Cited by 836NF-κB Signaling PathwaysInvertebrate Immune Response MechanismsImmune Response and Inflammation

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A CONSERVED SIGNALING PATHWAY: The Drosophila Toll-Dorsal Pathway

Belvin, Marcia P., Anderson, Kathryn V.·1996·Annual Review of Cell and Developmental Biology

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Abstract

▪ Abstract The Toll-Dorsal pathway in Drosophila and the interleukin-1 receptor (IL-1R)-NF-κB pathway in mammals are homologous signal transduction pathways that mediate several different biological responses. In Drosophila, genetic analysis of dorsal-ventral patterning of the embryo has defined the series of genes that mediate the Toll-Dorsal pathway. Binding of extracellular ligand activates the transmembrane receptor Toll, which requires the novel protein Tube to activate the cytoplasmic serine/threonine kinase Pelle. Pelle activity controls the degradation of the Cactus protein, which is present in a cytoplasmic complex with the Dorsal protein. Once Cactus is degraded in response to signal, Dorsal is free to move into the nucleus where it regulates transcription of specific target genes. The Toll, tube, pelle, cactus, and dorsal genes also appear to be involved in Drosophila immune response. Because the IL-1R-NF-κB pathway plays a role in vertebrate innate immunity and because plant homologues of the Toll-Dorsal pathway are important in plant disease resistance, it is likely that this pathway arose before the divergence of plants and animals as a defense against pathogens.

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Crossref · 10.1146/annurev.cellbio.12.1.393

