As well known, in a few starfish species, individual autotomized arms can regenerate to produce new complete adults. Gene 654, 2335. The expression patterns of genes of ECM components showed diverse regulation patterns within specific families. Regeneration of Spines and Pedicellariae in Echinoderms: a Review. Echinoderms are sexually dimorphic and release their eggs and sperm cells into water; fertilization is external. 2001 c; Thorndyke and Candia Carnevali 2001) exploring the basic mechanisms of the regenerative phenomenon and its cellular and molecular aspects. what type of symmetry do echinoderms have? More so when we realize the multiplicity of processes, other than regeneration, that are taking place following evisceration, including activation of the immune system to deal with incoming pathogens, wound healing responses in the injured tissues and metabolic changes due to the loss of the main digestive organ, among others. Crinoids are suspension feeders, capturing planktonic organisms in a network of mucus produced by soft appendages, called tube feet, contained in grooves on the tentacles, or arms. 17 (1), 119. Genes 12 (7), 1063. doi:10.3390/genes12071063, Yuan, J., Gao, Y., Sun, L., Jin, S., Zhang, X., Liu, C., et al. Ser. In ascidians the potential for asexual development is expressed during colony formation by developing functional individuals from adult tissues. The more primitive, so-called regular, sea urchins are omnivorous or vegetarian browsers, either scraping algae and other small organisms from rocks with their hard teeth or eating seaweed. Move, exchange oxygen and carbon dioxide, capture food and release waste. Salamander-like Tail Regeneration in the West African Lungfish. Most echinoderms spawn on an annual cycle, with the spawning period normally lasting one or two months during spring or summer; several species, however, are capable of spawning throughout the year. Among the results of the study that should be highlighted are 1) the discovery of an increased expression of transposons that can be associated with the regeneration process (discussed extensively below) and 2) the finding that Myc, a Yamanaka factor (associated with induced cell plasticity) was upregulated in the early stages of radial nerve regeneration. Front. Holothurians as a Model System to Study Regeneration. In spite of this apparently clear and well-established difference between epimorphosis and morphallaxis, recent results obtained in the same model in different experimental conditions (see crinoid arm regeneration, Candia Carnevali and Bonasoro 2001b) suggest that the traditional classification of these two processes is too reductive and that the mechanisms at the tissue/cellular level can be largely interchangeable and interpretable in a more plastic and dynamic light. A similar expression pattern to that of Wnt genes, was seen in Hox1 and Hox3 with maximum expression peaks in the 3-dpe regenerating intestine. Other developmental genes commonly associated with regeneration, such as Hox and Wnt, were not found at any stage. Model Systems for Regeneration: Hydra. Do echinoderm Genomes Measure up. At the same time the experiments established a path on how the function of other candidate genes could be tested. In the case of one Japanese feather star (Crinoidea), spawning is correlated with phases of the Moon and takes place during early October when the Moon is in the first or last quarter. Sea cucumbers often discharge parts of their internal organs if they perceive danger. 332 (7), 245257. 19 (1), 119. The authors suggest that VEGF is not strictly required for skeleton formation but might be needed for establishing the patterning of the spicules. Expression of Wnt9, TCTP, and Bmp1/Tll in Sea Cucumber Visceral Regeneration. Larger eggs with greater amounts of yolk may develop into a larval form that is planktonic but subsists upon its own yolk material, rather than feeding upon small organisms, before eventually transforming into a juvenile echinoderm. The class Asteroidea contains the true sea stars . (2019). Biol. Echinoderms feed in a variety of ways. Sea urchins exhibit the lowest regenerative capacity but still can regenerate parts of its their test, broken or lost spines and pedicellariae (Dubois and Ameye, 2001). Complementary to this revision we would like to direct the readers to two previous reviews on the topic that provide some of the first insights into this area of research (Thorndyke et al., 2001; Mashanov and