1. What are the three general areas of the developing vertebrate limb? 2. What embryonic regions contribute to the developing limb bud?

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1 Study Questions - Lecture 17 & What are the three general areas of the developing vertebrate limb? The three general areas of the developing vertebrate limb are the proximal stylopod, zeugopod, and the autopod. 2. What embryonic regions contribute to the developing limb bud? The embryonic regions that contribute to the developing limb bud are the lateral plate mesoderm and from the myotome responsible for limb muscle precursor cells. 3. What type of gene expression (gene family name) is associated with the A-P positioning of developing limb buds? The genes that are involved in the A-P positioning of the developing limb bud are Hox genes along the axes. 4. Describe the expression patterns of FGF8 and FGF10 in the developing limb bud and during limb outgrowth. Outline the model for the signalling pathway by which these expression patterns are consolidated and maintained. FGF10 is initially expressed throughout the lateral plate mesoderm. The expression pattern is then restricted to the sites of limb bud formation by Wnts specific for the fore limb and hind limb area. This localized expression in the lateral plate mesoderm then regulates FGF8 and Wnt3a in the ectoderm of limb buds. These are all necessary for limb outgrowth.

2 5. What genes are differentially expressed in the developing fore limb vs. hind limb? The developing fore limb will express Tbx5 while the hind limb will express Tbx4. 6. What is the apical ectodermal ridge and what is its function? The apical ectodermal ridge runs along the distal margin of the limb bud. It functions as a major signaling center with 3 major roles: -maintaining plasticity of underlying mesenchyme so that proliferation will lead to proximal to distal limb growth -maintaining expression of molecules that generate the A-P axis -interacting with proteins specifying A-P and D-V axes so that each cell can differentiate 7. What group of cells in the developing limb bud specify the character (fore vs. hind) of the developing limb? The group of cells that specify the character of the developing limb is known as the progress zone.

3 8. How was it determined that the specification of proximal vs. distal structure in the developing limb is not generated by signals from the AER? It was proven that the AER did not carry all the information as these were proven to be false: -older AERs combined with younger mesoderm would make distal structures in proximal positions -younger AER combined with older mesoderm would make repeated structures 9. What is the progress zone model of proximal/distal specification in the developing limb? What evidence suggests that this model is not correct? In the progress zone model, it was thought that the amount of time the mesodermal cell spends dividing in the progress zone determined how distal it s specification was. So if it had more time, it would result in a more distal part. However, when FGF was knocked out in mice, distal elements formed normally while proximal ones did not. 10. What is the early allocation and progenitor expansion model of the proximal/distal specification in the developing limb? What experimental evidence supports this model? In the early allocation and progenitor expansion model, it was thought that early cells are already specified and division just increases the number of discrete cell populations. Support came from experimental removal of the AER, zones of distal structures could be destroyed and lead to the formation of only proximal structures. 11. What is the developmental function of the zone of polarizing activity? The developmental function of the zone of polarizing activity is to specify A-P axis of a limb. 12. What important paracrine factor is expressed specifically in the ZPA? How was it determined that this gene is sufficient and necessary for ZPA function? The important paracrine factor expressed in the ZPA is Shh. It was determined that Shh was sufficient and necessary for ZPA function by placing a cell pellet secreting Shh onto the presumptive anterior region while normal ZPA was present at the posterior and forming a limb with two axes. 13. What does my cat have extra toes? The cat has extra toes because of a gain of function mutation in Shh. 14. What evidence suggests that interdigitary tissue is also involved in digit specification?

