Developmental Biology

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Official Journal of the Society for Developmental Biology www.sciencedirect.com/journal/developmental-biology

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Developmental Biology@devbiol· 20d

#DBfeature A tribute to Gail R Martin, mouse geneticist and mentor extraordinaire (April 12, 1944 - Feb 9, 2026) By Alexandra Joyner doi.org/10.1016/j.ydbio.2026.05.007

Gail Martin looking at the camera, sitting on a bench in front of a lily pad pond in her garden in Berkeley. Photo from Brigid Hogan.

Developmental Biology@devbiol· 21d

#DBfeature #ICYMI 🐸 MicroRNA miR-196a controls neural crest patterning by repressing immature neural ectoderm programs in Xenopus embryos By Alice Godden, Grant Wheeler, et al. tinyurl.com/4mne33bu

Depletion of miR-196a expands neural plate progenitors at the expense of neural crest specification and neuronal differentiation. Embryos were injected into one dorsal blastomere at the 4-cell stage of development with 300 pg of GFP mRNA or of lacZ developed with Red-gal. In all panels the injected side is to the right. Whole mount in situ hybridisation of zic1, (stage 14), zic3, (stage 14), msx2, (stage 14), sox2, (stage 14), pax6, (stage 15) hairy1, (stage 18), eng2, (stage 15) n-tub (stage 18) and elrc1, (stage 18), following MO mediated miRNA KD. Embryos are positioned in dorsal view with anterior at the top. Whole mounts are shown as well as transverse sections (i). Injection of the MO caused an expansion in the NPB markers Zic1 (n = 28/30) and Zic3 (n = 34/37) but a loss in the anterior region of the NPB marker Msx2 (n = 18/19) (A–C). The neural marker Sox2 was expanded in a lateral direction and appeared to lose the intense dark region of expression seen in the control side (see black arrow) (n = 37/37) (D). Expression of the neural marker Pax6 was slightly enlarged across the anterior neural fold (lens primordia) (see white arrow) but reduced in the position of the forebrain stripe (see black arrow) (n = 18/24) (E). The downstream target of Notch – Hairy1 had a reduced but broadened expression (n = 22/25) (F) Eng2 showed a reduced and broadened expression as well as a shift more posterior (n = 19/25) (G). Both neuronal differentiation markers N-tub (n = 24–30) and ELRC (n = 23/24) were reduced following miR-196aMO injection although this was restricted to the anterior most part of the NP (see black arrows). The expression of these two markers in the trigeminal placodes had extended in both an anterior and posterior direction (see white arrows) (H,I).

Developmental Biology@devbiol· 23d

#DBfeature #LatestResearch 🧠 Microtubule binding protein Togaram1 is required for proper development of mammalian forebrain and neural primary cilia, with its knockout leading to a thin and irregular neuron layer By Clarissa Nassar, Savera Shetty, Noelle Dwyer doi.org/10.1016/j.ydbio.2026.05.011

Neuron layer of E12.5 Togaram1 KO cortex is thin and irregular.
(A) Control E12.5 cortical section shows a thick layer of Pax6+ NSCs in the ventricular zone topped by daughter cells forming layers of basal progenitors (Tbr2+) and neurons (Tubb3+).
(B-D) Examples of different neuron layer abnormalities found in Togaram1 KOs.
(B) KO #1 displays a deep ventricular indent and a small neuron cluster to the left of the indent. Neuron layer is thinner above the indent, but is continuous.
(C) KO #2 has a very thin patchy neuron layer with a large neuron cluster.
(D) KO #3 has a ventricular indent with a thin neuron layer with a gap above the indent. White dashed line marks ventricle, yellow dashed lines outline ectopic neuron clusters, yellow arrowheads indicate breaks in the neuronal layer. Scale bars: 100um.