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“Since the identification of the first Toll protein in D. melanogaster (Belvin and Anderson, 1996), Tolls or TLRs have been identified in many vertebrates and invertebrates.”
Background
Human β-Defensin-2 Induction in Human Foreskin Keratinocyte by Synergetic Stimulation with Foods and Escherichia Coli
Nishimura, E., Kato, M., & Hashizume, S. (2003) · Cytotechnology (Dordrecht) · doi:10.1023/b: · Cited by 3
Investigation of an emergent, tetracycline-resistant, abortifacient Campylobacter jejuni clone in a pregnant guinea pig model
Burrough, E. R. (2011) · doi:10.31274/etd-180810-581 · Cited by 3
Nuclear Factor-kappa B
Clem, K. W., & Ip, Y. (2006) · doi:10.1007/978-0-387-40049-5_19 · Cited by 3
Effects of microbial infection on key gene expression in the Toll signaling pathway and immune response in Myzus persicae.
Zhang, C., He, L., Yang, H., & Yang, M.-F. (2025) · Pesticide Biochemistry and Physiology · doi:10.1016/j.pestbp.2025.106611 · Cited by 2
The formin-like regulates innate immunity and antibacterial immune responses in Scylla paramamosain
Li, Y., Zhang, Y.-C., & Zhu, F. (2025) · Aquaculture · doi:10.1016/j.aquaculture.2025.742167 · Cited by 2
The Molecular Toll Pathway Repertoire in Anopheline Mosquitoes
Rhodes, V., Waterhouse, R., & Michel, K. (2024) · bioRxiv · doi:10.1101/2024.09.12.612760 · Cited by 2
“Activation of Pelle leads to the subsequent phosphorylation and degradation of a key inhibitor of Toll signaling, Cactus (Grosshans et al. 1994; Belvin and Anderson 1996).”
Expression and Role of Toll-like Receptors in Facial Nerve Regeneration after Facial Nerve Injury
Lee, J.-M., Yeo, S., Jung, S. Y., Jung, J., Kim, S. S., Yoo, M., Rim, H., Min, H. K., Kim, S. H., & Park, D. (2023) · International Journal of Molecular Sciences · doi:10.3390/ijms241411245 · Cited by 2
“TLR3, which recognizes dsRNA produced during viral replication through a MyD88independent pathway and triggers the production of type I interferon (IFN α/β) [18], also plays a role in other cellular processes.”
Background
Serine protease inhibitors LmSPN2 and LmSPN3 co-regulate embryonic diapause in Locusta migratoria manilensis (Meyen) via the Toll pathway
Feng, S., Zhang, N., Chen, J., Zhang, D., Zhu, K., Cai, N., Tu, X.-B., & Zhang, Z.-H. (2023) · Journal of Integrative Agriculture · doi:10.1016/j.jia.2023.05.019 · Cited by 2
Large-scale analysis of the N-terminal regulatory elements of the kinase domain in plant Receptor-like kinase family
Fu, Q., Liu, Q., Zhang, R., Chen, J., Guo, H.-C., Ming, Z., Yu, F., & Zheng, H.-P. (2022) · bioRxiv · doi:10.1186/s12870-024-04846-7 · Cited by 2
“associated Kinases (IRAKs) from mammals (Belvin and Anderson, 1996; Cao et al., 1996).”
Expression and Function of Toll Pathway Components in the Early Development of the Wasp Nasonia vitripennis
Pers, D., Buchta, T., Özüak, O., Roth, S., & Lynch, J. (2022) · Journal of Developmental Biology · doi:10.3390/jdb10010007 · Cited by 2
Immunity vs Sociality: Adaptive evolution tests suggest social lifestyle exerts greater selection pressures than host-pathogen coevolution in the beesPreprint
Mee, L., & Barribeau, S. M. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.09.07.459261 · Cited by 3
“…been known to play a role in the innate immune response of insects, having been linked to anti-microbial, -fungal and -viral responses through activation of the Toll pathway (Belvin and Anderson, 1996; Silverman and Maniatis, 2001; Bangham et al., 2006; Ferreira et al., 2014; Sheehan et al., 2018).”
Background
Low adaptive and neutral genetic diversity in the endangered Antioquia wren (Thryophilus sernai)
Zapata, D., Rivera-Gutiérrez, H., Parra, J., & González-Quevedo, C. (2020) · Conservation Genetics · doi:10.1007/s10592-020-01313-2 · Cited by 2
RNAi screening identifies a new Toll from shrimp that restricts WSSV infection through activating Dorsal to induce antimicrobial peptides
Li, H.-Y., Yin, B., Wang, S., Fu, Q., Xiao, B., Lu, K., He, J.-G., & Li, C.-Z. (2018) · bioRxiv · doi:10.1101/365197 · Cited by 2
“mammalian TLRs function mainly in immunity, Drosophila Toll1 functions not only in 97 developmental processes (6, 7), but also in innate immunity response to bacterial, fungal, and 98 viral infections (8-10).”
Background
Amphioxus ortholog of ECSIT, an evolutionarily conserved adaptor in the toll and BMP signaling pathways
Lin, Y., Zhang, W., Li, J. W., Zhang, H. W., & Chen, D. Y. (2017) · Molecular Biology · doi:10.1134/s0026893317010125 · Cited by 3
“These expression patterns indicated that AmphiTLR and AmphiECSIT may play a role in establishing the dorsal-ventral axis of the embryo, as did the Toll gene in Drosophila [13, 14].”
Background
Ndae1 Expression and Regulation in Drosophila Embryos
Tevy, M., Seyres, D., Traina, C., Perrin, L., & Capovilla, M. (2014) · PLoS ONE · doi:10.1371/journal.pone.0092956 · Cited by 2
“Establishment of DV asymmetry in the early Drosophila embryo is largely under maternal control [35–38].”
Background
Characterization of ecdysteroid receptor isoforms and their influence on other developmental pathways in Drosophila melanogaster cells
Hönl, C. (2010) · OPen Access Repositorium der Universität Ulm (OPARU) (Ulm University) · doi:10.18725/oparu-1874 · Cited by 2
“After the degradation of cactus, dorsal is released to enter the nucleus (BELVIN AND ANDERSON 1996).”
BackgroundMethodInfluential
Conserved mechanisms of signal transduction by Toll and Toll-like receptors
Gangloff, M., Weber, A., & Gay, N. (2005) · Journal of Endotoxin Research · doi:10.1177/09680519050110050601 · Cited by 2
Development of the early immune system: impact on allergic diseases
Schroeter, C., Gibbons, F. K., & Finn, P. (2002) · doi:10.1016/s0889-8561(02)00012-7 · Cited by 2
“The Toll protein in Drosophila and IL-1R in humans can stimulate signal transduction pathways, which induce the activation of transcription factors such as nuclear factor-k B (NF-kB) [18].”
Background
Cutting edge: recognition of Gram-positive bacterial cell wall components by the innate immune system occurs via Toll-like receptor 2.
Yoshimura, A., Lien, E., Ingalls, R., Tuomanen, E., Dziarski, R., & Golenbock, D. (1999) · Journal of Immunology · doi:10.4049/jimmunol.163.1.1 · Cited by 2
Pathogen-Associated Molecular Patterns, Damage-Associated Molecular Patterns, and Their Receptors in Acute Kidney Injury
Emlet, D. R., Gómez, H., & Kellum, J. A. (2019) · Elsevier eBooks · doi:10.1016/b978-0-323-44942-7.00021-2 · Cited by 4
Toll Family Receptors
Hyvert, Y., & Imler, J. (2010) · Elsevier eBooks · doi:10.1016/b978-0-12-374145-5.00159-5 · Cited by 2
