clinical features    Intermittent chronic diarrhea of unclear etiology (no evidence of infectious or inflammatory bowel disease) Autoimmunity: RA, IBD, interstitial pneumonitis, psoriasis Granulomatous hepatitis (possibly related to disseminated BCG) 
    Splenomegaly Splenomegaly  
    Dentigerous cyst Granulomatous skin diseases  
     Cytopenia; neutropenia; thrombocytopenia (2/15 patients each)  
     Cancer: HPV vulva; LGL-T; SCC  
Immunoglobulins    Hypogammaglobulinemia of IgG, IgA, IgM Hypogammaglobulinemia of IgG and IgA; some with low-normal IgM Hypogammaglobulinemia of IgG, IgA; IgM low/normal/elevated 
Baseline ↓ IgG1 ↓ IgG1 Normal levels of IgG, IgM    
 ↓ IgG2a, IgG2b, IgA ↓ IgE ↓ IgA    
   (Some mouse models had ↓ IgG2) Normal levels of IgG1, IgG2a    
   ↓ IgG2b, IgG3    
Response to vaccination Intact response to T cell–independent antigen (based on IgM and IgG3 response to trinitrophenol-Ficoll) Intact response to T cell–independent antigen (based on IgM and IgG3 response to trinitrophenol-Ficoll) Poor (nearly absent) response to ovalbumin immunization    
 ↓ T cell–dependent responses (based on ↓ IgG1, ↓ IgE) ↓ T cell–dependent responses (based on ↓ IgG1, ↓ IgG2a, ↓ IgE)     
Lymphocytes No effect on T or B cell development from thymus or bone marrow No effect on T or B cell development from thymus or bone marrow Thymus: no effect on T cells Circulating absolute T cell counts: decreased Circulating absolute T cell counts: normal Circulating absolute T cell counts: normal 
    Circulating CD4+ T cell subset: decreased Circulating CD4+ T cell subset: variable Circulating CD4+ T cell subset: normal 
 No effect on overall composition of mature B and T cell subsets in spleen No effect on overall composition of mature B and T cell subsets in spleen Spleen: no effect on T cells; decreased B220+ IgM+ cells Circulating CD8+ T cell subset: decreased Circulating CD8+ T cell subset: variable Circulating CD8+ T cell subset: normal 
    Memory T cell counts: decreased Memory T cell counts: variable Memory T cell counts: normal 
   Bone marrow: increased B220+ CD25+ and B220+ CD43 Circulating Tfh (CD4+ CXCR5+ CD45RA): low–normal Reduction in circulating Tfh (defined as CD4+ CXCR5+ CD45RAReduction in circulating Tfh (CD4+ CXCR5+ CD45RA
    T reg cells: normal T reg cells: normal T reg cells: normal 
    B cell counts: decreased in childhood; intermittently within reference range in adulthood B cell counts: normal when diagnosed in childhood; decreased when diagnosed in adulthood B cell counts: decreased 
    Circulating naive B cells: increased Circulating naive B cells: high (children); decreased (adults) Circulating naive B cells: high (children) 
    Circulating switched memory B cell cells: decreased Circulating switched memory B cell cells: decreased Circulating switched memory B cell cells: decreased 
      NK cell counts: decreased 
Lymphoid organs ↓ number and size of GC formation ↓ number and size of GC formation Absent lymph nodes; abnormal lymphoid architecture in thymus and spleen    
Myeloid cells Not reported Not reported ↓ CD11b+ monocytes in spleen Progressive neutropenia Neutropenia (n = 2) Not reported 

The immunological findings in the mouse (top) and clinical features in humans (bottom) are listed. BCG, Bacillus Calmette-Guérin; RA, rheumatoid arthritis; IBD, inflammatory bowel disease; LGL-T, large granular lymphocyte T cells; SCC, squamous cell carcinoma.

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