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Determine should your aim is to buy wide distribution and maximum exposure. The mean vector copy number VCN of the bulk culture first fluctuated around one copy per diploid genome Fig. From passage 5 to 12, the VCN increased and followed the kinetics of the upcoming white clone. This increase can be explained by the reduction in vector-negative cells together with the clonal selection of the white cells with a VCN of at least 3.
This loss of barcode diversity became even more obvious when we analyzed the dynamics of individual barcodes. Overall, clonal complexity can be assessed by using the Shannon index Fig. In a very complex sample, we observed a high amount of different barcodes with low read counts and consequently also a high Shannon index.
In samples with a loss in clonal diversity, the Shannon index dropped as the number of individual barcodes became fewer and the read counts for some barcodes became dominant. This general loss of clonal complexity in the expanded MSC-EM cultures is further demonstrated by using stacked frequency charts Fig.
The contribution of each individual barcode to all sequences over time is displayed as a colored area. The first severe loss of diversity was observed within the first three passages already. Another major loss of barcodes occurred after passage 5, where mainly three mCherry, two Cerulean, and one Venus barcode became dominant. In flow cytometry at the same time, mainly fluorescence-negative, purple, red, and some white cells were detected.
Consequently, three mCherry barcodes were likely to be found in the red, purple, and white cell population. The two Cerulean barcodes were assumed to be a tag of the purple and white cell population, and the Venus barcode was found only in the white cell fraction. At passage 9, one of the mCherry barcodes started to disappear, at the same time when the red cell population was not detectable by flow cytometry anymore. After passage 12, only three dominant barcodes were found in the sequencing result, one of each color.
These results highlight the opportunity to follow global culture dynamics in flow cytometry and to assign unique barcodes to individual cell fractions. Explant cultures from independent UCPs showed similar decreases in barcode variability supplemental online Fig. The first loss of clonal complexity is usually found very early after culture initiation, when all cells lacking an MSC phenotype disappear from the culture.
A second drop of barcode diversity can be observed during expansion, when specific clones become dominant. This change in clonal complexity was also recapitulated by the proliferation rate of the mass culture, which changed depending on the clonal composition over time supplemental online Fig. A : Flow cytometric analysis of the clonal development. The graph displays the percentage of fluorescence-positive cells, and the color of the symbol indicates the color of the respective cell population black indicates fluorescent-negative cells.
The development of the mass culture during the first 12 passages P1. B : Vector copy number analysis of mass cultures by quantitative polymerase chain reaction. C : Shannon index describing the diversity of the barcodes in the mass culture. D : Area plot of the barcode contribution. Each area represents an individual barcode from one of the vector constructs different shades of the colors red, blue, and green refer to the RGB construct the barcode originated from.
To answer this question, we compared consecutive primary P1. We observed only a limited fluctuation in the color composition until passage P1. Further analysis by barcode sequencing revealed a drastic restriction of diversity to only two mCherry barcodes, which were identical between the primary and secondary MSC-EMs from UCP 2 Fig.
This selection of identical barcodes from consecutive MSC-EMs suggests a long-lasting, but limited, fraction of progenitor cells, which continuously give rise to new MSC cultures. Colors of the symbols and areas indicate fluorescent proteins coded by the vectors.
A : Flow cytometric analysis of the first 13 passages P1. B : Flow cytometric analysis of the first 3 passages P2. C : Flow cytometry analysis of the first 5 passages P1. D : Flow cytometry analysis of the first 3 passages P2. E : Area plot of the barcode distribution of the first 5 passages P1. F : Area plot of the barcode contribution of the first 3 passages P2. Finally, we investigated whether different clones possess different properties.
As a proof of principle, we chose a Matrigel assay in which MSCs can form tube-like structures. This tube formation assay was previously described by Gomes et al. A freshly initiated MSC-EM was expanded to a six-well size before the cells were sorted by fluorescent color.
Technical replicates of three resulting populations green, yellow, and negative were cultivated for 1 week in an endothelial growth medium EGM Afterward, cells were seeded on Matrigel, where they normally form tube-like structures. Four hours after seeding, tube formation was analyzed by microscopy. All replicates of the fluorescent-positive cells had a strongly reduced tube formation potential, whereas the negatively sorted population showed normal tube-like structures Fig.
