HOMEWORK RUBRICS MECHANOTRANSDUCTION UNIT: HOMEWORK #1 (20 pts towards your grade)

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1 HOMEWORK RUBRICS MECHANOTRANSDUCTION UNIT: HOMEWORK #1 (20 pts towards your grade) 1. Mesenchymal stem cells (MSC) cultured on extracellular matrices with different stiffness exhibit diverse lineage commitment owing to the extracellular mechanical stimuli sensed by the cells. Based on the results of Engler et al. (paper discussed in class) of how cytoskeletal tension affects differentiation indicate how likely or unlikely the following experimental observations might be. In each case briefly indicate the reason for your answer (3 pts each/ 12 pts total) i) Expression of constitutive active Rho promotes osteogenesis Likely. Rho activation will result in increased cytoskeletal tension as it activates myosin II. This manipulation will mimic the mechanical cues provided by a stiff/rigid matrix that according to the paper promotes osteogenesis. ii) Inhibition of ROCK potentiates neurogenesis Likely. Inhibition of ROCK will result in decreased cytoskeletal tension as it will inhibit myosin II phosphorylation and activation. This manipulation will mimic the mechanical cues provided by a soft matrix that according to the paper promotes neurogenesis. iii) Inhibition of focal adhesion formation prevents cell spreading and promotes osteogenesis Unlikely. According to the paper increased cell spreading at high mechanical tension (sensed at focal adhesions) are necessary for osteogenesis. Therefore a manipulation that inhibits these processes will impair osteogenesis. iv) Treatment with latrunculin blocks MSC differentiation. Likely. Mecahnotransduction requires acto-myosin contraction. In addition, inhibition of actin polymerization will affect many processes including the ability of the cell to sense and respond to mechanical cues by changes cytoskeleton organization and morphology; as well as the activation of gene transcription (translocation to nucleus and/or retention in the cytoplasm of certain transcription factors by actin, regulation of transcription machinery, etc) 2. The data on the left was taken from a paper studying myotube differentiation in various microenvironments. The results (Panels A-B) show that myoblasts (C2C12 cells) plated directly on glass (lower myotube) or on top of a myotubes monolayer (upper); develop differently with only the cells on the upper layer acquiring a muscle-like organization (presence of actomyosin striations-panels C-D). The authors conclude that myotubes differentiate optimally on substrates with tissue-like stiffness. Based on these results: what would you anticipate would happen if the following experiments are carried out? In your answer please indicate what would you expect to see when you look for acto-myosin striations formation and clearly indicate the reason why. (4 pts each/ 8 pts total) i) Myoblasts are plated on top of a fibroblast layer (E= 2-5 kpa) From the data above, we can conclude that a fibroblast layer will act as a soft substrate. The lower elastic modulus of this substrate (2-5 kpa) compared to that of muscle (12-15 KPa) will not be optimal for the cells to differentiate and develop acto-myosin striations. ii) Myoblasts are transplanted to a fibrotic, dystrophic skeletal muscle lesion (E >10 MPa) The results of this experiment will likely mimic those shown above for cells grown directly on glass (infinitve stiffness). The fibrotic environment is too rigid and the myoblast will not develop acto-myosin striations. 1