Developmental Biology@devbiol· 26d

#DBfeature #ICYMI Injury-induced Neuregulin-EGFR signaling from muscle mobilizes stem cells for whole-body regeneration in acoels By Brian Stevens, Riley Popp, Heather Valera, Kyle Krueger, Christian Petersen tinyurl.com/4h52at7f

Model. (A) Early wound signaling activates expression of nrg-1 in muscle near the wound site by 6 hours and subsequently in the distal blastema region from 1 to 5 days. As regeneration progresses, nrg-1 expressed from muscle signals through egfr-1, and receiving cells of this signaling could either be neoblasts (gray arrows), muscle, or other cell types. A downstream consequence of the signaling results in localization of neoblast proliferation near the blastema region, which correlates in space and time approximately with the locations of nrg-1's late expression phase. The activity of nrg-1 and egfr-1 promotes blastema outgrowth. (B) Regulatory model identifying a critical role for nrg-1 and egfr-1 functionally downstream of egr. Wounding activates immediate early expression of egr, which is responsible for transcriptional activation of nrg-1 and runt-1. nrg-1 is functionally required for regeneration and acts through egfr-1, and these factors exert critical responses downstream of egr in order to localize proliferation to wound sites and also cause blastema outgrowth. Gray lines: contributions from nrg-1 signal feedback and/or roles for pre-existing NRG-1 protein in the wound activation process (see text). (C) Phylogeny showing the conserved use of Neuregulin and/or EGFR signaling to functionally promote regeneration. Acoels are believed to be either early-branching Deuterostomes within Xenacoelomorpha or form a sister group to all Bilaterian animals (dotted lines).

Developmental Biology@devbiol· 27d

#DBfeature #LatestResearch 🐣 Gene knockout in chick reveals a new in vivo role for Cxcl14 during trigeminal ganglion development By Hugo Urrutia, Junpeng Gao, Marianne Bronner doi.org/10.1016/j.ydbio.2026.06.001

Perturbation of Cxcl14 specifically in placode cells disrupts trigeminal ganglion development. A. Schematic illustrating the in ovo experimental design to perturb Cxcl14, specifically targeting placode cells. Experimental (red) reagents were delivered unilaterally into the ectoderm of HH11 (13 somites) chick embryos. B-E″. Either a Cxcl14 specific gRNA-CRISPR/Cas9 construct (B-C′) or Cxcl14 MO (D-E′) were introduced onto the right side of the embryo. Immunolabeling of SOX10 (magenta) and TUJ1 (green) reveals the effects of loss of Cxcl14 within the trigeminal ganglion, specifically in placode cells at HH16. Arrows indicate position of neural crest-derived cells. Arrowheads indicate position of placode-derived cells.

Developmental Biology@devbiol· 27d

#DBfeature #ICYMI 👁️ Unraveling the developmental heterogeneity of human retinal ganglion cells within the developing retina By Emil Kriukov, Jonathan Soucy, Everett Labrecque, Petr Baranov tinyurl.com/mtvnssdy

Schematic of workflow. Public datasets re-analysis; processing (UMAPs shown); integration and downstream analysis (cartoon of cell fate trajectories and cell-cell interactions); generation of a reference atlas (annotated UMAP); application by sequencing new data and automatically embedding in the atlas and annotating cell types.

Developmental Biology@devbiol· 29d

#DBfeature #LatestResearch 🧠🐁 Neurogenesis and gliogenesis in the developing neocortex are imbalanced in mice lacking the proteoglycan Tsukushi By Sabrina Quaresima, Valerio Licursi, Tenta Ohkubo, Giuseppe Lupo, et al. doi.org/10.1016/j.ydbio.2026.05.009

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Alt text
TSK is expressed in the cortical plate of the developing mouse neocortex.
(A to E) Immunofluorescence with an anti-β-gal antibody (A, B), or X-gal staining (C to E), in coronal brain sections of E14.5 (A), P0 (B, C), P14 (D), or adult (E) TSK+/- mice, showing β-gal expression (green in (A, B); light blue in (C to E)) in the cortical plate between E.14.5 and P14. Hoechst nuclear staining is shown in blue (A, B). Scale bars, 500 μm (A, B) or 1000 μm (C to E).
(F to O) Double immunofluorescence in coronal brain sections of E14.5 TSK+/- mice with an anti-β-gal antibody (green staining) and anti-PAX6 (F, K), anti-Nestin (G, L), anti-TBR2 (H, M), anti-βIII-Tubulin (I, N) or anti-TBR1 (J, O) antibodies (red staining), showing β-gal expression in the cortical plate, partially overlapping with the TBR1+ domain. Images in (K to O) show magnifications of the boxed areas in (F to J). Hoechst nuclear staining is shown in blue. Scale bars, 500 μm.

