Mediterranea Journal

[Cyndi Barca]

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Scripta Mediterranea was an annual refereed journal published by the Canadian Institute for Mediterranean Studies. The journal published articles studying all aspects of Mediterranean culture, past and present, with a special interest in interdisciplinary and cross-cultural research and issues.

Mediterranea. International Journal on the Transfer of Knowledge is an international journal focusing on various areas of knowledge transfer from Late Antiquity to the Early Modern period, covering the Middle East and the Mediterranean basin, and paying special attention to philological, philosophical, scientific, cultural and religious fields of research.

If there is one thing that characterizes the powerful process of knowledge exchange between the Near East and the Latin West it is the passion for knowledge and the discovery of its secrets that inspired scholars of the period. This led to long journeys and rich encounters, and the mutual exchange between cultures that have repercussions up to the present day.

Mediterranea is a project focused on combining efforts, by linking highly qualified research institutions with expertise in the field of transfer of knowledge within the different areas of study that will be addressed in the journal.

Copyright: 2018 Schmidt et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

In contrast to mammals, some invertebrates maintain highly potent stem cells lifelong. The stem cells of the planarian S. mediterranea, collectively termed neoblasts, constitute around 25% of all adult body cells and a substantial fraction of them are pluripotent [12]. This abundance of adult stem cells is the basis of the outstanding regenerative abilities and tissue plasticity of adult planarians (reviewed in [13]). The ability to modify neoblast dynamics by tissue amputation or feeding and to transplant them between individual animals, allowing full repopulation of a neoblast-depleted animal from a single donor neoblast, together with their amenability to RNA interference (RNAi) make planarians a powerful model organism for studying stem cell regulation in vivo. In a recent study, we identified distinct AS patterns of planarian neoblasts and of their differentiated progeny and showed that they are shaped by the RNA binding protein bruli, a CELF family member, and its antagonists of the MBNL family, respectively [14]. Depletion of these factors disturbs not only splicing patterns but also the balance of neoblast self-renewal and differentiation confirming their role in fate regulation. Similarly, a member of the Sm family of proteins, which associate with UsnRNAs forming small nuclear ribonucleoproteins (snRNPs) of the spliceosome, has been implicated in neoblast proliferation [15].

(A, B) UsnRNA variant expression, quantified by qRT-PCR, in animals depleted of neoblasts by h2b RNAi (blue) or lethal irradiation (green) and in cell populations isolated by FACS (red). Shown are the variants with significantly changed expression relative to the respective control. Results for other variants are shown in S3 Fig. Expression levels are the averages of three biological replicates, normalized to those of gapdh (error bars represent standard deviation; two-sided t-test, * p

Next, we employed the UsnRNA processing assays to RNA of X1 cells isolated by FACS from ints RNAi animals. For all UsnRNAs analyzed, we observed even higher average accumulation of the unprocessed forms in ints RNAi X1 cells than for RNA extracted from entire ints RNAi animals. Except for U4A2, all unprocessed forms were significantly increased in either ints9 or ints3 RNAi X1 cells, or both, compared to X1 cells of control RNAi animals (Fig 3B).

Neoblasts are strictly required for tissue turnover and regeneration in planarians [13]. Hence, an essential role of Integrator complex-mediated UsnRNA processing in neoblast function would be reflected by defects in these processes upon ints depletion. Monitoring intact ints RNAi animals for several weeks, we found that they developed head regression and ventral curling, phenotypes indicative of a neoblast defect [44], and died after a median survival time of 27 days (ints9 RNAi) or 33 days (ints3 RNAi) after the first dsRNA injection (Fig 4A, S5A Fig). Interestingly, amputated ints RNAi animals initiated regeneration but developed only a very small regeneration blastema and completely failed to regenerate photoreceptors or pharynges (Fig 4B). 7 days following amputation, when 1 or both eyes were already visible in the majority of control trunk and tail fragments, none of the ints RNAi fragments had regenerated eyes (Fig 4C). Like homeostatic RNAi animals, they eventually underwent ventral curling and died approximately 3 weeks after the first RNAi treatment.

To test whether ints depletion indeed affects the neoblast population, we performed flow cytometry of cell suspensions prepared from regenerating ints RNAi vs. control RNAi animals and observed a strong reduction of the X1 fraction at 7 days post amputation (dpa; 17 days of RNAi; Fig 4D). Interestingly, while we observed a strong effect of ints RNAi on UsnRNA processing in neoblasts already at 13 days of RNAi (Fig 3B), the proportion of X1 cells at that time-point was still comparable to that of control RNAi animals. This demonstrates that neoblasts are progressively lost during regeneration of ints RNAi animals and suggests that the disruption of UsnRNA processing is an indirect cause of this loss. To exclude that the neoblast loss is merely a consequence of the observed regeneration defect, we performed the same analysis on homeostatic ints RNAi animals, which showed a similar, progressive reduction of the neoblast fraction (Fig 4D). In line with a slower turnover of neoblasts during homeostasis, their proportion decreased at a slower rate in homeostatic compared to regenerating ints RNAi animals.

We next confirmed the observed neoblast loss using Fluorescent In Situ Hybridization (FISH) of smedwi-1 mRNA, the prototypical stem cell marker in planarians [44], which was strongly diminished in regenerating ints3 and ints9 RNAi animals at 20 days of RNAi (Fig 4E). Immunofluorescence co-staining of the SMEDWI-1 protein, which is maintained in the immediate progeny of neoblasts (smedwi-1-/SMEDWI-1+ cells), revealed that in both ints RNAi conditions these cells were strongly reduced, albeit to a lesser extent than neoblasts, likely as a consequence of defective neoblast maintenance [45] (Fig 4E). Depletion of a third putative Integrator complex subunit, Smed-ints11, pheno-copied the regeneration defects and neoblast loss observed upon ints3 and ints9 RNAi (S5B Fig).

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