. 2023 Feb 3;12(2):246. doi: 10.3390/pathogens12020246

Table 1.

Non-canonical host cells of Leishmania parasites.

Host Cell/Origin	Leishmania Species	Main Outcome	Detection Method	Reference
PROFESSIONAL PHAGOCYTES
Dendritic cells (DCs)
Langerhans cells from mouse skin ^❖	L. major PMs	No or low PMs uptake	LM/Diff-Quik, TEM, FL/AO+EtBr	[53,54,55]
	L. major AMs	AMs uptake and internalisation, no or weak multiplication	LM/Diff-Quik, TEM, FL/AO+EtBr, ICC	[54,56,57]
Mouse lymph node DCs ^♦○	L. major AMs	Presence of AMs	LM/G, IHC	[56,58,59]
Mouse spleen DCs ^❖	L. major PMs/AMs, L. m. mexicana PMs	PMs/AMs uptake	LM/G	[59,60]
Mouse bone marrow DCs ^❖	L. major PMs/AMs, L. mexicana PMs, L. amazonensis AMs/PMs	PMs/AMs uptake in all; multiplication reported only in L. amazonensis	LM/Diff-Quik, ICC, FC	[61,62,63,64]
	L. infantum PMs/PMs (CFSE)	PMs uptake, transformation into AMs	LM/G, FC/CFSE-PMs	[65]
Human immature monocyte-derived DCs^❖	L. amazonensis, L. braziliensis, L. infantum PMs	PMs uptake, internalisation	CLSM/Dapi	[25]
Human immature monocyte-derived DCs^❖	L. donovani PMs	PMs uptake, transformation into AMs	LM/MGG	[66]
Mast cells (MCs)
Mouse peritoneal MCs ^❖	L. tropica, L. donovani PMs (CFSE)	PMs uptake in L. tropica, but not in L. donovani	FC+CLSM/CFSE-PMs	[22,67]
Mouse bone marrow MCs ^❖	L. major, L. infantum PMs	PMs uptake, transformation into AMs, multiplication leading to cell lysis and AMs release	LM/MGG	[21]
Eosinophils
Human peripheral eosinophils ^❖	L. donovani PMs	PMs uptake and killing after 2 h p.i.	LM/Diff-Quik	[68]
	L. donovani AMs	AMs uptake, not efficient killing	LM, TEM	[69]
Rat peritoneal eosinophils ^❖	L. major PMs	PMs uptake and killing	LM/MGG, ICC	[70]
Rat peritoneal eosinophils ^♦^○^❖	L. m. amazonensis PMs/AMs	PMs/AMs uptake and killing	TEM	[71]
Mouse eosinophils in skin lesion ^♦○	L. m. mexicana AMs	AMs uptake, not efficient killing	TEM	[72]
Histiocyte-like cells				[7]
Sticker dog sarcoma 503 cells ^❖	L. donovani, L. mexicana, L. m. mexicana, L. braziliensis, L. b. pifanoi, L. t. major PMs/AMs	PMs/AMs uptake, multiplication, continuous passages	LM/G, TEM	[73,74,75,76,77,78]
	L. m. mexicana PMs	PMs uptake, transformation into AMs, multiplication after day 3 p.i., transformation into PMs	LM/G, TEM	[43]
	L. adleri, L. hoogstraali, L. agamae PMs	Low PMs uptake, transformation into AMs	LM/G	[43]
NON-PROFESSIONAL PHAGOCYTES
Lymphocytes
Human B (Daudi) and T (HUT78) cells ^❖	L. donovani PMs/AMs	PMs/AMs uptake, PMs transformation into AMs, viability up to 2 weeks after infection with AMs	LM/G, TEM	[79]
Fibrocytes
Mouse peripheral blood fibrocytes ^❖	L. amazonensis PMs	PMs uptake, transformation into AMs, low multiplication, clearance by 72 h p.i.	LM/G, FL/Dapi, TEM, SEM	[80]
Fibroblasts
Canine skin fibroblasts ^♦□	Leishmania sp.	Presence of AMs	TEM, LM/HE, G, PAS	[81]
	L. donovani	Presence of AMs	IHC	[82]
Human skin fibroblasts ^♦□	L. tropica	Presence of AMs	LM/G, TEM	[83,84]
	Leishmania sp. (cutaneous)	Presence of AMs	TEM	[85]
Human skin fibroblasts ^❖	L. amazonensis PMs	PMs uptake, transformation into AMs, multiplication	TEM	[14]
	L. m. amazonensis AMs	AMs uptake, multiplication, killing of AMs by day 8 p.i.	LM/G, TEM, ICC	[12]
	Leishmania sp. (mucocutaneous), L. donovani PMs	PMs uptake in Leishmania sp. (not in L. donovani), transformation into AMs, no or low multiplication, decline during a 3-week period p.i.	LM, TEM, SEM	[86]
