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Results for “Developmental Biology”

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bioRxiv2026

Real-time, label-free assessment of cell fusion dynamics by high-content imaging

Cell-cell fusion is a fundamental biological process underlying diverse physiological and pathological phenomena, yet its quantitative analysis remains methodologically challenging due to its dynamic, heterogeneous, and multistep nature. Existing approaches to assess fusion largely rely on endpoint assays or manual scoring, limiting temporal resolution, scalability, and reproducibility. Here, we present a label-free, high-content live-cell imaging pipeline for real-time quantification of cell fusion dynamics, developed and validated using trophoblast syncytialization as a model system. The method integrates automated image acquisition with a reproducible, stepwise analysis workflow combining supervised texture-based segmentation, morphology-based measurements, and intensity-independent texture analysis. We define quantitative metrics, including the ratio of total cluster area to the number of detected clusters and cytoplasmic granularity features, that together discriminate bona fide fusion events from non-fusion-related cellular clustering or proliferation. Using canonical pharmacological inducers and inhibitors of fusion, we demonstrate the specificity and sensitivity of these parameters for detecting fusion-associated remodeling over time. We further demonstrate the scalability of the pipeline through high-throughput screening of biologically relevant growth factors, hormones, and inhibitors, enabling classification of modulators based on their independent, synergistic, or antagonistic effects on fusion dynamics. Consistent results obtained in an independent model further support its potential applications to additional fusion systems. By providing a robust, reproducible, and adaptable framework for time-resolved fusion analysis, this methodology bridges the gap between qualitative observation and quantitative kinetic assessment. Thus, the approach could be readily extended to other cell fusion systems following system-specific parameter optimization, offering a versatile platform for both mechanistic studies and discovery-driven screening applications.

developmental biology↗

bioRxiv2021

A temporally controlled sequence of X-chromosomeinactivation and reactivation defines female mouse in vitro germ cells with meiotic potential

The early mammalian germ cell lineage is characterized by extensive epigenetic reprogramming, which is required for the maturation into functional eggs and sperm. In particular, the epigenome needs to be reset before parental marks can be established and then transmitted to the next generation. In the female germ line, reactivation of the inactive X-chromosome is one of the most prominent epigenetic reprogramming events, and despite its scale involving an entire chromosome affecting hundreds of genes, very little is known about its kinetics and biological function. Here we investigate X-chromosome inactivation and reactivation dynamics by employing a tailor-made in vitro system to visualize the X-status during differentiation of primordial germ cell-like cells (PGCLCs) from female mouse embryonic stem cells (ESCs). We find that the degree of X-inactivation in PGCLCs is moderate when compared to somatic cells and characterized by a large number of genes escaping full inactivation. Nevertheless, PGCLCs that fail to undergo X-inactivation show an abnormal gene expression signature and deficiencies in meiotic entry. Subsequent to X-inactivation we observe gradual step-wise X-reactivation, which is mostly completed by the end of meiotic prophase I. Cells deviating from these progressive kinetics and undergoing X-reactivation too rapidly fail to enter a meiotic trajectory. Our data reveals that a fine-tuned X-inactivation and -reactivation cycle is a critical feature of female germ cell developmental competence towards meiosis and oogenesis

developmental biology↗

bioRxiv2023

Safety Assessment of Intravenous Injection of Rat Embryonic Proteome Extract for in-vivo Regenerative Therapies