Garca-Arrars, 2011) and to a recent comprehensive review by Dolmatov (2021) on the molecular aspects of regeneration mechanisms in holothurians. Comp. Echinoderms feed on a variety of marine life in a variety of ways. One of these families encoded a group of prostatic secretory proteins of 94 amino acids (PSP94)-like genes while the other was a family of fibrinogen-related protein (FREP) genes. However, the extent of an organisms regenerative properties can vary significantly within a taxonomic group. (2018). The presence of these genes in a highly regenerative species, led the authors to posit that these proteins played a role in the organisms intestinal regeneration process. Most studies performed to determine the role of particular molecules or pathways during echinoderm regeneration have been done using specific drugs that inhibit enzymes, receptors or components of a signal pathway. In the species examined so far most results throw light on aspects related to wound healing, growth, morphogenesis and differentiation, but in most cases many crucial questions remain largely unanswered, especially those related to specific cellular and molecular aspects, and much work is still required. doi:10.1186/1471-2164-10-262, Pasten, C., OrtizPineda, P. A., and GarcaArrars, J. E. (2012a). Ecol. Zoolog. II. 25 (5), 828835. Among significant genes with highest expression were craniofacial development protein 1 (CFDP2), THO complex (THOC1), transitional endoplasmic reticulum atpase (TER94) and adenosine kinase (adk). Tech. Among the stages, the first and second stages (the blastema-like formation and 50% differentiation stage, respectively) shared a high number of differentially expressed genes (488 upregulated and 743 downregulated). 4 (8), a008078. (2020). As expected, the identified genes were similar to those found in their previous in vivo study of 7-days regenerating blastema-like structure (Burns et al., 2011). What type of feeders are echinoderms? The first report of gene knockdown in adult echinoderms was performed using the regenerating radial nerve cord of holothurians as a model system. TABLE 2. 27 (2), 105110. Attention was indeed paid to echinoderms in our recent Review 1. Scientific Rep. 11 (1), 114. Cytoskeletal genes showed specific expression profiles with alpha-tubulin, beta-tubulin, and actin showing high expression, while myosin, gelsolin and gamma-tubulin appeared downregulated. Front. Specifically, alpha-tubulin-1 and -2, and actin-1 and -2 were upregulated at all time points, whereas gelsolin, myosin, and actin-3 were downregulated. Development (Cambridge, England) 134 (3), 479489. 458 (1), 1231. Lipopolysaccharides Induce Intestinal Serum Amyloid A Expression in the Sea Cucumber Holothuria Glaberrima. Microsc. Some of the represented GO terms are metabolic processes, cellular signaling, binding processes, cell adhesion, and transcription/translation processes. Echinoderms. doi:10.1111/j.1440-169X.2012.01368.x, Keywords: echinoderm, regeneration, gene function, transcriptome analyses, signaling pathways, dedifferentiation, Citation: Medina-Feliciano JG and Garca-Arrars JE (2021) Regeneration in Echinoderms: Molecular Advancements. One of the initial studies done in Holothuria glaberrima led to the identification of the presence of the first serum amyloid A (SAA) ortholog in an invertebrate deuterostome (Santiago et al., 2000). Lond. These phenomena, which are also quite common in asteroids, provide evidence of the wide exploitation and implications of regenerative potential in echinoderms. Biol. In this case, amputation is not a predictable event and the morphogenetic mechanisms imply phenomena of substantial rearrangement of the old structures, which appear to be unique to regeneration and not shared by embryonic development. Regenerating Echinoderms: a Promise to Understand Stem Cells Potential, in Stem Cells in marine Organisms (Dordrecht: Springer), 165186. CHAPTER 23 and 24 Flashcards | Quizlet doi:10.1007/s00441-014-2040-4, Mashanov, V. S., Zueva, O. R., and Garca-Arrars, J. E. (2012a). Taken together, the molecular studies of arm regeneration in brittle stars reinforce what has been seen at the cellular level, where common cellular events are taking place such as apoptosis, proliferation, migration, and differentiation. Noteworthy, they also identified the translation regulation pathways as being associated with the brittle star early arm regeneration. Res. Many echinoderms have remarkable powers of