Studies on Visceral Regeneration in Sea-Stars. doi:10.1002/jez.b.22899, Kondo, M., and Akasaka, K. (2010). 4 (5), 170140. doi:10.1098/rsos.170140, Ferrario, C., Czarkwiani, A., Dylus, D. V., Piovani, L., Carnevali, M. D. C., Sugni, M., et al. Pharmacological studies in brittle star arm regeneration and sea cucumber intestinal regeneration. Sci. (2020) on the effect of Wnt inhibitors (iCRT14) and activators (LiCl) is determined in the size of the regenerating gut rudiment when compared to those of vehicle treated controls. Many of the genes identified in this study were not mentioned in previous reports, probably because of the differences in their analysis approach. Dev. In A. japonicus, the tissues include the nerve ring, including the calcareous structure, as well as the associate muscles of animals that were 1) eviscerating animals, 2) 3-h post evisceration and 3) non-induced to eviscerate (normal). Scientific Rep. 11 (1), 114. These studies provide for the study of candidate genes, in particular those that have been previously associated with regenerative or developmental processes, to be searched in the regenerating tissues of echinoderms. Echinoderms, which thanks to their spectacular regenerative capabilities were the favourite models of the pioneer regenerationists of the 19th and early 20th centuries, after a long period of unexplainable neglect, were reproposed to our attention by a series of recent papers (for review see Candia Carnevali and Bonasoro 2001b; Candia Carnevali et al. Other genes upregulated in the in vitro regenerating explants, are also involved with proliferation, migration, and differentiation, such as frizzled, tetraspanin, and selenoprotein W, respectively. In one of the earlier studies, Afuni, a member of the BMP family, was shown to be expressed in A. filiformis regenerating arm (Bannister et al., 2005). doi:10.1186/s12915-020-00937-7, Purushothaman, S., Saxena, S., Meghah, V., Swamy, C. V. B., Ortega-Martinez, O., Dupont, S., et al. Do echinoderms have a complete digestive tract. Similarly, a study on the Iberian ribbed newt, Pleurodeles waltl, also showed that specific TEs, including Gypsy elements were upregulated during limb regeneration and that these appear to have been expanded in this species genome (Elewa et al., 2017). Developmental Dyn. Pharmacological inhibition of the Notch pathway was done in sea urchins using DAPT (Reinardy et al., 2015). Noteworthy, they also identified the translation regulation pathways as being associated with the brittle star early arm regeneration. Phylum Echinodermata | Biology for Majors II - Lumen Learning Among the most important are the experiments by (Patruno et al., 2002; Patruno et al., 2003) documenting the expression of putative members of the Transforming Growth Factor (TGF) and Bone Morphogenetic Protein (BMP) families in cells of the regenerating arm of the crinoid Antedon bifida. This makes further exploratory molecular studies on crinoids, echinoids, and asteroids essential in order to obtain a broader overview of the molecules that act during regeneration. Most people know if a sea star loses one or more arms, the appendage(s) will eventually grow back. The coelomate bilateria 4, 1763. When animals are treated with an inhibitor of the Wnt pathway, it can be modulating the expression (and thus, the role) of all these genes simultaneously. 407 (2), 315322. In fact, Sox17 is one of the identified genes in the transcriptome profile of the polian vesicles of eviscerated sea cucumbers, where production of coelomocytes occurs (Shi et al., 2020). (2015). doi:10.1371/journal.pone.0232981, Mashanov, V. S., and Garca-Arrars, J. E. (2011). No use, distribution or reproduction is permitted which does not comply with these terms. Here we review the available data to determine the genes and signaling pathways that have been proposed to be involved in regenerative processes. B., and Sansom, O. J. Genet. Ophiuroids with arms branched in a complex manner may feed in a way similar to that of the crinoids. Furthermore, DSP-1, another development-related gene involved in Hox regulation, was also found to be upregulated in the distal portion of the arm explant. Collectively, these reveal the grand requirement of resources and mechanisms for the regeneration of the lost tissue. In contrast, transforming growth factor beta-2 (Tgfb2), AT-rich interaction domain 1b (ARID1b), SIPA, and N-ethylmaleimide-sensitive fusion protein (NSF) were at the top of the downregulated transcripts list. The subphylum Stelleroidea contains the two classes of sea stars. The Eupentacta Fraudatrix Transcriptome Provides Insights into Regulation of Cell Transdifferentiation. J. Dev. Biol. Even though echinoderms are closely related to chordates, they are little known in the biomedical field, and therefore their uses to study pharmacological effects