Back groundTissue differentiation and organogenesis are exclusively embryonic-phase activities, and these two activities are strictly under the control of the embryonic proteome, the developing embryo is a prime source of molecules for regenerative therapy. Many ex-vivo rudimentary organoid culture studies confirmed the inherent biological function of embryonic factors. Before using embryonic proteins for in vivo therapies, their safety should be confirmed first. Hence, we aimed for an in vivo study to inject rat embryonic proteome extract (EPE) through the intravenous route and investigate the impact on immunological, biochemical, and hematological parameters in the adult rats. MethodsIn this study, we isolated rat embryonic proteins from the 14th, 16th, and 19th embryonic days by homogenization of embryos and isolated protein extracts through ultra-centrifugation. Six pairs of rats have been taken; six are allotted for control, and six are for the embryonic extract injection. The isolated embryonic protein extract was injected intravenously into the treatment group of rats and the normal saline into the control group. After the injections, blood samples are collected from both the treated and control groups to analyse immunological markers Il6, CRP, biochemical parameters creatinine, urea, sugar, proteins, albumin, globulin, and hematological parameters total leucocyte count, neutrophils, and lymphocyte percentage ResultAfter the EPE injection, biochemical parameters, immunological markers, and hematological parameters were analyzed in both control and treated groups. All the above mentioned parameters are within normal limits. Statistical analyses were done using the t-test for unequal variance (p< 0.05). We observed no significant difference between the control and treated groups, so in vivo injections of embryonic protein extract is safe for the use of in vivo regenerative therapies. ConclusionDespite extensive in vitro studies confirmed the biological function of embryonic growth factors for organ differentiations and rudimentary organoids, but in vivo clinical applications are not yet started because of immunogenicity, heterogeneity, and tumorigeniety. In our study we injected EPE intravenously and showed that EPE is non-immunogenic, non-heterogeneic, and non-tumorigenic. This study concludes that EPE is safe for in vivo injection, so that further studies can be continued for intra-organ injections for organ regenerative therapy.

developmental biology↗

bioRxiv2020

Single cell RNA analysis of trunk neural crest cells in zebrafish identifies pre-migratory populations expressing markers of differentiated derivatives

The neural crest is a migratory population of stem-like cells that contribute to multiple traits including the bones of the skull, peripheral nervous system, and pigment. How neural crest cells differentiate into diverse cell types is a fundamental question in the study of vertebrate biology. Here, we use single cell RNA sequencing to characterize transcriptional changes associated with neural crest cell development in the zebrafish trunk during the early stages of migration. We show that neural crest cells are transcriptionally diverse, and identify pre-migratory populations already expressing genes associated with differentiated derivatives. Further, we identify a population of Rohon-Beard neurons that are shown to be sources of Fgf signaling in the zebrafish trunk. The data presented identify novel genetic markers for multiple trunk neural crest cell populations and Rohon-Beard neurons providing insight into previously uncharacterized genes critical for vertebrate development.

developmental biology↗

bioRxiv2021

Perivascular cells support folliculogenesis in the developing ovary

Granulosa cells, supporting cells of the ovary, are essential for ovarian differentiation by providing a nurturing environment for oogenesis. Sufficient numbers of granulosa cells are vital for establishment of follicles and the oocyte reserve; therefore, identifying the cellular source from which granulosa cells are derived is critical for understanding basic ovarian biology. One cell type that has received little attention in this field is the perivascular cell. Here we use lineage tracing and organ culture techniques in mice to identify ovarian Nestin+ perivascular cells as multipotent progenitors that contribute to granulosa, thecal, and pericyte lineages. Maintenance of these progenitors was dependent on vascular-mesenchymal Notch signaling. Ablation of postnatal Nestin+ cells resulted in a disruption of granulosa cell specification and an increased incidence of polyovular ovarian follicles, thus uncovering key roles for vasculature in ovarian differentiation. These findings may provide new insights into the origins of female gonad dysgenesis and infertility.

developmental biology↗

bioRxiv2022

The addition of antibiotics to embryo culture media caused altered expression of genes in pathways governing DNA integrity in mouse blastocysts

Antibiotics are common components of embryo culture media and minimize the risk of microbial contamination and infection during assisted reproductive technology procedures (ART). The effects of two aminoglycoside antibiotics (gentamicin, streptomycin) and penicillin on the global profiles of gene expression (DE) were assessed by RNA-seq of individual mouse blastocysts. Zygotes were cultured in an optimized defined medium formulation (KSOM) to which a dose range of each antibiotic was added. A dose-dependent retardation of the rate of zygote development to morphologically normal blastocyst was observed and this was accompanied by a reduction in the number of cells present within the resulting blastocysts. The lowest dose of each antibiotic tested (similar to the concentrations used in clinical grade media) caused significant differential expression of approximately 1800 genes. In most cases antibiotic treatment caused a reduction in gene expression and gene ontology analysis showed that down regulated genes were enriched for several biological processes related to the maintenance of genomic integrity. All three antibiotics caused the downregulation of Brca2, Blm, Rad51c and Rad54l, genes involved DNA homologous recombination pathways and also several p53-dependent genes. Immunolocalization studies showed that each antibiotic also reduced level of BRCA2 and RAD51C detected within blastocysts. The present study shows that the supplementing embryo culture media with antibiotics is associated with wide ranging alterations in gene expression in a manner that could potentially compromise the genomic integrity of the resulting embryos.

