Final Exam - Somite development Flashcards
(34 cards)
Somitogenesis
Concomitant (happening at the same time) with neurulation
Neurulation
- neural tube closure
- Concomitant (happening at the same time) with somitogenesis
- the process in vertebrates in which the future brain and spinal cord formed from the ectodermal neural plate
- as a result, largely of localized changes in cell shape, the neutral plate develops a central groove (the neural groove) with folds rising up on either side (neural folds)
- the folds eventually meet and fuse along the midline to form a tubular structure (the neural tube) that develops into the brain and spinal cord in birds and mammals
- the neural plate gives rise to the brain and spinal cord is formed from stem zone
Notochord
- organizing centre
- a transient stiff, rod-like cellular structure in vertebrate embryos that runs from head to tail and lies centrally beneath the neural tube
- it is derived from mesoderm and its cells eventually become incorporated into the vertebral column
Somites
- blocks of mesoderm that segment from the mesoderm on either side of the notochord
- they give rise to trunk and limb muscles, the vertebral column and ribs and the dermis
Somites
- paired segmented blocks of tissue that form on either side of the notochord
- Each somite develops in a precise temporal and spatial order
- blocks of mesoderm that segment from the mesoderm on either side of the notochord
- they give rise to trunk and limb muscles, the vertebral column and ribs and the dermis
- repeating pattern
- are paraxial (on either side of the notochord)
- Patterning of the somites provides much of the anterior-posterior organization of the vertebrate body plan
- In chick, a new somite forms every 90 minutes in a precise anterior to posterior direction
Paraxial mesoderm
- mesoderm lying on either side of the dorsal midline and which gives rise to the somites
- paraxial=on either side of the notochord
- Patterning of the somites provides much of the anterior-posterior organization of the vertebrate body plan
Node
in avian and mammalian embryos, the embryonic organizing center analogous to the Spemann-Mangold organizer of amphibians
- also known as Henson’s node in birds
- in plants, that part of the stem at which leaves and lateral buds form
Stem zone
-in avian and mammalian embryos, an arc of self-renewing epiblast cells on either side of the primitive streak immediately posterior to the regressing node that give rise to the trunk neural tube and the medial parts of the somites
Stem zone
- located in and around the node containing stem cells that divide and populate the pre-somitic mesoderm
- in avian and mammalian embryos, an arc of self-renewing epiblast cells on either side of the primitive streak immediately posterior to the regressing node that give rise to the trunk neural tube and the medial parts of the somites
Primary body formation
-in avian and mammalian embryos, the formation of the head and trunk from a stem cell population originating in the node and the adjacent arc of epiblast (called the stem zone)
Secondary body formation
- formation of the most posterior region of the body from the tailbud in mouse and chick embryos
- Tail is formed from a pool of stem cells called the tail bud stem cells
Tail bud stem cells
- Tail is formed from a pool of stem cells called the tail bud stem cells
- Originates from primitive streak and surrounding epiblast
Pre-somitic mesoderm
- the unsegmented mesoderm between the node (in chick and mouse) and the already formed somites
- it will form somites from its anterior end
- This region is produced from the stem cells surrounding the node (e.g. stem zone)
- The stem zone cells are self-renewing
- Daughters contribute to the pre-somitic mesoderm which is a region located between the last somite and the regressing node
- During node regression, the pre-somitic mesoderm region remains fairly constant in length
- Changes in cell shape and intercellular connections give rise to the blocks of somite tissue as they form from this pre-somitic mesoderm
Fgf and Wnt signalling
- Fgf and Wnt are signalling molecules
- expressed in node area (adjacent arc epiblast) of the mouse
- Maintain a pool of undifferentiated and proliferating (2 properties of stem cells) stem cells in the stem zone
- maintains the length of the pre-somitic mesoderm
- The highest levels of Fgf / Wnt are in the posterior with highest levels in the node
- patterns the anterior-posterior axis
- Fgf 8 mRNA is produced by the node and becomes degraded as cells move away from the node
- A Fgf 8 protein gradient is created without diffusion from a point source
- Wnt-3a, FGF Receptor 1
- Mutations in genes in the mouse result in truncation of the body axis
- A defect in maintaining the pre-somitic mesoderm
- A key transcriptional target of these pathways is expression of Brachyury
- In mouse, mutations in brachyury also cause axis truncation, probably due to the inability to regulate pre-somitic mesoderm