regeneration. R. Soc. Mar. This will provide the basis for comparative analyses establishing unique molecular characteristics that might be responsible for the amazing regenerative processes observed in echinoderms. FIGURE 1. Fission is a form of asexual reproduction, during which sea cucumbers constrict around the middle region due to the changes in the connective tissue of the body wall. echinoderm, any of a variety of invertebrate marine animals belonging to the phylum Echinodermata, characterized by a hard, spiny covering or skin. doi:10.2307/1539385, Anderson, J. M. (1962). ossicles (spines) water vascular system. Echinoderm Structure and Function - Advanced ( Read ) | Biology Thus, we expect that this review will serve to entice researchers into exploiting echinoderm model systems for in depth biomedical studies in muscle . The authors exploited one of the main advantages of using echinoderm models; the possibility of comparing regenerative responses to embryonic development. (2017). Echinoderms belong to the phylum Echinodermata. 316 (2), 323335. Strikingly, this gene has been shown to be highly expressed in myoblasts proliferation (Loflin et al., 2006), which is in concordance with previous reports suggesting it is associated with the myocyte differentiation that takes place in early regeneration stages of A. filiformis (Biressi et al., 2010). Proc. doi:10.1016/j.ydbio.2017.05.015, Kudtarkar, P., and Cameron, R. A. open Sci. Components of the proteasome were found to be overexpressed during the early stages of intestinal regeneration (Pasten et al., 2012a). A recent study focused arm regeneration in another brittle star species O. brevispina (Table 1) (Mashanov et al., 2020). doi:10.1016/j.jembe.2011.06.032, Burns, G., Thorndyke, M. C., Peck, L. S., and Clark, M. S. (2013). Both drugs caused a decrease in the size of the regenerating rudiment and a significant reduction in cell division and cell dedifferentiation. Phylum Echinodermata Groups & Reproduction | How Do Echinoderms Among the enriched pathways identified are ribosome, proteasome, development, signaling (e.g., Notch, TGF-beta, Wnt), and metabolic pathways. In most asteroids the larval form in the next stage of development is called a brachiolaria, which has three additional arms used for attaching the larva to the seafloor. (2012). They identified a total of 1,733 differentially expressed transcripts among all samples, from which 791 were sequenced. The differential expression of genes associated with development, remodeling, transcription, and metabolic activity provide the molecular effectors that underlie the initial events of the regeneration process. The most enriched processes from this comparison were those related to serine family amino acid and polyol metabolism. Dev. Skeletal Regeneration in the Brittle star Amphiura Filiformis. Tissue and Cell 59, 19. Bridging the regeneration gap: insights from echinoderm models Reproduction is carried out by the release of sperm and eggs into the water. In general, results coincided with those found in the initial studies of H. glaberrima; many of the similar expression trends were those of developmental, cytoskeleton and ECM related family genes. On the other hand, and different to that found before, KLF-6 showed downregulation at 3-, 7- and 14-dpe, but upregulation at 21-dpe. doi:10.1038/nature25458, Nusse, R., and Clevers, H. (2017). (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. Regeneration in Echinoderms - Echinodermata - Barnard Health Care The phylum Echinodermata is composed of five major classes: Crinoidea (feather stars), Asteroidea (sea stars), Echinoidea (sea urchins), Ophiuroidea (brittle stars), and Holothuroidea (sea cucumbers) ( Figure 1 ). doi:10.1186/1471-213X-9-3, Santiago, P., RoigLpez, J. L., Santiago, C., and GarcaArrars, J. E. (2000). Transcriptome profiling of regenerating tissues in brittle stars. J. Exp. doi:10.1086/BBLv221n1p93, Mashanov, V. S., Zueva, O. R., and GarcaArrars, J. E. (2012c). The more advanced irregular echinoids, which usually lack teeth, are burrowers and pass small organisms to the mouth with the aid of spines and tube feet. Therefore, rather than assessing tissue specific transcripts, they aimed to identify genes that might stand out in regenerating tissues. Development 148 (10), dev180760. Regeneration is a phenomenon present, to some degree, in all metazoans from sponges to vertebrates. How do echinoderms use their water vascular system. Echinoderms, with their