4 Experiments where interdigitary tissue was removed would lead to duplications of certain digits. For example, if the tissue between digit 2 and 3 were removed, both of them would now result in digit 2 identity. 15. What signalling molecule associated with specification of dorsal vs. ventral in the vertebrate limb? How is this related to nail-patella syndrome in humans? The signalling molecule associated with specification of D-V axis in the vertebrate limb is Wnt which is expressed in the dorsal ectoderm of limb buds. This signalling activates the transcription factor Lim1 in dorsal mesenchyme which is fuccient and necessary for dorsal fate. In humans, mutants of Lim1 will lack dorsal limb structures. This causes nail-patella syndrome as dorsal structures will not form correctly. 16. What type of signalling operates at the junction of dorsal and ventral mesoderm in the developing limb to establish the position of AER? Notch signalling is important at the junction of dorsal and ventral mesdoderm in developing limb to establish the position of AER. It will relay the dorsal ventral signals to the cells forming the AER> In the dorsal region, Wnt is expressed which activates Lmx1 leading to radical fringe expression. However, in the ventral region engrailed is expressed and it downregulates both Wnt and radical fringe. The AER will form at this junction between dorsal and ventral. 17. How do duck s feet develop the webbing whereas the chicken s does not? How could a researcher experimentally make chickens with webbed feet? Apoptosis of interdigitary tissue is directed by the presence of BMPs. In the duck, webbing develops as interdigitary tissue contains a BMP antagonist known as gremlin. Chickens do not have anything similar to this. A researcher could create a chicken with webbed feet by introducing Gremlin or BMP antagonists into interdigitary regions. This will prevent apoptosis of the tissue, leading to webbed chicken feet. 18. BMP signalling may either induce apoptosis or induce the formation of bone and cartilage? How is thought that such different outcomes result from the same signalling molecule? Such radical differences in outcomes are a result of other factors interacting with BMP. When BMP is in the presence of Wnt and B-catenin, it will induce cartilage formation. However, if BMP is now in the presence of FGF, it will lead to the activation of Dkk-1 and apoptosis will follow.

5 Study Questions - Lecture What are germ cells? Germ cells are different than somatic cells in that they are responsible for reproductive function. They will undergo meiotic cell divisions to generate gametes. 2. What type of specification is involved in mammalian germ cell specification? What type of specification is seen in Xenopus, Zebrafish, Drosophila, and C. elegans? In the specification of germ cells in mammals, conditional or regulative specification is used. This occurs when cells are specified in the epiblast through inductive signalling from the extraembryonic ectoderm and endoderm. However, in Xenopus, Zebrafish, Drosophila, and C. elegans, the germ cells are determined by autonomous specification through invariable cleavage. This is a result of localized germ cell determinants in a distinct region of the oocyte cytoplasm being segregated into the germ cell lineage through cleavage. 3. What is the germ plasm? The germ plasm is a special area of cytoplasm within the oocyte that houses these protein and mrna determinants. 4. Why do you suppose primordial germ cells in different systems are all transcriptionally quiescent? Primordial germ cells in different systems are transcriptionally quiescent as they must remain undifferentiated until the gonads and other structures have begun to develop. Active transcription will lead to proper migration, and further differentiation will occur as the organism approaches a reproductive stage. 5. Outline experimental evidence from Drosophila that suggest that germ line determinants are composed mostly of RNA. In an experiment, an untreated embryo s pole plasm was isolated. The pole plasm was then fractionated to produce a protein portion and a RNA portion. These samples were introduced into two embryos without pole plasm and it was discovered that the pole plasm protein was unable to induce pole cell formation while the RNA was able to produce a fertile adult. 6. In Drosophila, why would a mutation that is specific to germ line determinant (required for germ line specification or germ line viability) be identified from a line that produces grandchildless females?

6 A mutation that is specific to germ line determinant could be easily identified from a line that produces grandchildless females as these individuals would be sterile. Since these mutations do not disrupt the development of the organism, mutations can be identified. 7. What are the identities of some of the pole plasm determinants in Drosophila? Which of these have also been identified to specify the germ line in frogs, fish, and even in mammals? In Drosophila, pole plasm determinants include: -germ cell-less mrna (necessary for gene silencing in pole cells) -polar granule component mrna (inhibits transcription by preventing phosphorylation of RNA Pol II) -posterior group determinants (such as Oskar and Nanos that inhibit translation of certain messages) -RNA binding proteins (e.g. Vasa) -mitochondrial ribosomes In frogs and fish, there are also granules, mitochondrial factors, different mrnas including homologues of nanos and vasa. 8. Germ cells tend to be specified in a location that is remote from major sites of signalling activity, then they migrate to populate the forming gonad. What type of cues are used to guide primordial germ cells to their destination? Compare Drosophila to fish and frogs. Primordial germ cells are guided by repulsive and attractive interactions. This includes wumen and HMG-CoA reductase respectively in Drosophila. In Zebrafish, chemoattraction of PGCs occurs when the developing gonad secretes Sdf1. The PGCs express a receptor for Sdf1 called CXCR4. In Xenopus, migration is dependent on fibronectin in the extracellular matrix, but it also contains Sdf1 attractant and CXCR4 receptor on the PGCs. 9. Mammalian primordial germ cells migrate with a group of cells that are not PGCs. What is the function of these other cells? PGCs in mammals migrate alongside cells secreting stem cell factor (SCF). These cells could potentially be like a travelling stem cell niche, keeping the PGCs in a relatively undifferentiated form.

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