Lu, T.-M. | bioRxiv Science

bioRxiv2025

The xenacoelomorph gonopore is homologous to the bilaterian anus

The bilaterian through gut with an anal opening is a key invention in animals, since it facilitates effective food processing, which allows animals to grow to a larger body size. However, because non-bilaterian animals lack a through gut, the evolution of anus is still debated. The formation of bilaterian hindgut is governed by the spatial expression of several transcription factors (e.g. Caudal and Brachyury) under the control of Wnt signaling. This conserved pattern has been used to support the homology of the anus of protostomes (insects, snails) and deuterostomes (sea urchins, humans). Here we show, that these bilaterian "hindgut" marker genes are expressed around the male gonopore of several xenacoelomorphs, which have a blind gut without an anal opening. These findings suggest a deep evolutionary relationship between the xenacoelomorph male gonopore and the bilaterian anus. Since xenacoelomorphs are the potential sister group to all remaining Bilateria, our results suggest that the bilaterian anus evolved from a male gonopore that came in contact with the digestive endoderm to form the posterior opening.

developmental biology↗

bioRxiv2021

The evolution of the metazoan Toll receptor family and its expression during protostome development

BackgroundToll-like receptors (TLRs) play a crucial role in immunity and development. They contain leucine-rich repeat domains, one transmembrane domain, and one Toll/IL-1 receptor domain. TLRs have been classified into V-type/scc and P-type/mcc TLRs, based on differences in the leucine-rich repeat domain region. Although TLRs are widespread in animals, detailed phylogenetic studies of this gene family are lacking. Here we aim to uncover TLR evolution by conducting a survey and a phylogenetic analysis in species across Bilateria. To discriminate between their role in development and immunity we furthermore analyzed stage-specific transcriptomes of the ecdysozoans Priapulus caudatus and Hypsibius exemplaris, and the spiralians Crassostrea gigas and Terebratalia transversa. ResultsWe detected a low number of TLRs in ecdysozoan species, and multiple independent radiations within the Spiralia. V-type/scc and P-type/mcc type-receptors are present in cnidarians, protostomes and deuterostomes, and therefore they emerged early in TLR evolution, followed by a loss in xenacoelomorphs. Our phylogenetic analysis shows that TLRs cluster into three major clades: clade is present in cnidarians, ecdysozoans, and spiralians; clade {beta} in deuterostomes, ecdysozoans, and spiralians; and clade {gamma} is only found in spiralians. Our stage-specific transcriptome and in situ hybridization analyses show that TLRs are expressed during development in all species analyzed, which indicates a broad role of TLRs during animal development. ConclusionsOur findings suggest that the bilaterian TLRs likely emerged by duplication from a single TLR encoding gene (proto-TLR) present in the last common cnidarian-bilaterian ancestor. This proto-TLR gene duplicated before the split of protostomes and deuterostomes; a second duplication occurred in the lineage to the Trochozoa. While all three clades further radiated in several spiralian lineages, specific TLRs clades have been presumably lost in others. Furthermore, the expression of the majority of these genes during protostome ontogeny suggests their involvement in immunity and development.

evolutionary biology↗