This shows that MSC mass cultures are composed of cells with different functional properties. Tube formation assay with different fluorescence sorted clones. A freshly initiated mesenchymal stem cell explant monolayer was first expanded, before cells were sorted by the present colors green, yellow, and negative. A tube formation assay was performed in triplicate with the different sorted populations. Depicted is microscopic fluorescence overlay after 4 hours.
The yellow and green population had a strongly reduced tube formation capability compared with cells from the fluorescence negative population. Despite their broad application in clinical trials for the treatment of a variety of diseases, the definition of unique surface markers to directly isolated MSCs has not been achieved so far [ 25 ].
Regardless of the tissue source, MSC monolayer cultures have to be expanded to yield clinically relevant cell numbers [ 26 ]. During the first 2 weeks of culture establishment, the cell population is quite heterogeneous and only develops the typical homogenous phenotype during expansion.
How many cells are able or necessary to initiate such a culture, and how the clonal composition changes over time, are still unanswered questions [ 27 ]. RGB barcode-marking technology is a powerful tool to analyze clonal development on different empirical levels [ 18 ]. In our current study, we used this technique to investigate umbilical cord tissue as a constant source of MSC cultures and observed a general clonal selection pressure during the inevitable expansion phase, already shortly after initiation.
Previously, Otte et al. Whereas monolayer cultures alone showed signs of senescence after 6 weeks, separately cultured UCPs kept initiating new MSC-EMs over months, raising the question of a stem cell-preserving niche in the tissue piece. This would represent an advantage over other MSC sources, where this microenvironment is normally destroyed by enzymatic digestion of the tissue during isolation.
MSCs or their direct progenitors have to proliferate inside the UCP and migrate toward the periphery of the pieces, to initiate monolayer cultures. We applied Ki immunohistochemistry to identify actively dividing cells in freshly isolated or cultured UCP.
In contrast, cultivated pieces showed dividing cells at the periphery and around the collapsed umbilical vessel wall. Several groups suggested a pericytic origin of MSC [ 10 , 28 ], which would be in line with our observation that nonvascularized tissue pieces lost their MSC-EM initiation potential over time. Because we did not know the exact localization of these cells, we tried different time points and methods of lentiviral transduction. Additionally, fresh pieces were either cut in half to increase the surface, mildly digested to promote access of virus to the inner cells, or injected with viral supernatant to reach cells deeper in the UCP.
Enzymatic digestion shortly before the time of transduction turned out to be incompatible with gene transfer, further cultivation of intact UCP, and MSC-EM initiation. This was likely because of the fact that remaining enzymes after incomplete removal also negatively affect the viral particles. Simple addition of vector supernatant to UCP resulted in reproducible but mainly superficial gene marking.
Injection into pieces yielded transduced spots together with labeled cells at the periphery, which could be explained by backflow of viral supernatant from the injection canal into the medium. These results imply that we did reach a population of MSC-initiating cells, but failed to access another population located in deeper tissue layers, possibly around the vessel structures. In line with this finding, analysis of the barcode diversity of whole pieces did not reveal a significant difference in the labeling efficiency between different transduction methods.
Shortly after transduction of fresh pieces, the total number of labeled cells in fluorescence microcopy was low in comparison with later time points, when the same pieces were analyzed again. In contrast, UCP, which had been transduced after 5 weeks in culture, showed a high amount of initially labeled cells, but no further increase in gene marking. However, the number of fluorescence-positive MSC-EMs growing out from the pieces was not different between fresh and cultivated pieces.
This observation is important, because it suggests that we transduced mostly short-lived cells in the older pieces and that there is only a limited pool of MSC-I in the umbilical cord tissue. This assumption is further supported by the observation of a lower barcode variability in secondary and tertiary MSC-EMs from the same piece.
Initially starting with hundreds of different barcodes and often a diverse color composition, only a few barcodes became dominant over time as fluorescent markers showed a reduced complexity. In line with our observations, Schellenberg et al. Additionally, we observed a change of proliferation rate in the mass cultures at time points of barcode diversity reduction.
This happened predominantly after passage 5—9 and potentially displayed the growth rate of those clones, which were about to become dominant. Because of the limited transduction efficiency of whole tissue pieces and possibly the location of MSC-I cells, we also observed a high contribution of unlabeled cells in our MSC-EM cultures. Sometimes the transduced cells diminished, when unlabeled cells became selected, whereas at other times the marked cells dominated over time.