developmental biology↗

bioRxiv2022

High throughput expression-based phenotyping and RNAi screening reveals novel regulators of planarian stem cells

The complexity of cell types and states revealed by single-cell RNAseq atlases presents a challenge for the systematic analysis of fate determinants using traditional screening methodologies. Differentiation in the planarian Schmidtea mediterranea exemplifies this problem, as these animals continuously produce over 100 differentiated cell types for homeostasis and regeneration using neoblast adult pluripotent stem cells. The signaling factors enabling neoblast self-renewal and selective differentiation of these many fates are still incompletely understood. We developed a method using high-throughput expression profiling by qPCR and whole-animal RNAseq to simultaneously assess numerous cell fate markers as the phenotypic readout in large-scale RNAi screens. Applying this method, we performed an RNAi screen of 400 kinases, receptors, and other regulatory molecules to reveal specific functions for 30 previously unknown factors in neoblast biology. 17 genes were required for neoblast maintenance, including factors likely involved in cell-cycle regulation, nutrient sensing, and chromatin modification. Multidimensional expression information additionally revealed several specific regulators of other neoblast activities, including a mink1 kinase regulating global neoblast differentiation, the energy responsive kinase adenylate kinase-2 regulating intestine specification within the neoblast population, an RNA acetyl transferase nat10 regulating epidermal differentiation, and a pak1 kinase restricting neoblast localization to prevent tissue outgrowths. These results identify several new regulators of neoblast activities and demonstrate the applicability of expression-based screening for systematic analysis of stem cell phenotypes in whole animals.

developmental biology↗

bioRxiv2023

Turing pattern prediction in three-dimensional domains: the role of initial conditions and growth

Reaction-diffusion systems have been widely used to model pattern formation in biological systems. However, the emergence of Turing patterns in three-dimensional (3D) domains remains relatively unexplored. A few studies on this topic have shown that extending pattern formation from 2D to 3D is not straightforward. Linear stability analysis, which is commonly used to associate admissible wave modes with predicted patterns in 1D and 2D, has yet to be applied in 3D. We have used this approach, together with finite element modelling of a Turing system with Schnakenberg kinetics, to investigate the effects of initial conditions and growing domains on the competition between admissible modes in 3D Turing pattern emergence. Our results reveal that non-random initial conditions on the activator play a stronger role than those of the inhibitor. We also observe a path dependency of the evolving pattern within a growing domain. Our findings shed new light on the mechanisms ensuring reliable pattern formation in 3D domains and have important implications for the development of more robust models of morphogen patterning in developmental processes.

developmental biology↗

bioRxiv2023

Single cell transcriptomics of the human parasite Schistosoma mansoni first intra-molluscan stage reveals tentative tegumental and stem cell regulators.

BackgroundSchistosomiasis is a major Neglected Tropical Disease, caused by the infection with blood flukes in the genus Schistosoma. To complete the life cycle, the parasite undergoes asexual and sexual reproduction within an intermediate snail host and a definitive mammalian host, respectively. The intra-molluscan phase provides a critical amplification step that ensures a successful transmission. However, the cellular and molecular mechanisms underlying the development of the intra-molluscan stages remain poorly understood. MethodsS. mansoni mother sporocysts were dissociated into single cell suspensions, and live cells enriched and sequenced using the single cell 10X Genomics Chromium platform. We defined somatic and stem/germinal cell clusters, identified cell type-enriched Gene Ontology (GO) terms, and predicted transcription factor binding sites for key marker genes. ResultsSix cell clusters comprising stem/germinal, two tegument, muscle, neuron, and parenchyma were identified and validated by Fluorescence in situ Hybridisation (FISH). GO term analysis predicted key biological processes for each of the clusters. Using the Self-Assembling Manifold (SAM) algorithm, three sub-clusters were identified within the stem/germinal cell population. Furthermore, transcription factor binding sites and putative transcription factors were predicted for stem/germinal and tegument clusters. ConclusionsWe report a spatially validated single cell transcriptomic analysis of the first intra-molluscan stage of S. mansoni. Key cell regulators were identified, paving the way for future analyses to unveil their role during the parasite development and interaction with its intermediate host.