Human foreskin fibroblasts ^❖	L. donovani PMs	PMs uptake, transformation into AMs, no multiplication, viability up to day 14 p.i.	LM, TEM, SEM	[87]
Human foreskin fibroblasts ^❖	L. major PMs (SPIONs)	PMs uptake	LM/SPIONs-PMs+Prussian blue, TEM	[88]
	L. major PMs (AO, Dil)	PMs uptake	FL/AO-PMs, Dil-PMs	[89]
Mouse skin fibroblasts ^♦○	L. amazonensis PMs	AMs presence	LM/HE, Lennert’s G	[90]
Mouse skin fibroblasts ^❖	L. major PMs/AMs	PMs/AMs uptake	ICC	[9]
	L. amazonensis PMs	PMs uptake and killing of PMs after day 3 p.i.	LM/G, TEM, FL/Dapi	[91]
	L. infantum, L. mexicana PMs	PMs uptake, transformation into AMs, multiplication	LM/G, TEM	[10]
Hamster skin fibroblasts ^❖	L. infantum, L. mexicana PMs	PMs uptake, transformation into AMs, multiplication	LM/G, TEM	[10]
Rat skin fibroblasts ^❖	L. infantum, L. mexicana PMs	PMs uptake, transformation into AMs, no multiplication	LM/G, TEM	[10]
Human fibroblasts in lymph node ^♦□	Leishmania sp.	AMs presence	LM/G	[92]
Mouse fibroblasts in lymph node ^❖	L. major PMs/AMs	PMs/AMs uptake	TEM, ICC	[9]
Draining lymph nodes of healed mice (presumably fibroblasts) ^♦○	L. major PMs	Presence of AMs, parasite survival or limited killing	IHC	[9]
Mouse embryonic fibroblasts ^❖	L. donovani PMs	PMs uptake, transformation into AMs, efficient host defence via IFN-inducible guanylate binding proteins	LM/G, CLSM/Dapi	[93]
	L. amazonensis PMs (RFP)	PMs uptake, transformation into AMs, multiplication	LM/HE, TEM, CLSM+FC/RFP-PMs	[8]
	L. major PMs	PMs uptake	CLSM/Dapi	[94]
Mouse tumour fibroblasts (L cells) ^❖	L. amazonensis, L. major AMs (GFP)	AMs uptake, internalisation (low in L. major), multiplication (not in L. major)	CLSM/GFP-AMs	[15]
Fibroblasts from embryonic chick brain ^❖	L. donovani AMs	AMs uptake, viability up to day 17 p.i., transformation into PMs	LM/G	[11]
Fibroblast-like cells from embryonic chick muscle ^❖	L. donovani/presumably AMs	AMs uptake, no multiplication, degeneration after day 20 p.i.	LM/HE	[95]
Mouse perineurial cells ^♦○	L. amazonensis PMs	Presence of AMs	TEM	[96]
Adipocytes
Mouse brown and white adipose tissue ^♦○	L. infantum PMs	PMs uptake, viable AMs present for up to 40 weeks p.i.	IHC, qPCR	[46]
Mouse adipocytes derived from primary pre-adipocytes from subcutaneous white adipose tissue ^❖	L. infantum PMs (GFP)	PMs uptake (further progress not reported)	TEM, qPCR, CLSM/GFP-AMs	[46]
Human adipocytes derived from adipose tissue primary progenitor cells ^❖	L. infantum PMs (GFP)	PMs uptake (further progress not reported)	qPCR, CLSM/GFP-AMs	[46]
Mouse adipocytes differentiated in vitro from 3T3-L1 fibroblasts ^❖	L. amazonensis, L. braziliensis PMs/AMs	PMs/AMs uptake, PMs transformation into AMs, viability up to 144 h p.i. and ability to transform into PMs	LM/G, FL/Dapi, TEM	[45]
	L. amazonensis AMs (GFP)	AMs uptake, viability up to 144 h p.i.	FL/GFP-AMs	[45]
Epithelial cells
Human epithelial cells of eccrine sweat gland (HIV patient) ^♦□	Leishmania sp., L. infantum	AMs presence	LM/HE	[97,98]
Human retinal pigmented epithelial cells (ARPE-19) ^❖	L. amazonensis PMs	PMs uptake, internalisation	LM/G, IHC, TEM	[99]
Human amnion epithelium ^❖	L. donovani, L. b. pifanoi PMs	PMs uptake, transformation into AMs, clearance by day 29–32 p.i.	LM/G	[100]
Human amnion epithelium ^❖	L. donovan PMs	PMs uptake, transformation into AMs, multiplication (not clear whether PMS or AMs)	LM/G	[101]
A549 (human adenocarcinomic alveolar basal epithelium) cells ^❖	L. donovani PMs	PMs uptake, transformation into AMs, efficient defence via IFN-inducible guanylate binding proteins	LM/G, CLSM/Dapi	[93]