outstanding regenerative abilities, can provide important insights into the molecular basis of regeneration. Mar. Echinoderms; potential model systems for studies on muscle regeneration Biochem. Dev. Vavilov J. Genet. Therefore, the need to perform experiments that probe the identified genes to determine their function during the regeneration process. During metamorphosis, the fundamental bilateral symmetry is overshadowed by a radial symmetry dominated by formation of five water-vascular canals (see below Form and function of external features). (D) Results from Bello et al. Regeneration in Echinoderms and Ascidians - Wiley Online Library 15 (10), e2003790. This extensive and strategic employment of regenerative phenomena throughout the phylum indicates that in echinoderms regeneration actually represents an essential component of the life cycle, and this has a wide range of biological implications. Ophiuroids with arms branched in a complex manner may feed in a way similar to that of the crinoids. Analysis of conservation demonstrated the positive selection of Wnt signaling pathway genes among echinoderms. Conversely, at the 7-dpe stage there was a high representation of sequences corresponding to matrix metalloproteinases (MMPs), which are related to the ECM remodeling. genomics 22, 19. Insights into Intestinal Regeneration Signaling Mechanisms. In holothuroids it is also ubiquitous but has been explored in terms of mechanisms only in a few cases (Dolmatov and Ginanova 2001; Garcia-Arraras and Greenberg 2001). In this study, a total of 1,978 upregulated and 2,434 downregulated transcripts were identified in samples treated with a Notch pathway inhibitor. 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. Brittlestars Contain Highly Sulfated Chondroitin Sulfates/dermatan Sulfates that Promote Fibroblast Growth Factor 2-induced Cell Signaling. However, no effect on apoptosis or cell dedifferentiation occurred. (E) Holothuroidea represented by the sea cucumber Holothuria glaberrima. doi:10.1387/ijdb.200227dq, Ramachandra, R., Namburi, R. B., Dupont, S. T., Ortega-Martinez, O., van Kuppevelt, T. H., Lindahl, U., et al. doi:10.1371/journal.pbio.2003790, Zhu, W., Kuo, D., Nathanson, J., Satoh, A., Pao, G. M., Yeo, G. W., et al. Therefore, an extensive section of the article focuses on ophiuroids and holothuroids, being the only species, whose regeneration has been studied from in situ characterization up to functional studies. Other interesting transcripts were also identified as highly expressed at one or more stages, such as melanotransferrin (Mtf), centaurin and many other unknown transcripts. doi:10.1002/jemt.1187, Candia-Carnevali, M. D., Thorndyke, M. C., and Matranga, V. (2009). internal skeleton. 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). D: Genomics Proteomics 30, 143157. Regeneration of the radial nerve has also been studied in H. glaberrima. Finally, four different Wnt (A, 4, 6 and 16) were detected in regenerating internal organs of E. fraudatrix and each shown to have a distinct expression profile during the regeneration process (Girich et al., 2017). Microsc. Echinoderm regeneration studies have advanced in recent years, and will become more numerous, thanks to the accessibility and increase of genomic and molecular data. Coelomic Expression of a Novel Bone Morphogenetic Protein in Regenerating Arms of the Brittle star Amphiura Filiformis. Pisma v Vavilovskii Zhurnal Genetiki i Selektsii= Lett. Regenerating Echinoderms: A Promise to Understand Stem - ResearchGate RNA-seq Reveals Dynamic Changes of Gene Expression in Key Stages of Intestine Regeneration in the Sea Cucumber Apostichopus Japonicas. Zoolog. Expression of Skeletogenic Genes during Arm Regeneration in the Brittle star Amphiura Filiformis. (2020). Several species of sand dollars sometimes feed on suspended organisms carried to them by ocean currents as they lie on the seafloor. Biol. (Dolmatov et al., 2019). Mobile Genet. Reconstitutive regeneration of arms is particularly frequent in crinoids and ophiuroids which have fragile arms often involved in self-induced or traumatic mutilations. (2001). In the same report, VEGF was also studied in embryos and adults, by blocking its receptor with the drug axitinib. For instance, in ophiuroids, studies differ on the regions being dissected and on the description of regeneration stages with some focusing on differentiation index and other on days post amputation. Nature 554 (7690), 3435. 