developmental biology↗

bioRxiv2025

XIST Drives X-Chromosome Inactivation and Safeguards Female Extraembryonic Cells in Humans

Dosage compensation of sex chromosomes through X-chromosome inactivation (XCI) is required for mice extra-embryonic tissue growth and embryo development. The species specificity in mechanisms and timing leading to XCI during early embryogenesis, however, left the key question of the interdependence between XCI and human development open. Here, we show that the differentiation of naive human pluripotent stem cells to trophoblast stem cells and extraembryonic mesoderm cells triggers XCI. The inactive X chromosome, however, displays an atypical chromatin state, lacking classical enrichment of heterochromatin markers and DNA methylation. We demonstrate that extraembryonic differentiation and XCI are kinetically and functionally linked. Using loss of function approaches, we prove that XIST is required for human XCI establishment. We also reveal that XCI is key for the survival of human female extraembryonic cells. Our work therefore links XCI to the formation of extraembryonic annexes, with important consequences for human reproductive biology. HIGHLIGHTSO_LINaive hPSCs to EXMCs and TSCs differentiation recapitulates human XCI C_LIO_LIThe Xi has an unusual chromatin status in human extraembryonic cells C_LIO_LIXIST is required for the establishment of human XCI C_LIO_LIXCI supports healthy development of human female extraembryonic cells C_LI

developmental biology↗

bioRxiv2019

Neural crest cells bulldoze through the microenvironment using Aquaporin-1 to stabilize filopodia

Neural crest migration requires cells to move through an environment filled with dense extracellular matrix and mesoderm to reach targets throughout the vertebrate embryo. Here, we use high-resolution microscopy, computational modeling, and in vitro and in vivo cell invasion assays to investigate the function of Aquaporin-1 (AQP-1) signaling. We find that migrating lead cranial neural crest cells express AQP-1 mRNA and protein, implicating a biological role for water channel protein function during invasion. Differential AQP-1 levels affect neural crest cell speed, direction, and the length and stability of cell filopodia. Further, AQP-1 enhances matrix metalloprotease (MMP) activity and colocalizes with phosphorylated focal adhesion kinases (pFAK). Co-localization of AQP-1 expression with EphB guidance receptors in the same migrating neural crest cells raises novel implications for the concept of guided bulldozing by lead cells during migration.

developmental biology↗

bioRxiv2019

The C. elegans homolog of the Evi1 proto-oncogene, egl-43, coordinates G1 cell cycle arrest with pro-invasive gene expression during anchor cell invasion

Cell invasion allows cells to migrate across compartment boundaries formed by basement membranes. Aberrant cell invasion is a first step during the formation of metastases by malignant cancer cells.\n\nAnchor cell (AC) invasion in C. elegans is an excellent in vivo model to study the regulation of cell invasion during development. Here, we have examined the function of egl-43, the homolog of the human Evi1 proto-oncogene (also called MECOM), in the invading AC. egl-43 plays a dual role in this process, firstly by imposing a G1 cell cycle arrest to prevent AC proliferation, and secondly, by activating pro-invasive gene expression. We have identified the AP-1 transcription factor fos-1 and the Notch homolog lin-12 as critical egl-43 targets. A positive feedback loop between fos-1 and egl-43 induces pro-invasive gene expression in the AC, while repression of lin-12 Notch expression by egl-43 ensures the G1 cell cycle arrest necessary for invasion. Reducing lin-12 levels in egl-43 depleted animals restored the G1 arrest, while hyperactivation of lin-12 signaling in the differentiated AC was sufficient to induce proliferation.\n\nTaken together, our data have identified egl-43 Evi1 as a critical factor coordinating cell invasion with cell cycle arrest.\n\nAuthor summaryCells invasion is a fundamental biological process that allows cells to cross compartment boundaries and migrate to new locations. Aberrant cell invasion is a first step during the formation of metastases by malignant cancer cells.\n\nWe have investigated how a specialized cell in the Nematode C. elegans, the so-called anchor cell, can invade into an adjacent epithelium. Our work has identified an oncogenic transcription factor that controls the expression of specific target genes necessary for cell invasion, and at the same time inhibits the proliferation of the invading anchor cell.\n\nThese findings shed light on the mechanisms, by which cells decide whether to proliferate or invade.