on muscle formation and/or regeneration have been extremely limited. Many of the ECM related genes (i.e., collagen, tenascin, laminin, and echinonectin), showed upregulation at all regenerating stages, while the MMP genes returned to normal expression at 14-dpe. Certainly, although the analysis of E. fraudatrix intestinal regeneration gene profiles had a specific focus on transdifferentiation processes, the identified transcription factors might also represent additional processes occurring simultaneously. Microscopic Overview of Crinoid Regeneration. Regeneration of the radial nerve has also been studied in H. glaberrima. All three caused a decrease in the size of the rudiment, suggesting that MMP activity is necessary for normal regeneration of the intestine. Cel Tissue Res. In fact, they are highlighted in Fig. Echinoderms. This is probably the limiting factor, at present, to advance the molecular analyses of regeneration. brittle star, also called serpent star, any of the 2,100 living species of marine invertebrates constituting the subclass Ophiuroidea (phylum Echinodermata). Regeneration in Echinoderms: Molecular Advancements Studies on Visceral Regeneration in Sea-Stars. Many asteroids are active predators on shellfishes and even upon other starfishes; other asteroids are mud swallowers. Melanotransferrin: New Homolog Genes and Their Differential Expression during Intestinal Regeneration in the Sea Cucumber Holothuria Glaberrima. 316 (2), 323335. DAPT was also tested during arm regeneration of the brittle star Ophioderma brevispina (Mashanov et al., 2020). Regeneration of Spines and Pedicellariae in Echinoderms: a Review. Similarly, another study on lungfish tail regeneration, the authors found 16 TEs upregulated in the regenerating tail blastema (Verissimo et al., 2020). In morphallaxis, extensive phenomena of rearrangement/recycling from existing tissues take place: no blastema is involved and there is only limited and localized proliferation of cells derived from existing tissues by dedifferentiation, transdifferentiation and/or migration. On the other hand, Wnt14 showed continuous upregulation at all regenerating stages and BMP-1 was only upregulated at 7-dpe. Moreover, the expression of some of these molecules has been shown to differ significantly at some stage of intestinal regeneration. However, these genomes have not been explored or discussed in relation to the animals regeneration capacity. open Sci. Physiol. I. Regeneration of Pyloric Caeca in Henricia leviuscula (Stimpson). J. Exp. The effect of FGF signaling inhibition was studied in the regenerating arm of the brittle star A. filiformis (Czarkwiani et al., 2021). 52 (3-4), 166180. These and other studies underscore the impressive regeneration capacities of larval echinoderms, opening the possibility of using larvae to probe molecular mechanisms of the regeneration processes. Regeneration also occurs in echinoids,but is limited in terms of extent and degree of capabilities and only a few examples have been investigated (Dubois and Ameye 2001; Bonasoro et al. The identified genes were representatives of the common processes mentioned before, such as metabolic processes and translation regulation. Many of the pathways and processes identified in these studies correlate with those described in the regeneration of other tissues and organs. These dynamic changes are exemplified by focusing on those genes previously recognized as important for regeneration. J. Exp. doi:10.1016/j.cbpb.2017.06.005, Lin, G., and Slack, J. M. (2008). Therefore, this decrease of transcription activity at this stage compared to the earliest, demonstrate that an increment in cellular activity is needed once regeneration is initiated. One of the most recent, is the transcriptome profiling of A. japonicus polian vesicles (Shi et al., 2020). (2020). doi:10.1007/978-1-4939-7737-6_12, Cary, G. A., Wolff, A., Zueva, O., Pattinato, J., and Hinman, V. F. (2019). Wnt and Frizzled Expression during Regeneration of Internal Organs in the Holothurian Eupentacta Fraudatrix. 55 (6), 403426. 55 (6), 427437. The phylum Echinodermata consists of about 7000 living species and the phylum is divided into five smaller classes. Echinoderms, with their outstanding regenerative abilities, can provide important insights into the molecular basis of regeneration. The blastula invaginates at one end to form a primitive gut, and the cells continue to divide to form a double-layered embryo called the gastrula. In this study the focus was on ECM-related genes, as this change in the strength of the connective tissue would not be possible without ECM remodeling. University of Puerto Rico, Ro Piedras Campus, Ro Piedras, Puerto Rico. Comp. (2017). It has fragile arms formed by lobes of ciliated bands and is supported by fragile rods of calcite, the skeletal material. The third