Developmental Biology@devbiol· Jul 13

#DBfeature #ICYMI 🐠 Single, double, and triple mutant combinations reveal redundant and novel functions of scube genes during zebrafish development By Quoc Duy Tran, Peter D. Currie et al. tinyurl.com/3d8tj7sf

scube mutants display Hh loss-of-function phenotypes. Loss of slow muscle fibres in scube mutant families at 30 hpf. In wild-type zebrafish, slow muscle fibres form a regular array with clearly defined V-shaped somites. In scube1−/−, scube3−/−, and scube1−/−scube3−/−, morphology of slow muscle fibres are comparable to wild-type siblings. In contrast, scube2−/−, scube1−/−scube2−/−, scube2−/−scube3−/−, and the triple scube mutant show severe disruption of slow muscle fibres along the body, including fibre loss (white arrows), abnormal fibre shapes (yellow arrows), and the presence of U-shaped somites (indicated by white dashed lines). The triple mutant shows the most severe phenotype, characterised by the near-complete loss of slow muscle fibres and pronounced abnormalities in the remaining fibres.

Developmental Biology@devbiol· Jul 10

#DBfeature #LatestResearch 🐠 Skeletal remodeling is responsible for the foramen in the skull for the hyoid artery in zebrafish By Miki Iwasaki, Junpei Kuroda, Masanori Nakae, Hironori Wada doi.org/10.1016/j.yd...

Formation of the Ch foramen by bone deposition. Bone, cartilage and endothelial cells are labeled in cyan, green and magenta, respectively. (E′) A single optical section of the boxed region in E. Channel for green and magenta fluorescence is shown in the right panel. n = 3. Anterior is to the left.

Developmental Biology@devbiol· Jul 9

#DBfeature #ICYMI 🪱 C. elegans sperm activation for fertilization requires robust activity of the TAT-5 lipid flippase By Katherine Maniates, Ann Wehman et al. tinyurl.com/5fu7c49e

Cartoon of wild type and tat-5 mutant mature sperm and cell membrane. Wild type sperm is motile and functional thanks to internalised PtdEth in the membrane. Mutant spermatocytes lose membrane asymmetry without tat-5 activity, losing their activation an motility.

Developmental Biology@devbiol· Jul 8

#DBfeature #LatestResearch 🐠 Regulation of the yolk microtubule and actin cytoskeleton by Dachsous cadherins during zebrafish epiboly By Gina Castelvecchi, Lila Solnica-Krezel, et al. tinyurl.com/58dmfkc7

Bundled yolk microtubule array structures in dchs triple mutants A. Confocal images at 50% epiboly of WT and dchs triple mutants stained with DAPI (blue) and for DM1α (microtubules, green). Arrowhead indicates microtubule bundling. B. Quantification of yolk microtubule bundling through measurement of bare yolk areas lacking microtubules at 50% epiboly. ∗∗p < 0.01. WT N = 25 embryos, MZdchs1a−/−; Zdchs1b−/−; MZdchs2−/− N = 32 embryos. C. Confocal images of yolk microtubule meshwork marked with DM1α, just vegetal of YSL of WT and dchs triple mutants at 50% epiboly. D. Quantification of yolk microtubule alignment coherency. ∗∗∗∗p < 0.0001. WT N = 25 embryos, MZdchs1a−/−; Zdchs1b−/−; MZdchs2−/− N = 32 embryos.