2 MECHANOTRANSDUCTION UNIT: HOMEWORK #2 (20 pts. towards your grade) Background: Angiogenesis is controlled by soluble growth factors such as VEGF as well as by physical interaction between cells and the ECM. To gain insights into the mechanisms whereby mechanical signals integrate with other environmental cues during development researchers look at the role of small GTPase RhoA; a modulator of mechanical force and cell shape; and the expression of a transcription factors (TFII-I, GATA2) which regulate expression of VEGF receptor (VEGFR2) in endothelial cells. Stress-induced distortion of the endothelial cell cytoskeleton regulates Rho activity by controlling p190rhogap (an upstream inhibitor of RhoA). In addition p190rhogap can bind to TFII-I sequester it in the cytoplasm. Based on this information and the data presented in the panels below, please answer the following: Panel a: VEGFR2 and TFII-I mrna expression mrna expression following knockdown of TFII-I Panel b: Western blot following knockdown for TFII-I and/or p190rhogap. Panel d: VEGFR2 mrna expression following knockdown of p190rhogap and/or GATA2. 1. Does TFII-I control transcription of VEGFR2? Yes/No? Explain (4pts) Yes. From panel a,b we can observed that both mrna and protein levels for VEGFR2 are upregulated following knockdown of TFII-I knockdown with sirna; while overexpression of TFII-I using lentivirus (virus) downregulate VEGFR2 levels (panel a, only). So we can speculate that TFII-I is a negative regulator of VEGFR2 expression. 2. Does p190rhogap control transcription of TFII-I? Yes/No? Explain? (4 pts) From the data we can only conclude that p190rhogap is necessary for TFII-I protein expression as the treatment with sirna targeting p190rhogap, decreases the levels of TFII-I detected by western blot (panel b, third lane). Whether this regulation occurs at the level of transcription is not addressed directly by these experiments, but cannot be ruled out. Of note, it is stated that p190rhogap can sequester TFII-I in the cytoplasm, therefore p190rhogap effects on TFII-I expression might be partially mediated by a posttranscriptional mechanism. 3. Does p190rhogap controls transcription of GATA2? Yes/No? Explain (4 pts) The data in panel d only shows that GATA2 affects transcription of VEGFR2; specifically it shows that upregulation of VEGFR2 expression following p190rhogap knockdown; is reversed after GATA2 knockdown. Therefore, we can conclude that the upregulation of VEGFR2 mrna expression requires GATA2, but from these data; we cannot draw any conclusion as the control of p190rhogap on GATA2 transcription. 2

3 Endothelial cells were plated in matrices of different stiffness (150Pa, 4000 Pa and glass- panel c) and the levels of VEGFR2 protein expression were measured before and after knockdown of TFII-I, GATA2 and/ or p190rhogap. 3. Is VEGFR2 expression responsive to matrix elasticity? Yes/No? Explain (4 pts) Yes. VEGFR2 expression responds to differences in matrix elasticity. Panels c (top & bottom) show that the levels of VEGFR2 protein expression are up-regulated in stiffer matrices (4000 Pa, glass) compared to soft (150 Pa). In glass (infinite stiffness) VEGFR2 levels are higher than in the soft substrate (150 Pa) but still lower than at the optimal elastic modulus (4000 Pa) for vascular endothelial cells. 4. Based on the data how does RhoA activity promotes/inhibits VEGFR2 expression? In your answer clearly explain the combination of factor that promotes VEGFR2 expression and how p190rhogap affects that balance. (4 pts) Knockdown of p190rhogap, which negatively regulates RhoA will led to increased actomyosin contractility which mimics the condition of stiffer matrices, and therefore upregulates VEGFR2 expressions in soft matrices as shown in panel d (right). This up-regulation requires GATA2, as we can see that when this transcription factor is knockdown; the level of VEFGFR2 expression is down-regulated (with or without p190rhogap activity). From panel (d) we can conclude that TFII-I and GATA2 which are both transcription factors; affect VEGFR2 expression in opposite manner. TFII-I inhibits VEGFR2 expression; while GATA2 promotes it. This is demonstrated by the increase on VEGFR2 expression following knockdown of TFII-I with sirna; and by the decrease of VEGFR2 expression following GATA2 knockdown in soft matrices (150 Pa). Optimal conditions for VEGFR2 expression required downregulation of TFII-I and the presence of GATA2. These conditions appear to be favored in stiffer substrate (4000 Pa), which will promote a higher activation of Rho, such as that present in the absence of p190rhogap. Downregulation of p190rhogap will also result in reduction TFII-I levels as explained in question 2. 3