Activation of the Innate immune System: The Toll-like Receptor 4 and Signalling through Ubiquitylation
Gomperts, B. D., Kramer, I. M., & Tatham, P. E. R. (2009) · Elsevier eBooks · doi:10.1016/b978-0-12-369441-6.00015-5
Inflammation and Neuronal Susceptibility to Excitotoxic Cell Death
Malva, J. O., & Bernardino, L. I. (2007) · Springer eBooks · doi:10.1007/978-0-387-70830-0_1 · Cited by 1
Host Defensive and Pharmacological Study of Chlorella Vulgaris Strain CK
Hasegawa, T., Kumamoto, S., Nomoto, K., & Yoshikai, Y. (2005) · Studies in natural products chemistry · doi:10.1016/s1572-5995(05)80046-8 · Cited by 2
Invertebrate Cytokines and the Phylogeny of Immunity
Beschin, A., & Müller, W. E. (2004) · Progress in molecular and subcellular biology · doi:10.1007/978-3-642-18670-7 · Cited by 2
“The Toll pathway has been originally identified due to its pivotal role in the dorsoventral patterning of the Drosophila embryo (Belvin and Anderson 1996).”
BackgroundInfluential
Redox signaling in development: An emerging frontier.
Ulfig, A., & Jakob, U. (2026) · Developmental Cell · doi:10.1016/j.devcel.2026.07.005 · Cited by 1
When Small Meets Smaller: Immune Modulation and Virulence Strategies in Insect–Bacteria Interactions
Bianchi, T., Mastore, M., Banfi, D., Loulou, A., Quadroni, S., & Brivio, M. (2026) · Insects · doi:10.3390/insects17050515 · Cited by 1
“This releases the NF-κ B-like transcription https://doi.org/10.3390/insects17050515 factors Dorsal and Dif, which translocate into the nucleus and induce transcription of AMPs genes, whose transcripts are key elements in bacterial clearance [28].”
Influential
Transcriptional regulation of Toll-like Receptors in Pinna nobilis: Insights into immune response under varying pathogen loads and effect of individual variation
Lattos, A., Coupé, S., Papadopoulos, D. K., & Giantsis, I. (2026) · Gene Reports · doi:10.1016/j.genrep.2025.102415 · Cited by 1
Antimicrobial peptide SCY2 with its interacting proteins Scyreprocin mediate the innate immune defense of Scylla paramamosain against Pseudomonas putida infection.
Li, H., Wang, Y., Bai, Y., Chen, F.-Y., & Wang, K.-J. (2025) · International Journal of Biological Macromolecules · doi:10.1016/j.ijbiomac.2025.149596 · Cited by 1
Cephalic ganglia transcriptomics of the American cockroach Periplaneta americana (Blattodea: Blattidae)
Levy, I., & Arvidson, R. S. (2024) · Journal of Insect Science · doi:10.1093/jisesa/ieae113 · Cited by 1
Trans regulation of an odorant binding protein by a proto-Y chromosome affects male courtship in house fly
Delclós, P. J., Adhikari, K., Mai, A. B., Hassan, O., Oderhowho, A. A., Sriskantharajah, V., Trinh, T., & Meisel, R. P. (2024) · eLife · doi:10.7554/elife.90349 · Cited by 4
“The Toll signaling pathway regulates cellular processes, including embryonic development and immune response to infection ( Belvin and Anderson, 1996 ; Valanne et al., 2011 ).”
Background
Caspar specifies primordial germ cell count and identity in Drosophila melanogaster
Das, S., Hegde, S., Wagh, N., Sudhakaran, J., Roy, A. E., Deshpande, G., & Ratnaparkhi, G. (2024) · bioRxiv · doi:10.7554/elife.98584 · Cited by 1
“Pathogenic invasion of insects induces both the humoral and cellular immune response ( Belvin and Anderson, 1996 ; Williams, 2007 ).”
Influential
The effects of Gram-positive and Gram-negative bacterial toxins (LTA & LPS) on cardiac function in Drosophila melanogaster larvae
Brock, K. E., Elliott, E. R., Abul-Khoudoud, M. O., & Cooper, R. L. (2023) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2023.104518 · Cited by 1
“It is interesting to note that both the toll genes and initial immunological relationship to bacterial infection with Toll receptors were first described in Drosophila (Anderson et al., 1985; Imler and Hoffmann, 2001; Belvin and Anderson, 1996; Lemaitre et al., 1996).”
Background
Single Cell Analysis of the Fate of Injected Oncogenic RasV12 Cells in Adult Wild Type Drosophila
Chen, D., Lan, X., Huang, X.-M., Huang, J., Zhou, X., Miao, Z., Ma, Y.-T., Goto, A., Ji, S.-M., & Hoffmann, J. A. (2023) · Journal of Innate Immunity · doi:10.1159/000529096 · Cited by 1
“We further observed in the violin plots that most genes in the Toll pathway [23, 24], especially those encoding the receptor Toll, the adaptor Myd88 and the negative regulator kinase Cactus were abundantly expressed in most cells of both samples (day 0 and day 11), but their average expression…”
Background
A non-canonical Raf function is required for dorsal–ventral patterning during Drosophila embryogenesis
Lusk, J. B., Chua, E., Kaur, P., Sung, I. C. H., Lim, W.-K., Lam, V., Harmston, N., & Tolwinski, N. (2021) · Scientific Reports · doi:10.1038/s41598-022-11699-3 · Cited by 1
Cellular Behaviors Underlying Pattern Formation and Evolution
Mizutani, C. M. (2016) · Elsevier eBooks · doi:10.1016/b978-0-12-800049-6.00132-3 · Cited by 1
“Toll” Extending Its Gate from Drosophila Development to T Cell Response: Implication in Innate Immunity, Adaptive Immunity and Immunotherapy
Chattopadhyay, S., Pal, S., Pratheek, B. M., Meena, V., Singh, S. K., & Maiti, P. (2011) · doi:10.4137/int.s7422 · Cited by 1
A new era for sepsis treatment? Understanding sepsis as a consequence of host immune response
Hung, D., & Baden, L. (2002) · doi:10.1517/13543776.12.2.181 · Cited by 1
“The gene product or receptor was found to be a homologue of a previously characterised Drosophila melanogaster receptor Toll, which plays a role in the innate immunity of Drosophila and thus, the mouse receptor was called Toll-like receptor (TLR) [10,11].”
Background
Les protéines PGRP, un chaînon manquant de l’immunité innée de la drosophile
Royet, J. (2001) · doi:10.1051/medsci/200117121359 · Cited by 1
“Découverte pour son rôle dans la mise en place de la polarité dorso-ventrale de l’embryon, la voie Toll est bien caractérisée au niveau moléculaire [6].”
Background
How Do Gene Networks Promote Morphological Evolution
Sadier, A. (2019) · Old Questions and Young Approaches to Animal Evolution · doi:10.1007/978-3-030-18202-1_10 · Cited by 1
Antiviral Immune Responses to Invertebrate Iridescent Virus 6 in Drosophila
West, C. (2018) · doi:10.13028/m25962 · Cited by 1
“Before Toll was shown to play a key role in immune function, it was shown to be required during development in embryonic dorsal-ventral patterning by Christiane Nüsslein-Volhard’s laboratory (Belvin and Anderson, 1996).”
Method
Mesoderm Formation in the Drosophila Embryo
Wakabayashi-Ito, N., & Ip, Y. (2006) · Springer eBooks · doi:10.1007/0-387-32963-3_3 · Cited by 1