Developmental Biology@devbiol· Jul 7

#DBfeature #ICYMI 🪱 Expression of pal-1, the C. elegans Caudal homologue, is required for correct positioning of motor neuron cell bodies in the larval ventral nerve cord and is regulated by the nuclear hormone receptor SEX-1 By N Noblett, T Roenspies, S Flibotte, A Colavita tinyurl.com/2u963pvx

(E) Schematic showing DD and DA neuron organization along the VNC. (F) Representative images of anterior DA and DD neurons in pal-1(zy117), sex-1(zy130) and double mutants. Asterisks mark dorsally mispositioned neurons. Scale bar = 20 μm. (G) Quantification of worms with dorsally mispositioned neurons at L1. Statistics: (C) one-way ANOVA with Tukey's Post-hoc Comparisons, and (G) Chi-squared test with Monte Carlo simulation for 10,000 replicates, followed by a pairwise analysis using Fisher's exact test with Monte Carlo simulation for 10,000 replicates and adjusted with Holm corrections.

Developmental Biology@devbiol· Jul 6

#DBfeature #LatestResearch 🐠 A parallel reporter assay for zebrafish enables simultaneous, quantitative analysis of multiple transcriptional reporters, and cell type-specific comparisons of cis-regulatory module activity By Gloria Ligunas, Stefan Materna tinyurl.com/357w6usv

Parallel reporter analysis reveals cell-type–specific CRM activity in sorted vascular endothelial cells. Activity of barcoded reporters was quantified in sorted vascular endothelial cells relative to the inactive control (C120). Reporters containing the fli CRM showed the strongest activation in this cell type. ubb-containing reporters were significantly activated over controls, but to a lesser extent than in whole embryos. dll4in3 reporter activity was elevated relative to baseline, although the difference from C120 was not statistically significant. Each CRM was tested using two independent barcoded reporters across four biological replicate experiments. Individual points represent barcode-specific measurements from independent biological replicates.

Developmental Biology@devbiol· Jul 3

#DBfeature #ICYMI Loss of CMTR1 leads to gastrulation failure and disrupted anterior-posterior patterning, with unique gene expression profiles between XX and XY embryos By Janelle Welton, Kimberly Tremblay, Jesse Mager tinyurl.com/4ruh8b5k

Cmtr1 KOs exhibit sex specific differences (a) Whole mount E7.5 male and female control and KO embryos. (B) Quantification of embryo size from E7.5 litters using ImageJ. (C) Principal component analysis (PCA) indicating male and female KOs cluster separately. (D) Normalized counts for select non-X or –Y linked differentially expressed genes which are higher in male KOs than female KOs. (E) Normalized counts (DESeq2) for select differentially expressed non-X or –Y linked genes which are higher in female KOs than male KOs. (F) RT-PCR validation of male/female differential expression in mutant embryos

Developmental Biology@devbiol· Jul 2

#DBfeature #LatestResearch Single-cell RNA-seq uncovers cellular diversity and dynamic gene regulatory networks along the progenitor-to-chondrocyte trajectory of late synovial joint development By Ji-Hye Yea, Qin Bian, Patrick Cahan, et al. tinyurl.com/3cwut5ft

Dynamic GRNs associated with inferred transcriptional progression between progenitor-like and chondrogenic states. (A) Heatmap of genes dynamically expressed along the chondrocyte trajectory. Epoch divides cells (columns) and genes (rows) into stages or epochs. Results of enrichment analysis of genes up-regulated in each epoch are shown to the right of the heatmap. The Epoch algorithm also reconstructs dynamic gene regulatory networks (GRNs). The top regulators of each epoch, determined by the network importance metrics of centrality and betweenness, are listed to the right of the enrichment results. (B) Sub-networks of that exemplify genes specific to the early (Clec3b) and late (Col9a3 and Prg4), and their regulators.

Developmental Biology@devbiol· Jul 1

#DBfeature #ICYMI Review: Critical roles of the primitive endoderm in in fetal cell fate specification and patterning By Marcelio Shammami, Alyssa Virola-Iarussi, Ian McCrary, Amy Ralston tinyurl.com/dy5js2f7

PrE signaling before and after implantation. A) During preimplantation development, spontaneous differences in FGF4/FGFR1,2/MEK signaling sort PrE from EPI. B) After implantation and prior to gastrulation, the anterior visceral endoderm (AVE) secretes antagonists of BMP/Nodal/WNT signaling to direct anterior/posterior patterning.

