Complexity and Controversies over the Cytokine Profiles of T Helper Cell Subpopulations in Tuberculosis

Table 1

T helper cell (Th) subtypes and Th-related soluble mediators in human and experimental tuberculosis.

	Species	Putative role in tuberculosis	Reference
T helper subtype
Th1	Mouse and human	the growth and dispersion of M. tuberculosis	[7–11]
Th2	Human	in BALF associated with clinical progression	[12]
		Th2 response in active disease	[13, 14]
	Mouse	progression and reactivation of TB	[15]
T regulatory	Human	Treg cells in more severe active disease	[16]
		reactivation of latent TB	[17]
		TB-MDR	[18]
	Mouse	Treg cells in the early days of infection protects from severe disease	[19]
Th17	Human	Th17 cells in active TB	[20, 21]
	Mouse	neutrophil accumulation and tissue damage	[22, 23]
		recruitment of IFN- producing cells	[24]
		Induce granuloma formation and remain as long-lived memory cells	[25–27]
Th22	Human	in healthy M. tuberculosis-exposed individuals	[20]
		in PBMC culture from patients with active TB	[28]
		in pleural fluid from patients with active TB	[29–31]
Th9	Human	Th9 cells in tuberculous pleural effusion	[32]
T helper-related soluble mediator
IFN-	Human	↑ Mycobacterium-specific production after clinical cure	[33]
		in active TB patients at the site of infection	[34–37]
	Mouse	influx of neutrophils and neutrophil-associated tissue damage	[38]
		iNOS in infected macrophages	[39, 40]
	Human and mouse	autophagy	[41]
NO	Mouse	killing and growth inhibiting of virulent M. tuberculosis	[39]
TNF-	Human	↑ Mycobacterium-specific production after clinical cure	[33, 42–44]
	Mouse and human	Maintenance of the granuloma integrity	[45–48]
IL-4	Human	in the blood and BALF in TB patients with severe forms	[12, 49–53]
	Mouse	progression and reactivation of TB	[15, 54]
		Without any influence in disease susceptibility	[55, 56]
	Mouse and human	autophagic control of intracellular Mycobacterium tuberculosis	[57]
IL-10	Human	↑ Mycobacterium-specific production after clinical cure	[33]
		in active disease	[14, 58, 59]
		No difference between active-TB and clinically cured individuals	[60]
	Mouse	long-term lack of control of inflammatory responses and progression of the disease	[61]
IL-17	Mouse	Th1 induction following BCG vaccination	[62]
IL-9	Human	in patients with pulmonary TB	[63]
		IFN- expression by PBMCs in latent TB	[64]
IL-22	Human	in pleural fluid from patients with active TB	[29–31]
	Mouse	IL-22 deficiency does not alter the outcome of infection	[32, 65]