1989; Moss et al. This study was centered on the transdifferentiation mechanism of mesodermal cells as part of the regeneration process, since in this species, the coelomic epithelia of the anterior regenerating intestine gives rise to the luminal epithelia (Boyko et al., 2020). In brief, it is known that fibers and cells migrate from both nerve stumps forming a bridge that eventually gives rise to a new nerve cord region that is scar-free. Dev. The Mesentery as the Epicenter for Intestinal Regeneration, in Seminars in Cell & Developmental Biology (Academic Press), 92, 4554. The expression of the developmental genes upregulated at the first stage was different from those upregulated during the third stage, as would be expected due to the occurrence of different cellular events. Transcriptomic Analysis of Early Stages of Intestinal Regeneration in Holothuria Glaberrima. Echinoderms are known for their unique five-fold symmetry and their abilities of regeneration of limbs and wounded surfaces. 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. Table of Contents Home Science Bugs, Mollusks & Other Invertebrates Spiny-skinned Invertebrates echinoderm Asexual reproduction Asexual reproduction in echinoderms usually involves the division of the body into two or more parts ( fragmentation) and the regeneration of missing body parts. Most people know if a sea star loses one or more arms, the appendage(s) will eventually grow back. (2007). (2012). 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. Pharmacological studies in vivo and in vitro, provide additional (and much stronger) evidence for an active role. (2017). In comparison, eIF4G2, MSK1, myosin light chain phosphatase (MLCP) and Actin cytoskeleton transcripts showed upregulation. Among the top upregulated genes at 1-dpe were actin, serine/threonine-protein kinase NLK, translation elongation factor 2 (TEF2), and elongation factor 1 alpha (EEF1A). This extreme case clearly shows that in echinoderms regeneration is an indispensable complement of the programme of asexual reproduction which leads to the development of new individuals through fission mechanisms (Emson and Wilkie 1980; Mladenov and Burke 1994). In direct development the young usually are reared by the female parent. zoo chp 14 Flashcards | Quizlet These studies focused on perturbations in skeletogenesis and were accompanied by extensive analyses of gene expression that served as molecular markers. The study yielded an improved characterization of differentially expressed genes and their possible involvement in specific pathways and processes in A. japonicus. Inhibition of Cell Proliferation Does Not Slow Down Echinoderm Neural Regeneration. Embryos treated with the FGF receptor inhibitor failed to form the skeletal spicules needed to form the larval skeleton. doi:10.1242/jeb.02445, Eaves, A. Molecular studies of echinoderm regeneration can be grouped into three different categories. Biol. Sci. Echinoderms (starfish, brittle star, sea urchin, feather star, sea The first large-scale molecular studies of intestinal regeneration performed in a holothurian were done in H. glaberrima, using expressed sequence tags (EST) and microarrays (Rojas-Cartagena et al., 2007; Ortiz-Pineda et al., 2009). How Do Echinoderms Get Their Food - Micro B Life (2018). In these experiments MMP activity was shown to increase during intestinal regeneration, and this took place concomitantly with collagen degradation. PLoS Biol. In contrast, and surprisingly, Bmi-1 pluripotency factor appeared to be downregulated. In regenerative studies of non-traditional model species, such as sea cucumbers, these uncharacterized transcripts are an important aspect as they could lead to potential key players in the process. doi:10.1007/s101269900027, Nowoshilow, S., Schloissnig, S., Fei, J. F., Dahl, A., Pang, A. W., Pippel, M., et al. Regeneration is widespread among echinoderms -sea urchins regenerate spines and pedicellaria, sea cucumbers regrow their viscera after evisceration, brittle stars and sea stars regrow arms -but . 128 (1), 123. Terms in this set (9) Spiny skin. In some asteroids fragmentation occurs when two groups of arms pull in opposite directions, thereby tearing the animal into two pieces. In brief, soon after severing the arm, the injured tip is healed and re-epithelized. Regeneration in Echinoderms: Molecular Advancements - PMC However, the results are only correlative and there is no concrete proof that the listed genes are involved in regeneration.