Developmental Biology@devbiol· Jun 30

#DBfeature #LatestResearch Camk2g1 regulates Notch signaling and histone methylation to specify hematopoietic stem cells By Camden Kurtz, Sanyam Patel, Haerin Kim, Sarah Rothschild tinyurl.com/bdp9u37v

Elevated H3K27me3 levels in camk2g1 crispants. Increased levels of H3K27me3 are observed in the dorsal aorta (DA, white arrows) but not the posterior cardinal vein (PCV, green arrows) of camk2g1 fli1:GFP positive morphants (n = 24-30) compared to control embryos (n = 29-30)

Developmental Biology@devbiol· Jun 29

#DBfeature #ICYMI Review: Controllable targeted protein degradation as a promising tool for discovery of novel cellular and developmental mechanisms By De-Li Shi, Xuan Zhao, Chengtian Zhao, Ming Shao tinyurl.com/yc26fzy6

Schematic of the function of common protein degradation systems DeGradFP, AID, dTAG, and PROTAC.

Developmental Biology@devbiol· May 15

#DBfeature Loss of potassium channel Kcnj2 alters electrical activity in palate mesenchyme cells, which affects a network of calcium-induced transcription factors and causes cleft palate in mice. By Yunus Ozekin, Rebecca O'Rourke, Emily Anne Bates tinyurl.com/4c9w8saz

Loss of Kcnj2 affects transcription in Osr1 positive mesenchyme 4 cells. Immunohistochemistry shows that Asporin expression is expanded into more cells in Kcnj2KO/KO palatal shelves. A white line shows where fluorescence was quantified and the arrow shows the region of highest asporin expression.

Developmental Biology@devbiol· May 14

#DBfeature 🐸 TMBIM4 affects left-right patterning via pluripotency exit during gastrulation in X. tropicalis, and its loss replicates the phenotypes of human congenital heart disease patients by Nicholas Diab, Mustafa Khokha et al www.sciencedirect.com/science/arti...

GFP-Tmbim4 localizes to the membrane in Xenopus

Developmental Biology@devbiol· May 13

#DBfeature 🧑‍🎓 The Neural Tube CURE trains students in developmental biology methodology while offering a research course designed to increase opportunities for undergraduate research experience By Anneke Dixie Kakebeen, Joshua Fandel, Lee Niswander tinyurl.com/y2va2xpy

The neural tube course based undergraduate research experience
Scientific goal. Spina bifida: congenital defect, worldwide impact. Candidate human bifida genes --> validate candidate genes in chick embryos.
Course gaols. Developmental biology, research & society, techniques, science communication.
Student outcomes. Self-evaluations show significant gains in concepts, techniques, literature, writing, presenting, self-efficacy.
Experimental workflow. sgRNA/Cas9 in HH4 and HH10 chick embryo, look for neural tube defects.

Developmental Biology@devbiol· May 11

#DBfeature Long-term mitochondria-endoplasmic reticulum complexes in the oocytes and early embryos of a matrotrophic pseudoscorpion Chelifer cancroides By Izabela Jędrzejowska, Karol Małota, Arnold Garbiec tinyurl.com/45etdr9c

mt-ER complexes in postovulatory oocytes and early embryos. Postovulatory oocytes in the lumen of the ovary. Note numerous stick-like mt-ER complexes distributed in the ooplasm among lipid droplets and yolk spheres. Semi-thin sections stained with methylene blue. Ultrastructure of mt-ER complexes seen at the longitudinal (B) and cross (C) sections. TEM
Early embryo. Note that most mt-ER complexes fill the cytoplasm of micromeres (enlarged in inset). Semi-thin sections stained with methylene blue. The cytoplasm of blastomeres contains numerous mt-ER complexes, free mitochondria (encircled by a dotted line) (enlarged in inset E) and folded membranes of endoplasmic reticulum (encircled with a dashed line). TEM.
Abbreviations: c – mt-ER complexes; g, glycogen granules; ld, lipid droplet; oo, oocyte; ow, ovarian wall; solid arrowhead, longitudinal section of mitochondrion; hollow arrowhead, cross section of mitochondrion.

Developmental Biology@devbiol· May 10

#DBfeature 🌊 #DBreview Germ cell formation and reproduction in tunicates by Yasunori Sasakura et al www.sciencedirect.com/science/arti...

