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Kinetic studies of ionization of a tertiary halide in a protic solvent suggest that only the substrate participates in the rate-determining step (slow step). The nucleophile is involved only after the slowest step. The SN1 reaction takes place in a multiple-step mechanism. 
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Related Experiment Video

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Ex vivo Mechanical Loading of Tendon
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Excessive mechanical loading promotes osteoarthritis through the gremlin-1-NF-κB pathway.

Song Ho Chang1, Daisuke Mori1,2, Hiroshi Kobayashi1

  • 1Sensory and Motor System Medicine, Graduate School of Medicine, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo, 113-8655, Japan.

Nature Communications
|March 31, 2019
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Summary

Mechanical loading induces gremlin-1 in cartilage, promoting osteoarthritis by activating inflammatory pathways and inhibiting cartilage repair. Targeting gremlin-1 may offer a novel therapeutic strategy for osteoarthritis.

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Area of Science:

  • Biochemistry
  • Cell Biology
  • Orthopedics

Background:

  • Osteoarthritis pathogenesis is linked to excessive mechanical loading of articular cartilage.
  • Chondrocytes are key cells in cartilage homeostasis and response to mechanical stress.

Purpose of the Study:

  • To identify novel molecular factors involved in the pathogenesis of osteoarthritis induced by mechanical loading.
  • To elucidate the role of gremlin-1 in chondrocyte response to mechanical stress and its contribution to osteoarthritis.

Main Methods:

  • Investigated gremlin-1 expression in chondrocytes under mechanical loading (cyclic strain, hydrostatic pressure).
  • Utilized mouse models with intra-articular gremlin-1 antibody administration or chondrocyte-specific Gremlin-1 deletion.
  • Assessed activation of nuclear factor-κB (NF-κB) signaling and reactive oxygen species (ROS) production.
  • Examined the effect of gremlin-1 on anabolic gene expression (Sox9, Col2a1, Acan).

Main Results:

  • Gremlin-1 is induced by mechanical loading in chondrocytes, particularly in deeper cartilage layers.
  • Gremlin-1 activates NF-κB signaling, leading to increased catabolic enzyme production.
  • Inhibition of gremlin-1 (antibody or genetic deletion) decelerated osteoarthritis development in mice.
  • Mechanical loading-induced ROS production, mediated by ras-related C3 botulinum toxin substrate 1, activates RelA/p65, which upregulates Gremlin-1.
  • Gremlin-1 antagonizes bone morphogenetic protein-induced expression of anabolic genes.

Conclusions:

  • Gremlin-1 is a critical mediator of cartilage degeneration in response to excessive mechanical loading.
  • Gremlin-1 promotes an osteoarthritis-promoting catabolic environment by activating NF-κB and inhibiting anabolic pathways.
  • Targeting gremlin-1 presents a potential therapeutic avenue for mitigating osteoarthritis progression.