category comprises the studies where the functions of the candidate genes are tested. There are specific transcripts that support this as they have been identified to be involved in the generation of the digestive system during development, such as genes from the Sox and GATA family. Studies so far (and therefore this review) have been limited to exposing genes that are easily identifiable by database mapping, but very few have made efforts towards new gene discoveries. In general terms, animals in deuterostome clades that radiated before vertebrates show great regenerative capabilities (Bely and Nyberg, 2010). what type of symmetry do echinoderms have? Hyman, L. H. (1955). Therefore, this void serves as a reminder of the opportunities available for those that wish to focus on echinoderm regenerative biology. Genet. Echinoderms can perform asexual reproduction by fragmentation and cleaving off a portion of its body to regenerate a new one. Furthermore, there were development-associated genes that were expressed. For instance, at the earliest stage (3-dpe) one of the most represented transcripts was serum amyloid A (SAA) (Santiago et al., 2000; Santiago-Cardona et al., 2003). In this study, RNA sequencing (RNA-seq) was performed along with proteomic analyses, providing a higher level of confidence and confirmation of the molecular acitivtiy during this process. official Publ. Identification and Expression Characterization of WntA during Intestinal Regeneration in the Sea Cucumber Apostichopus Japonicus. (2013). In contrast, morphallaxis seems to be a more complicated and slower regenerative process which tends to follow traumatic mutilations, for instance in arm tip regeneration of asteroids (Mladenov et al. Proc. (2019). Thus, the Wnt pathway genes fulfill two of the requirements for a potential role (or roles) in the intestinal regeneration process: 1) genes associated with the pathway are expressed in cells of the regenerating intestinal rudiment and 2) some of these genes show differential expression associated with specific stages of the regeneration process. (2012). (2012). How Do Echinoderms Get Their Food - Micro B Life PLoS Biol. Notwithstanding, there is widespread ground to cover to clearly understand the molecular underpinnings of regeneration. doi:10.1016/j.margen.2015.02.004, Candia Carnevali, M. D., and Bonasoro, F. (2001). The drug used was SU5402, a specific inhibitor of FGF receptors that competes with ATP for the tyrosine kinase catalytic domain. Under artificial conditions, as in aquariums, echinoderms can survive apparent starvation for several weeks at a time. Although, experiments with pharmacological drugs are a step in the right direction, they remained, in many ways, inconclusive. Expression of Skeletogenic Genes during Arm Regeneration in the Brittle star Amphiura Filiformis. doi:10.1016/j.tice.2012.10.002, Kasahara, M., Kobayashi, C., Yamanaka, A., and Kitazawa, C. (2019). doi:10.1016/j.gene.2018.01.102, Viera-Vera, J., and Garca-Arrars, J. E. (2019). Many echinoderms aggregate before spawning, thus increasing the probability of fertilization of eggs. doi:10.1093/glycob/cwt100, Reinardy, H. C., Emerson, C. E., Manley, J. M., and Bodnar, A. G. (2015). Custom microarrays based on previous EST sequences were done, profiling differential gene expression of 3-, 7-, and 14-dpe regenerating intestines, and compared to normal intestines (Ortiz-Pineda et al., 2009). Res. Mar. These epimorphic mechanisms appear to reproduce rather closely what usually happens in embryonic developmental processes. 1998). BMC Dev. Dev. Among the enriched pathways identified are ribosome, proteasome, development, signaling (e.g., Notch, TGF-beta, Wnt), and metabolic pathways. (A) Stages of arm regeneration in the brittle star A. filiformis provide a baseline to detect the effect of regeneration modulating drugs as shown in Czarkwiani et al. In sexual reproduction, eggs (up to several million) from females and spermatozoa from males are shed into the water (spawning), where the eggs are fertilized. Both drugs reduced cellular proliferation in the intestinal rudiment, however MG132 treated animals showed a reduction in the size of the rudiment while E64d treated animals showed a delay in the degradation of collagen. Biol. TEs are genomic DNA sequences that are able to jump from one position to another by copying themselves and inserting the copy DNA into a new location or by excising themselves from the genome and reinserting into a new location. doi:10.1016/j.jinorgbio.2006.05.018, Mashanov, V., Akiona, J., Khoury, M., Ferrier, J., Reid, R., Machado, D. J., and Janies, D. (2020). Tissue and Cell 59, 19. Ecol. It is physically separated from most other tissues of the animal and can be easily isolated. The Mesentery as the Epicenter for Intestinal Regeneration, in Seminars in Cell & Developmental