Authors: Belvin, M. P., & Anderson, K. V.
Affiliations: University of California, Berkeley (US)
ORCID: Belvin, Marcia P.
Anderson, Kathryn V.
Venue: Annual Review of Cell and Developmental Biology
Publisher: Annual Reviews
Published: 1996-11
Volume: 12
Issue: 1
Pages: 393-416
ISSN: 1081-0706 (ISSN-L)
1530-8995
DOI: 10.1146/annurev.cellbio.12.1.393
OpenAlex: W2098336582
Cited by: 836
References: 0
Citations by year: 2026: 7 · 2025: 14 · 2024: 24 · 2023: 18 · 2022: 23 · 2021: 22 · 2020: 12 · 2019: 21 · 2018: 23 · 2017: 17 · 2016: 17 · 2015: 24 · 2014: 24 · 2013: 25 · 2012: 19
FWCI: 5.77
Type: journal article
Language: en
Open access: closed
Locations: Annual Review of Cell and Developmental Biology (publishedVersion)
PubMed (publishedVersion)
Topics: Life Sciences › Biochemistry, Genetics and Molecular Biology › Cancer Research › NF-κB Signaling Pathways
Life Sciences › Immunology and Microbiology › Immunology › Invertebrate Immune Response Mechanisms
Life Sciences › Immunology and Microbiology › Immunology › Immune Response and Inflammation
Keywords: Drosophila, Toll receptor, tubes, dorsoventral patterning, embryonic patterning, innate immunity, interleukin-1 receptor, NF-κB, evolution of signaling pathways, plant disease resistance, conserved signaling pathways
MeSH terms: Animals, Drosophila, Signal Transduction, Gene Expression Regulation, Developmental

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A CONSERVED SIGNALING PATHWAY: The Drosophila Toll-Dorsal Pathway

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