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Table 1 Candidate TEs located nearby immune-related genes. TE length is indicated as full-length (FL) or truncated (T). In the case of truncated elements, whether the insertion is a solo LTR, a TIR or 5′ truncated (5′) is also indicated. NS, not significant. NA, not analyzed

From: Regulatory regions in natural transposable element insertions drive interindividual differences in response to immune challenges in Drosophila


TE family (class)

TE start position

TE length

Evidences of selection

Gene (position)

Immune-related evidence


pogo (DNA)

3R: 29760415

1146 bp (T: TIR)

Fst, nSL [52]

kay (4434 bp 5′)

Expression. Component of the JNK pathway, essential for antimicrobial peptide release [56, 57]. Upregulated in imd and bsk mutant LPS-induced S2 cells, and downregulated in Rel mutants [58]. Upregulated in larvae infected with P. entomophila (gram-negative) [46], and after 4 h of infection with P. entomophila [41].


Doc (non-LTR)

3 L: 6040416

2305 bp (T: 5′)

Allele age [59]

Jon65Aiv (281 bp 3′)

Expression. Upregulated after septic injury with mixed bacteria: M. luteus (gram-positive) and E. coli (gram-negative) [60]. Downregulated after 4 h of infection with P. entomophila [41]. Downregulated after oral infection with the avirulent gacA P. entomophila [61].


BS (non-LTR)

3 L: 7130011

126 bp (T: 5′)

H12, nSL [52]

CG15829 (338 bp 3′)

Expression. Upregulated after infection by septic injury with mixed bacteria (gram-positive and gram-negative), and regulated by Rel [60]. Upregulated after 4 h of infection with P. entomophila [41]. Upregulated in transgenic flies expressing a SINV replicon (alphavirus) [62].

CG8628 (739 bp 5′)

Expression. Upregulated in microbiota associated flies vs. germ free flies [63], after infection with several pathogens (gram-positive and gram-negative, fungi, protozoa) [64], and downregulated after 4 h of infection with P. entomophila [41].


Burdock (LTR)

2R: 9151357

6413 bp (T: LTR)


CG8008 136 bp 3′)

Expression. Induced by LPS (gram-negative) in an IKK-dependent manner in S2 cell cultures [65]. Upregulated after E. coli (gram-negative) infection in S2 cells [66].


Juan (non-LTR)

3R: 15132112

2995 bp (T: 5′)


CG42788 (180 bp 5′)

Expression. Downregulated in response to P. rettgeri (gram-negative) infection in females [67].


Juan (non-LTR)

X: 8031495

4249 bp (FL)


CG2233 (12 bp 3′)

Expression. Downregulated in PEBP1 mutant L3 larvae, which are more resistant to M. luteus (gram-positive) and E. coli (gram-negative) infection [68]. Latitudinal expression differentiation after infection with E. coli and M. luteus mix in temperate vs. tropical populations [69].


G5 (non-LTR)

2R: 7317828

51 bp (T: 5′)


Dscam1 (46 bp 3′)

Expression. Required in hemocytes for efficient phagocytosis and binds to E. coli (gram-negative) [70].


BS (non-LTR)

3R: 16041234

102 bp (T: 5′)


Pnr (3′UTR)

Expression. pnr is a modifier of the Toll pathway and RNAi mutants show Imd pathway hyperactivation when infected with E. cloacae (gram-negative) and M. luteus and E. faecalis (gram-positive) [71].


BS (non-LTR)

2R: 18807871

800 bp (T: 5′)


CG15096 (479 bp 3′)

Expression. Downregulated in Oregon R and Rel-mutant flies with microbiota compared to axenic flies [72], and after P. entomophila infection [41].


Transpac (LTR)

3 L: 12863675

5500 bp (FL)


CG10943 (816 bp 5′)

Expression. Upregulated in Oregon R and Rel-mutant flies with microbiota compared to axenic flies [72], 24 h after infection with O. muscaedomesticae (protozoan) [64], and after P. entomophila infection [41].


pogo (DNA)

X: 21399382

2067 bp (FL)


lcs (2,067 bp 5′)

Expression. Involved in virus response, downregulated in males infected with sigma virus [73]. Upregulated in young fly guts [72].


invader4 (LTR)

3R: 16189464

347 bp (T: solo LTR)

CL test, TajimaD, Phenotypic [74]

Bin1 (5′UTR)

Survival. Mutant larvae are more sensitive to fungal A. fumigatus (fungi) infection [75].


roo (LTR)

2R: 9871090

434 bp (T: solo LTR)

TajimaD, iHS, H12, Phenotypic [76]

cbx (first intron)

Survival. Mutant flies are more sensitive to S. aureus (gram-positive) septic infection, but not to S. typhimurium (gram-negative) infection [77].


mdg1 (LTR)

X: 3785867

189 bp (T: LTR)

TajimaD [78]

tlk (intron)

Survival. Involved in antimicrobial humoral response to gram-negative [57]. tlk knockdown, together with other five genes knocked-down, reduces phagocytosis of E. coli (gram-negative) and S. aureus (gram-positive) in S2 cells [79].



3 L: 15554974

1732 bp (FL)


AGO2 (first intron)

Survival. Involved in defense response to virus infections [80, 81], and interacts with Imd pathway proteins during gram-negative infection [82].



3 L: 17799864

1732 bp (FL)


NUCB1 (first intron)

Survival. Mutants are more resistant to V. cholerae (gram-negative) oral infection [83].


1360 (DNA)

2 L: 17432071

48 bp (T: TIR)

iHS [52]

Dif (first intron)

Survival and expression. Transcription factor involved in defense response to fungus and gram-positive bacteria and mediates Toll pathway activation [84,85,86,87,88]. Dif mutants are susceptible to fungi and gram-positive bacterial infection [88, 89]. Upregulated in guts from P. entomophila infected flies [41].


BS (non-LTR)

2R: 9945496

131 bp (T: 5′)


Mef2 (first intron)

Survival and expression. Adult Mef2 mutant males are more sensitive to E. cloacae (gram-negative) and M. marinum (gram-positive) septic infection [90]. Upregulated after 4 h of infection with P. entomophila (gram-negative) [41].


297 (LTR)

2R: 23877783

414 bp (T: LTR)


TM4SF (1 bp 5′)

A tetraspanin transmembrane protein, which modulate immune-signaling in Drosophila [53].

ken (340 bp 3′)

JAK-STAT. Member of JAK-STAT pathway [91]. JAK-STAT pathway plays a role in immune response in D. melanogaster [54].