Biology (Academic Press), 92, 4554. Physiol. Mar. doi:10.1098/rspb.2002.2010, Piovani, L., Czarkwiani, A., Ferrario, C., Sugni, M., and Oliveri, P. (2021). A. doi:10.1002/jemt.1192, Verissimo, K. M., Perez, L. N., Dragalzew, A. C., Senevirathne, G., Darnet, S., Barroso Mendes, W. R., et al. Among the basket stars, a type of brittle star . Regeneration in Sea Lilies. J. Inorg. Furthermore, the study identified 11 transcription factors with potential roles in the control of genes during regeneration processes. Biol. Asexual reproduction in echinoderms usually involves the division of the body into two or more parts (fragmentation) and the regeneration of missing body parts. Dev. Evolution of Animal Regeneration: Re-emergence of a Field. Extensive studies have been done in ophiuroids and holothuroids. So far, only a few genes have been implicated in being differentially regulated during regeneration in echinoderms, including Hox1, and BMP2/4 and univin, from the TGF- superfamily of signaling molecules (Table 10.1). Specifically, regeneration has been mainly studied at this level in six species: the brittle stars A. filiformis, Ophionotus victoriae, and Ophioderma brevispina, where the focus is on arm regeneration and the sea cucumbers Eupentacta fraudatrix, A. japonicus, and H. glaberrima as models of intestinal regeneration. It has been documented that echinoderm larvae can regenerate certain structures and are even able to undergo fission and clone themselves to produce two embryos from the original one (Vickery et al., 2001; Eaves and Palmer, 2003). doi:10.1242/dev.180760, Czarkwiani, A., Dylus, D. V., and Oliveri, P. (2013). (2001). Wnt Signaling and Injury Repair. protein sequence of receptors or enzymes); in the present case in the VEGF receptor that is targeted by the axitinib. As seen before, their results showed that the extent of the differential expression in earlier stages was higher compared to those at later regenerative stages. Among ophiuroids, the brittle star Amphiura filiformis has been the main species for studies on specific genes associated with regenerative processes, particularly arm regeneration. 2 (5), 244246. Other highly expressed genes identified were associated to cell proliferation, division, and apoptosis (e.g., cleavage stimulation factor, polyubiquitin, proteasome, actin, and collagen). Characterization of Proteolytic Activities during Intestinal Regeneration of the Sea Cucumber, Holothuria Glaberrima. Myc Regulates Programmed Cell Death and Radial Glia Dedifferentiation after Neural Injury in an Echinoderm. Insights into Intestinal Regeneration Signaling Mechanisms. The first stage contained the highest percentage of differentially expressed genes, which suggested a higher transcription activity due to the numerous events required to form the blastema-like structure. Cell 169 (6), 985999. (2012). Echinoderms. Biol. doi:10.1016/j.cbd.2011.03.002, Sun, L., Yang, H., Chen, M., Ma, D., and Lin, C. (2013). Cell Dev. How do echinoderms reproduce? - WisdomAnswer Yes. Echinoderms have the ability to regenerate a missing arm, or ray. However, no effect on apoptosis or cell dedifferentiation occurred. Among significant genes with highest expression were craniofacial development protein 1 (CFDP2), THO complex (THOC1), transitional endoplasmic reticulum atpase (TER94) and adenosine kinase (adk). The main changes were in expression levels rather than in the presence or absence of gene expression. Certainly, the phylum Echinodermata contains some of the most suitable species to perform in depth studies in Regenerative Biology. There are about 2,000 species of sea stars living in the world's oceans in habitats from tropical coral reefs, kelp forests to the cold deep oceans (greater than 6 km), All sea stars are marine animals. doi:10.1371/journal.pbio.2003790, Zhu, W., Kuo, D., Nathanson, J., Satoh, A., Pao, G. M., Yeo, G. W., et al. doi:10.1016/j.semcdb.2018.09.001, Garca-Arrars, J. E., Lzaro-Pea, M. I., and Daz-Balzac, C. A. doi:10.1016/j.tree.2009.08.005, Ben Khadra, Y., Said, K., Thorndyke, M., and Martinez, P. (2014). genomics 9, 915. (2018). Visceral Regeneration in a Sea Cucumber Involves Extensive Expression of Survivin and Mortalin Homologs in the Mesothelium. (2011) was followed by a report using 7-days arm regenerating explants in vitro (Burns et al., 2012). An increase in the expression of the molecules at certain regeneration stages was associated with their possible involvement with cellular regenerative events, particularly with cellular migration. The degree to which these sequences are involved in this process is still unexplored, but as more genomic data is generated for distinct echinoderm species, we will be able to answer questions about these elements that were previously difficult to approach.

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