Developmental Biology@devbiol· May 9

#DBfeature #DBreview 🪱 Building bridges for oocyte growth: regulation of C. elegans germline architecture and function by oriented cell divisions By Réda Zellag, Kimia Zarnani, Abigail Gerhold, Jean-Claude Labbé tinyurl.com/3ne592zj

Spindle orientation during C. elegans germ cell mitosis positions the cytokinetic ring to favour cytoplasmic bridge duplication. The spatiotemporal pattern of LIN-5/NuMA (blue) cortical localization promotes spindle orientation parallel to the rachis surface in germ cells. This ensures that the cytokinetic ring forms in the orthogonal plane, allowing the cytokinetic ring to interact with the germ cell's rachis face and/or the stable cytoplasmic bridge during ring closure and thus favoring cytoplasmic bridge duplication.

Developmental Biology@devbiol· May 8

#DBfeature A new hypothesis about meiosis "rejuvenating" germ cells each generation by ensuring they continue to match the species' DNA specifications by Allan Spradling et al www.sciencedirect.com/science/arti...

Polarized germline cysts are formed in Drosophila, Xenopus and mice

Developmental Biology@devbiol· May 7

#DBfeature ShH10 efficiently targets all subtypes of supporting cells along the length of cochlear duct without impacting cochlear cellular architecture or auditory function By Yun Ji Bertken, Yeeun Kim, Juan Llamas, Assaf Beck, Aaron Nagiel, Ksenia Gnedeva tinyurl.com/mum9vktc

ShH10 minimally infects cells in the brain and contralateral ear. In mice injected with ShH10 at P1, GFP expression (green) is not seen in the brain at or P21. In contrast, high levels of GFP expression are seen in the brain of AAV-ie-K558R injected mice, including nearly 100 % of Purkinje cells in the cerebellum at P21. Cell nuclei are stained with DAPI in the brain sections (blue). Scale bar 50 μm.

Developmental Biology@devbiol· May 6

#DBfeature A modular enhancer mediates SCRT2 repression of ISLET1 in the spinal cord by Vitória S. Botezelli et al www.sciencedirect.com/science/arti...

SCRT2 represses ISLET1 expression in the dorsal domain

Developmental Biology@devbiol· May 5

#DBfeature Elongation of the nascent avian foregut requires coordination of intrinsic and extrinsic cell behaviors By Olivia Powell, Emily Garcia, Vanshika Sriram, Yi Qu, Nandan L. Nerurkar tinyurl.com/52pecyc5

Endodermal tissue deformations at the AIP during foregut elongation. (A) Velocity fields quantified from time lapse of cell movements between HH stages 8+ and 10+. Orientation and magnitude of arrows indicate orientation and speed of cell movements, respectively; underlying heat map indicates speed (n = 3); scale = 200 μm. (B) Principal strain rates computed from velocity fields in (A), where major and minor cruciform axes represent maximum and minimum principal strain rates D1 and D2, respectively, and their orientations represent the associated principal directions. Negative/compressive principal strain rates are indicated in blue, and positive/extensional strain rates are indicated in red; strain rate scale = 0.01 %/s; spatial scale is as in (A). (C) Maximum and minimum principal strain rates calculated from velocity fields at HH stage 8, following electroporation with 3 nls-GFP RNA control (left) and DeAct-GFP RNA (right) (n = 3);

Developmental Biology@devbiol· May 4

#DBfeature 🐣 Nkx2.5 triggers vascular remodeling in chicken but not in emu wing mesoderm, where Nkx2.5-positive limb cells belonged to muscle instead by Ingrid Cordeiro, Mikiko Tanaka et al www.sciencedirect.com/science/arti...

Increased expression of Scl is detected 24 h after Nkx2.5 overexpression, which was followed by vascular changes 48 h after electroporation

Developmental Biology@devbiol· May 3

#DBfeature A low-cost, high-throughput pipeline for 3D imaging of embryonic mouse hearts using lightsheet microscopy By Xiangyang Liu, Jianfeng Wang, Youshi Chen, Hongjun Shi tinyurl.com/yc46asz2