Fgf8 mRNA
- is produced by the node and becomes degraded as cells move away from the node
- A Fgf 8 protein gradient is created without diffusion from a point source
- part of Fgf/Wnt signalling
Wnt-3a
- Mutations in genes in the mouse result in truncation of the body axis
- A defect in maintaining the pre-somitic mesoderm
- part of Fgf/Wnt signalling
FGF Receptor 1 (FGFR1)
- Mutations in genes in the mouse result in truncation of the body axis
- A defect in maintaining the pre-somitic mesoderm
- part of Fgf/Wnt signalling
Brachyury
- a transcription factor
- important in somite development
- highly conserved
- driving specification from induction and patterning
- A key transcriptional target of the Wnt and Fgf signalling pathways is expression of Brachyury
- In mouse, mutations in brachyury also cause axis truncation
- Probably due to the inability to regulate pre-somitic mesoderm
autonomous
- somite development is autonomous and follows a pattern determined earlier in development
- describes any developmental process that can continue without a requirement for extracellular signals to be continuously present
- Experiment: Take pre-somatic mesoderm, flip it around, now inverted. Instead of taking on fate of its new position (more anterior), cells still become somites, in its normal place. Therefore, somite development is autonomous
- Temporal sequence of somite formation from the pre-somitic mesoderm is autonomous. Demonstrated through chick manipulation of the pre-somitic zone. Positional identity of pre-somitic mesoderm is already pre-determined. This suggests that the A-P axis is laid down earlier in development. Probably occurring during gastrulation (e.g. primitive streak formation and elongation).
opposing gradients of FGF and Retinoic Acid (RA)
- are expressed in opposing gradients
- retinoic acid antagonizes the Fgf/Wnt signalling pathways
- retinoic acid is expressed by the somites
- pattern pre-somitic mesoderm
- Retinoic acid expressed by the somites and diffuses towards the posterior
- This forms a retinoic acid gradient
- Opposing the Fgf 8 gradient is the retinoic acid gradient
- It antagonizes Fgf 8
- Functions to keep the pre-somitic mesoderm a consistent length (e.g. between the node and the last formed somite)
- Most embryos treated with excess retinoic acid have axis truncations
- The two gradients control the pre-somatic mesoderm and somite formation in a wavefront model
wavefront
- for somite formation
- the node migrates towards the posterior
- regulated by threshold of Fgf/Wnt signalling
- The position of the wavefront of somite formation is specified by a drop below a threshold level of Fgf/Wnt signalling
- when the threshold level of Fgf-8 and Wnt-3 signalling drops below the threshold, somite formation can occur
- Fgf-8 gradient moves with the node as it regresses that allows for the pre-somitic mesoderm to remain a constant length
- As cells pass the determination wavefront, they receive a second signal from an oscillating clock to form a somite at a precise time interval (e.g. 90 minutes)
Oscillating clock
- the clock is established by waves of c-hairy-1 expression
- allows for the precise timing of each somite
- A new somite forms every 90 minutes in the chick
- These blocks of tissue are determined by an internal clock
- This is represented by periodic cycles of gene expression
- c-hairy-1 is an example where its expression sweeps from the posterior to the anterior pre-somitic mesoderm with a period of 90 minutes
- After each wave, c-hairy-1 becomes restricted to the posterior region of each newly formed somite
- Each cycle, a given cell in the pre-somitic mesoderm experiences distinct phases when c-hairy-1 is or is not expressed
- More anterior somites have experienced fewer c-hairy waves/oscillations
- The control of c-hairy-1 expression is not well understood
- Notch-Delta signalling is necessary and is interacting with the Fgf-8/Wnt-3 gradient
- Over active or under active Notch signalling, there is a loss or irregularity of somite formation
- Fgf-8 gradient (high in node) interacts with the oscillating clock
- Timing and position of somite formation
c-hairy-1
- involved in somite formation
- part of oscillating clock
- c-hairy-1 is an example of a gene where its expression sweeps from the posterior to the anterior pre-somitic mesoderm with a period of 90 minutes = oscillating clock
- After each wave, c-hairy-1 becomes restricted to the posterior region of each newly formed somite
- Each cycle, a given cell in the pre-somitic mesoderm experiences distinct phases when c-hairy-1 is or is not expressed
- More anterior somites have experienced fewer c-hairy waves/oscillations
- The control of c-hairy-1 expression is not well understood
Delta-Notch signalling
- Notch-Delta signalling is necessary in the oscillating clock of somite formation
- Over active or under active Notch signalling, there is a loss or irregularity of somite formation
- important for timing and position of somite formation