HeLa (human cervix carcinoma) cells ^❖	L. t. major PMs	PMs uptake, transformation into AMs, multiplication, destruction of host cells after day 3	LM	[102]
HeLa (human cervix carcinoma) cells ^❖	L. donovani PMs	PMs uptake, transformation into AMs, decline after 5 h p.i.	LM/G	[103]
LLC-MK2 (rhesus monkey kidney epithelium) cells ^❖	L. donovani AMs	AMs uptake, multiplication	LM/G	[104]
Vero (monkey kidney) cells ^❖	L. chagasi, L. braziliensis PMs	PMs uptake, transformation into AMs, multiplication	LM/G, TEM	[105,106]
Chinese hamster ovary cells ^❖	L. amazonensis AMs	AMs uptake, multiplication	IHC, TEM	[42]
C. burnetii-infected Vero cells ^❖	L. amazonensis AMs	AMs uptake, multiplication	LM, TEM	[107]
C. burnetii-infected Chinese hamster ovary cells ^❖	L. amazonensis AMs	AMs uptake, multiplication	LM, TEM, CLSM/PI	[107,108]
Mesenchymal stem cells (MSCs)
Mouse bone marrow MSCs ^♦○❖	L. infantum PMs	PMs uptake, transformation into AMs	LM/G, ICC, FC	[109]
Human adipose tissue MSCs ^❖	L. donovani, L. infantum, L. major, L. tropica PMs	PMs uptake, transformation into AMs but AMs present only at day 1 p.i.; at day 7, 14, 21, and 28 only PMs detected	LM/G, microcapillary culture method, PCR	[16]
Myocytes
Canine skeletal/smooth muscles ^♦□	L. infantum, Leishmania sp.	Presence of AMs within myofibres	LM/HE, IHC	[110,111]
Mouse skeletal muscles ^♦○	L. amazonensis AMs	Presence of AMs within myofibres	LM/HE	[112]
Turtle heart cells ^❖	L. m. mexicana, L. adleri, L. hoogstraali PMs	PMs uptake (lower in L. adleri and L. hoogstraali), transformation into AMs (further progress not reported)	TEM (L. mexicana only), LM/G	[43]
Endothelial cells
Human endothelial cells of blood vessels ^♦□	L. donovani, Leishmania sp.	Presence of AMs	LM	[81,113,114]
Human endothelial cells of capillaries ^♦○	L. tropica PMs	Presence of AMs	LM	[115]
Human microvascular endothelial (HMEC-1) cell line ^❖	L. infantum PMs	No uptake of PMs	LM/G	[116]
Keratinocytes
Human keratinocytes (HIV patient) ^♦□	L. infantum	AMs presence	LM/HE	[98]
Human keratinocytes ^❖	L. infantum, L. major PMs	PMs uptake, transformation into AMs at low levels, no multiplication	LM/G, CLSM/Dapi	[117]
Unidentified cells in primary cultures
Hamster kidney cells ^❖	L. braziliensis, L. donovani PMs	PMs uptake, transformation into AMs, multiplication (not in L. donovani)	LM/G	[118,119], as cited in [7,49]
Chicken embryo muscles ^♦○	L. t. major PMs	PMs uptake, transformation into AMs, multiplication, destruction of host cells after day 3	LM	[102]

Abbreviations: AMs—amastigotes, AO—acridine orange, CFSE—carboxyfluorescein N-succinimidyl ester, CLSM—confocal laser scanning microscopy, DCs—dendritic cells, EtBr—ethidium bromide, FC—flow cytometry, FL—fluorescence microscopy, G—Giemsa, GFP—green fluorescent protein, HE—haematoxylin-eosin, ICC—immunocytochemistry, IHC—immunohistochemistry, LM—light microscopy, MCs—mast cells, MGG—May–Grünwald–Giemsa, MSCs—mesenchymal stems cells, PAS—periodic acid-Schiff, PCR—polymerase chain reaction, PI—propidium iodide, PMs—promastigotes, p.i.—post inoculation, qPCR—quantitative polymerase chain reaction, RFP—red fluorescent protein, SEM—scanning electron microscopy, SPIONs—superparamagnetic iron oxide nanoparticles, TEM—transmission electron microscopy. Symbols: ^❖—in vitro, ^♦—in vivo, ^□—clinical case, ^○—experimental infection. Note: Leishmania species names correspond to the names as provided in the original research articles.