developmental biology↗

bioRxiv2020

DYRK2 is a ciliary kinase involved 1 in vertebrate Hedgehog signal transduction

Primary cilia are organelles specialized for signaling. We previously defined the proteomes of sea urchin and sea anemone cilia to identify ciliary proteins that predate the origin of bilateria. This evolutionary perspective on cilia identified DYRK2, a kinase not been previously implicated in ciliary biology. We found that DYRK2 localizes to cilia and that loss of DYRK2 disrupts ciliary morphology. We also found that DYRK2 participates in ciliary Hh signal transduction, communicating between SMO and GLI transcription factors. Mutation of mouse Dyrk2 resulted in skeletal defects reminiscent of those caused by loss of Indian hedgehog (Ihh). Like Dyrk2 mutations, pharmacological inhibition of DYRK2 dysregulates ciliary length control and attenuates Hedgehog signaling. Thus, DYRK2 is required for ciliary morphology, for Hedgehog signaling in vitro, and for skeletal development. We propose that DYRK2 is part of the mechanism that transduces SMO to activate GLI transcription factors within cilia.

developmental biology↗

bioRxiv2021

Multiple morphogens and rapid elongation promote segmental patterning during development

The vertebrate hindbrain is segmented into rhombomeres (r) initially defined by distinct domains of gene expression. Previous studies have shown that noise-induced gene regulation and cell sorting are critical for the sharpening of rhombomere boundaries, which start out rough in the forming neural plate (NP) and sharpen over time. However, the mechanisms controlling simultaneous formation of multiple rhombomeres and accuracy in their sizes are unclear. We have developed a stochastic multiscale cell-based model that explicitly incorporates dynamic morphogenetic changes (i.e. convergent-extension of the NP), multiple morphogens, and gene regulatory networks to investigate the formation of rhombomeres and their corresponding boundaries in the zebrafish hindbrain. During pattern initiation, the short-range signal, fibroblast growth factor (FGF), works together with the longer-range morphogen, retinoic acid (RA), to specify all of these boundaries and maintain accurately-sized segments with sharp boundaries. At later stages of patterning, we show a nonlinear change in the shape of rhombomeres with rapid left-right narrowing of the NP followed by slower dynamics. Rapid initial convergence improves boundary sharpness and segment size by regulating cell sorting and cell fate both independently and coordinately. Overall, multiple morphogens and tissue dynamics synergize to regulate the sizes and boundaries of multiple segments during development. Author SummaryIn segmental pattern formation, chemical gradients control gene expression in a concentration-dependent manner to specify distinct gene expression domains. Despite the stochasticity inherent to such biological processes, precise and accurate borders form between segmental gene expression domains. Previous work has revealed synergy between gene regulation and cell sorting in sharpening borders that are initially rough. However, it is still poorly understood how size and boundary sharpness of multiple segments are regulated in a tissue that changes dramatically in its morphology as the embryo develops. Here we develop a stochastic multiscale cell-base model to investigate these questions. Two novel strategies synergize to promote accurate segment formation, a combination of long- and short-range morphogens plus rapid tissue convergence, with one responsible for pattern initiation and the other enabling pattern refinement.

developmental biology↗

bioRxiv2025

Tuning Mitotic Recombination with Patterned DNA Nicks for Precision Mosaic Analysis

CRISPR/Cas9-based mosaic analysis is a powerful tool for in vivo genetics but is limited by cytotoxicity and mutagenesis associated with DNA double-strand breaks (DSBs). Here, we establish Cas9-derived nickases as safer and more reliable alternatives for inducing mitotic recombination in Drosophila. We demonstrate that single-strand nicks are sufficient to generate mosaic clones and systematically dissect the parameters governing this process. We find that clone frequency can be controlled by the gRNA nicking pattern, with two distant nicks on the same DNA strand synergistically enhancing recombination by over nine-fold compared to a single nick. Based on these findings, we propose a mechanistic model for nick-induced crossover and provide a versatile toolkit for generating tissue-specific nickases. This work establishes nickase-based MAGIC as a superior method for high-fidelity clonal analysis, enabling more precise investigation of gene function in development and disease. SIGNIFICANCE STATEMENTThe CRISPR/Cas9-based mosaic technique, MAGIC, is a versatile tool for in vivo biological investigations. However, its reliance on DNA double-strand breaks (DSBs) can cause significant, unintended cell damage. Here we establish that Cas9-derived nickases, which create gentler single-strand nicks, are a superior alternative. We show that nickases safely induce genetic mosaics in Drosophila by avoiding this cellular toxicity. By systematically dissecting the process, we discovered principles of gRNA design that allow clone frequencies to be tuned for different experimental needs. This work provides a new mechanistic model for nick-induced genetic exchange, a high-fidelity "nickase-MAGIC" method, and a versatile toolkit for precision clonal analysis.

