Imaging and manipulating the segmentation clock
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Cellular and Molecular Life Sciences
REVIEW
Imaging and manipulating the segmentation clock Kumiko Yoshioka‑Kobayashi1,2,5 · Ryoichiro Kageyama1,2,3,4 Received: 24 June 2020 / Revised: 21 August 2020 / Accepted: 22 September 2020 © Springer Nature Switzerland AG 2020
Abstract During embryogenesis, the processes that control how cells differentiate and interact to form particular tissues and organs with precise timing and shape are of fundamental importance. One prominent example of such processes is vertebrate somitogenesis, which is governed by a molecular oscillator called the segmentation clock. The segmentation clock system is initiated in the presomitic mesoderm in which a set of genes and signaling pathways exhibit coordinated spatiotemporal dynamics to establish regularly spaced boundaries along the body axis; these boundaries provide a blueprint for the development of segment-like structures such as spines and skeletal muscles. The highly complex and dynamic nature of this in vivo event and the design principles and their regulation in both normal and abnormal embryogenesis are not fully understood. Recently, live-imaging has been used to quantitatively analyze the dynamics of selected components of the circuit, particularly in combination with well-designed experiments to perturb the system. Here, we review recent progress from studies using live imaging and manipulation, including attempts to recapitulate the segmentation clock in vitro. In combination with mathematical modeling, these techniques have become essential for disclosing novel aspects of the clock. Keywords Fluorescent reporter · Luminescent reporter · Notch signaling · Oscillation · Somite · Synchronicity
Introduction The animal body is formed by a series of repetitive structures along the head-to-tail axis, such as the vertebrate spine and ribs. In vertebrates, these “segment-like” structures are derived from somites, an embryonic tissue. Somites differentiate from the presomitic mesoderm (PSM), a proliferative progenitor population located at the caudal end of the embryo during the mid-gestation period. During this * Kumiko Yoshioka‑Kobayashi yoshioka.kumiko.4r@kyoto‑u.ac.jp * Ryoichiro Kageyama [email protected]‑u.ac.jp 1
Institute for Frontier Life and Medical Sciences, Kyoto University, Kyoto 606‑8507, Japan
2
Graduate School of Medicine, Kyoto University, Kyoto 606‑8501, Japan
3
Graduate School of Biostudies, Kyoto University, Kyoto 606‑8501, Japan
4
Institute for Integrated Cell-Material Sciences, Kyoto University, Kyoto 606‑8501, Japan
5
Institute for the Advanced Study of Human Biology (ASHBi), Kyoto University, Kyoto 606‑8501, Japan
differentiation process, a group of cells at the anterior end of the PSM are “pinched off” and differentiate into epithelialized somites, which are distributed on both sides of the neural tube. The development of these epithelialized somites occurs sequentially from head to tail, with regular intervals and periodicity. The periodicity is species-specific: 2 h in mice and 4–6
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