4 MECHANOTRANSDUCTION UNIT: HOMEWORK #3 (15 pts towards your grade) 1. Prolonged passage of normal cells on plastic Petri dishes often leads to spontaneous malignant transformation. How might altered mechanotransduction contribute to this phenomenon? (7.5 pts) The paper discussed in class demonstrated that increased ECM stiffness; similar to that reported for tumors in vivo; contributes to malignant transformation and tumor progression by promoting integrin clustering, Erk activation and increased Rho-mediated cytoskeletal tension. Cells growing on rigid substrates such as plastic Petri dishes for a prolonged period of time, might undergo similar activation of this integrin-rho-erk mechanosensitive axis, thus aiding to spontaneous malignant transformation. 2. Some malignant cells can grow and survive in suspension. How might altered mechanotransduction contribute to this phenomenon? (7.5 pts) Cells growing in suspension have lost regulation of the mechanosensitive axis, as they are not in contact with a substrate. Based on the model suggested by the paper we can speculate that these cells might harbor a mutation that mimics high cytoskeletal tension conditions, such those found in tumor/rigid matrices. Any mutation that leads to constitutive activation or loss of inhibition of the integrin/rho-erk pathway might contribute to this malignant phenotype. In the paper for example they used an integrin mutant that promotes integrin selfassociation (V737). Expression of this mutant allowed cells to display morphological changes and activation of focal adhesion in soft matrices to levels similar of those found in stiffer substrate. 4

5 MECHANOTRANSDUCTION UNIT: HOMEWORK #4 (20 points towards your grade) 1. The panels on the left show the results of in vitro rearrangements of zebrafish progenitors derived from two mutants: MZoep (red, ectoderm) and Cyclops (green,mesendoderm); after 2.5 hours (A,C) and 16 h (B,D). A,B correspond to the result of time-dependent sorting experiments; while C,D are the result of the engulfment experiment. E,F, show the results of mixing Cyclops (red) and Cyclops (green) cells. Briefly explain these results based on thermodynamic changes in aggregate cortical tension. (10 pts) The differential surface contraction (DSC) model proposes that, differences in cortical tension between cells and their environment rather than cell-cell adhesion drives cell sorting. In this experiment we can observe how cells from the ectoderm (red); which have the highest cortical tension aggregate in the center, surrounded (B) or engulfed (D) by the cells from the mesendoderm (green); which have a lower cortical tension. This arrangement reflects the tendency of the global aggregate area to decrease; driven by differences in aggregate surface tension. In E and F red and green cells derived from the mesendoderm, which have the same cortical tension, remain unsorted as expected. 2. The panels on the right show the sorting (A,B) and engulfment (C,D) assays of MZoep (red, ectoderm) and Cyclops (green, mesendoderm) cells. Panel A,C show the results using control non-treated cells; while panels B,D show the results after injection MZope cells with E- cadherin morpholinos. Briefly explain these results based on changes in aggregate surface tension. (10 pts) In this experiment what we observe is a reversal (B,D) of the original (A,C) self-sorting arrangement based on cortical tension: i.e. the ectoderm (red, higher cortical tension) surrounding the mesendoderm (green, lower cortical tension), following down-regulation E-cadherin, a molecule that mediates cell-cell adhesion. The explanation for this change in the context of the DSC model is related to how changes in cell-cell adhesion affect the overall aggregate surface tension. Aggregate surface tension is the result of both the cortical tension between cell-media interface (γ CM ); which is controlled by actomyosin tension only and is higher for ectoderm progenitors; and by cell-cell cortical tension (γ CC ), which is affected by both actomyosin tension minus the adhesion at the cell-cell interface. For cell-cortex tension to increase aggregate surface tension and to influence sorting as seen in panels A and C; it must increase the difference between γ CM and γ CC. Therefore, when cell adhesion is decreased as in panels B and D by reducing E-cadherin; the γ CC for ectoderm progenitors will increase, and the overall aggregate surface tension, which is the difference between γ CM and γ CC will decrease, causing the reversal of cell sorting. 5

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