This observation argues against the possibility that the clonal selection was generally triggered by insertional mutagenesis because of the lentiviral vector transduction. Generally, we observed only a minor overlap, suggesting the existence of several MSC-I in the tissue pieces. However, in some explant cultures, dominant clones carried identical barcodes.
The existence of subpopulations within MSC cultures has also been described by other groups according to differences in surface markers, whole genome sequencing, and functional characteristics [ 29 — 33 ]. The best resolution so far was achieved with single colony-derived strains and the linkage to functional properties by Sworder et al.
Finally, we analyzed whether different clones have different functional properties. We used a tube formation assay with cells sorted by color. Although the three different populations originated from the same MSC-EM, only one population efficiently formed tubes in all replicates, highlighting different capabilities among clones. Despite several functional in vitro tests, like trilineage differentiation or the tube formation assay, the predictive value in terms of clinical translation remains doubtful.
We believe the tube-like structure formation to be a specialized surrogate assay in terms of functionality. Because the secretion of various instructive cytokines by MSC is believed to be one of major factors accounting for their in vivo regenerative potential, future studies should focus on differences in the expression of these factors by specific clones. We observed clonal selection of UCP-derived MSC-EMs, which can be explained by complex proliferation dynamics and a small set of long-term proliferating progenitors.
Furthermore, we could also demonstrate functional differences among clones. In particular, temporary contribution of clones to the expanded culture might explain sources of variation seen in MSC in vitro experiments or potentially even clinical trials.
Our study also showed the suitability of UCP transduction with lentiviral vectors coding for fluorescent proteins and genetic barcodes as a tool for clonal development analysis. Future studies should aim to analyze the consequences of the clonal dynamics on functional difference between MSC cultures and the possibility to manipulate this outcome by the choice of different culture conditions.
This will help to identify conditions for the generation of clinically applicable MSCs with less heterogeneity and better reproducibility. Selich and M. Schambach: conception and design, financial support, data analysis and interpretation, manuscript writing, final approval of manuscript. Read article at publisher's site DOI : Stem Cell Res Ther , 12 1 :7, 06 Jan J Clin Med , 9 9 , 10 Sep Hass R.
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Significance Mesenchymal stem cells MSCs can easily be obtained from various adult or embryonal tissues and are frequently used in clinical trials. Free full text. Stem Cells Transl Med. Published online Mar PMID: Author information Article notes Copyright and License information Disclaimer.
Corresponding author. Correspondence: Michael Rothe, Ph. Telephone: ; E-Mail: ed. Received Jul 28; Accepted Jan This article has been cited by other articles in PMC. Go to:. Open in a separate window. Figure 1. Virus Production and Transduction Viral supernatants were produced in a four-plasmid split packaging system as described before [ 18 , 20 ].
Tube Formation Assay The tube formation assay was performed as described previously [ 23 ]. Data Processing and Statistics The preprocessing of the sequencing data was performed with customized Padre 0. Figure 2. Figure 3. Figure 4. Clonal Selection During Expansion of MSC Monolayer Cultures We further investigated whether monolayer cultures, once initiated, remained stable in their clonal repertoire, or if an expansion would lead to selection of certain clones.
Figure 5. Figure 6. Figure 7. Supplemental Data: Click here to view. Heterotopic of bone marrow. Analysis of precursor cells for osteogenic and hematopoietic tissues. Boxall SA, Jones E. Markers for characterization of bone marrow multipotential stromal cells. Stem Cells Int. Mesenchymal stem or stromal cells: a review of clinical applications and manufacturing practices. Immunological aspects of allogeneic mesenchymal stem cell therapies.
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|Richard bettinger art||This shows that MSC mass cultures are composed of cells with different functional properties. They also give Perception in the wertheim bettingen kindergarten readiness and ambitions of the businesses creating these merchandise, ta18 bitcoins with avenues of troubleshooting and repair service. Mesenchymal stem cells: A perspective from in vitro cultures to in vivo migration and niches. The mean vector copy number VCN of the bulk culture first fluctuated around one copy per diploid genome Fig. Further information available here. F : Area plot of the barcode contribution of the first 3 passages P2. Figure 5.|
|Gta sa inside track betting||Send Email. That is the way you identify the price of your ebook. Figure 3. Despite a homogenous appearance under the microscope, MSC cultures undergo massive clonal selection over time. This is about mathematical questions from highschool. Sometimes the transduced cells diminished, when unlabeled cells became selected, whereas at other times the marked cells dominated over time.|
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