developmental biology↗

bioRxiv2022

Activating and repressing gene expression between chromosomes during stochastic fate specification

DNA elements act across long genomic distances to regulate gene expression in processes including enhancer-promoter interactions and imprinting. During the gene-regulatory phenomenon of transvection in Drosophila, DNA elements on one allele of a gene act between chromosomes to increase or decrease expression of another allele of the gene. Despite the discovery of transvection over 60 years ago, little is known about its biological role. Furthermore, how different cis regulatory DNA elements contribute to the activation or repression of transvection at distinct times during development is unclear. Here, we studied the stochastic expression of spineless (ss) in developing photoreceptors in the fly eye to understand gene activation and repression between chromosomes. We identified a biological role for transvection in regulating expression of naturally occurring ss alleles. We characterized CRISPR-engineered deletions of sequences across the ss locus and identified DNA elements required for activating and repressing transvection. We found that different enhancers participated in transvection at different times during development to promote gene expression and specify cell fates. Bringing a silencer element on a heterologous chromosome into proximity with the ss locus "reconstituted" the gene, leading to repression. Our studies show that transvection regulates gene expression via distinct DNA elements at specific timepoints in development, with implications for genome organization and architecture.

developmental biology↗

bioRxiv2025

Stochastic Modeling of Hematopoietic Stem Cell Dynamics

The study of hematopoietic stem cell (HSCs) maintenance and differentiation to supply the hematopoietic system presents unique challenges, given the complex regulation of the process and the difficulty in observing cellular interactions in the stem cell niche. Quantitative methods and tools have emerged as valuable mechanisms to address this issue; however, the stochasticity of HSCs presents significant challenges for mathematical modeling, especially when bridging the gap between theoretical models and experimental validation. In this work, we have built a flexible and user-friendly stochastic dynamical and spatial model for long-term HSCs (LT-HSCs) and short-term HSCs (ST-HSCs) that captures experimentally observed cellular variability and heterogeneity. Our model implements the behavior of LT-HSCs and ST-HSCs and predicts their homeostatic dynamics. Furthermore, our model can be modified to explore various biological scenarios, such as stress-induced perturbations mediated by apoptosis, and successfully implement these conditions. Finally, the model incorporates spatial dynamics, simulating cell behavior in a 2D environment by combining Brownian motion with spatially graded parameters. *Summary StatementThis study addresses the challenge of characterizing hematopoietic stem cell (HSC) dynamics by developing a flexible, user-friendly stochastic spatial model of long-term and short-term HSCs. The model captures observed cellular variability and heterogeneity, predicts homeostatic dynamics, can be adapted to simulate stress-induced perturbations like apoptosis, and incorporates a spatial component to analyze HSC movement within a bone marrow niche.

developmental biology↗

bioRxiv2017

Organoid And Tissue Patterning Through Phase Separation: Use Of A Vertex Model To Relate Dynamics Of Patterning To Underlying Biophysical Parameters

AO_SCPLOWBSTRACTC_SCPLOWExactly a century ago, DArcy Thompson set an agenda for understanding tissue development in terms of underlying biophysical, mathematically-tractable mechanisms. One such mechanism, discovered by Steinberg in the 1960s, is adhesion-mediated sorting of cell mixtures into homotypic groups. Interest in this phase separation mechanism has recently surged, partly because of its use to create synthetic biological patterning mechanisms and partly because it has been found to drive events critical to the formation of organoids from stem cells, making the process relevant to biotechnology as well as to basic development. Here, we construct quantitative model of patterning by phase separation, informed by laboratory data, and use it to explore the relationship between degree of adhesive difference and speed, type and extent of resultant patterning. Our results can be used three ways; to predict the outcome of mix-ing cells with known properties, to estimate the properties required to make some designed organoid system, or to estimate underlying cellular properties from observed behaviour.

developmental biology↗