1a) displayed the typical bi-layered structure of a pseudostratified myoepithelium described in the literature for other starfish (Wood and Cavey 1981; Rieger and Lombardi 1987), where apical squamous/cubic peritoneocytes and subapical longitudinally oriented myocytes are placed on the same basal lamina (Fig. Peritoneocytes are releasing apical cytoplasmic portions (asterisks). Our parallel proteomic analysis supported this, as in non-regenerating samples no proteins specific to stem/pluripotent cell production were detected. 3e). 2010). Ballarin L, Karahan A, Salvetti A, Rossi L, Manni L, Rinkevich B, Rosner A, Voskoboynik A, Rosental B, Canesi L, Anselmi C, Pinsino A, Tohumcu BE, Jemec Kokalj A, Dolar A, Novak S, Sugni M, Corsi I, Drobne D. Front Immunol. Unauthorized use of these marks is strictly prohibited. (JPG 301 KB). More than a simple epithelial layer: multifunctional role of echinoderm coelomic epithelium. 2022 Aug 3;20(8):503. doi: 10.3390/md20080503. We have used the transgenic mouse model mWt1/IRES/GFP-Cre (Wt1cre) crossed . 2a). These circulating elements are considered the immune cells involved in the clearance of foreign materials, pathogens, and cell debris, as well in haemostatic functions, since they form clots at the wound sites (Pinsino et al. 2010 Jan;52(1):57-68. doi: 10.1111/j.1440-169X.2009.01159.x. From a histological point of view, these vesicular cells belong to the circular muscle layer itself and can therefore be considered part of the myo-epithelial layer, as also suggested by the presence of scattered myocyte among them. Scale bar = 400m. Cell 123:889901. Fibers undergo rearrangement phenomena indicated by evident modifications of their contractile apparatus (arrows). Wound Rep Reg 23(4):611622, Ben Khadra Y, Ferrario C, Benedetto CD, Said K, Bonasoro F, Carnevali MD, Sugni M (2015b) Regrowth, morphogenesis, and differentiation during starfish arm regeneration. 2019). Russ J Mar Biol 33:411416. Each colored circle corresponds to one of the three biological replicates analyzed by LCMS/MS: in pink, number of proteins identified from A1E1 replicate; in green, number of proteins identified from B1F1 replicate; in blue, number of proteins identified from C1D1 replicate. Origin of coelom 1 1875-80; <Greek kolma cavity, equivalent to koil-, variant stem of koilon to hollow out (verbal derivative of kolos hollow) + -ma noun suffix denoting result Other words from coelom ajCE of 1week p.a. Apical secretion phenomena are evident (asterisks). Macrophages expressing this receptor have also been found in mammals during the healing phase of acute inflammation, in chronic inflammation, and in wound healing tissue (Fabriek et al. The mesonephros acts as transient fetal kidney in mammals and in females regresses around 55 days post coitum (dpc) in cow (term ~ 283 d) [19] and 75 dpc in sheep (term ~ 147 d) [3]. https://doi.org/10.1038/35008673, Miller PW, Pokutta S, Mitchell JM et al (2018) Analysis of a vinculin homolog in a sponge (phylum Porifera) reveals that vertebrate-like cell adhesions emerged early in animal evolution. https://doi.org/10.1007/s00441-006-0314-1, Dylus DV, Czarkwiani A, Blowes LM et al (2018) Developmental transcriptomics of the brittle star Amphiura filiformis reveals gene regulatory network rewiring in echinoderm larval skeleton evolution. https://doi.org/10.1016/j.immuni.2006.12.005, Yeo JC, Wall AA, Luo L, Stow JL (2016) Sequential recruitment of Rab GTPases during early stages of phagocytosis. BMC Biol 19:9. https://doi.org/10.1186/s12915-020-00937-7, Pobre KFR, Poet GJ, Hendershot LM (2019) The endoplasmic reticulum (ER) chaperone BiP is a master regulator of ER functions: getting by with a little help from ERdj friends. This study examined the role of coelomocytes outside circulation, named marginal coelomocytes and small undifferentiated cells of the coelomic epithelium in coelomocyte replenishment in Asterias rubens. At present, in adult echinoderms, CE is supposed to play fundamental roles acting as follows: Source of stem/multipotent cells (undifferentiated coelomocytes), particularly during regenerative phenomena (CandiaCarnevali et al. More than a simple epithelial layer: multifunctional role of echinoderm coelomic epithelium. Scale bar = 2m. Scale bar = 2m. A Search Tool for the Retrieval of Interacting Genes/Proteins (STRING) (Szklarczyk et al. Proteins identified in the M. glacialis CE by LCMS/MS (a, b). Curr Opin Genet Dev 32:2430. (3) Does the CE display other key functions involved in regeneration as well as in overall animal physiology? 2017). 2018; Ferrario et al. In fact, exo- and endocytosis work as coupled processes allowing the maintenance of the vesicle pool while preserving plasma membrane functionalities (Liang et al. The Rab GDP dissociation inhibitor regulates the GDP/GTP exchange reaction by inhibiting the dissociation of GDP from them, and the subsequent binding of GTP. These cells appeared scarcely differentiated, although bearing an apical flagellum and containing numerous electron-lucent vesicles, occasional large phagosomes, and myelin figures: in addition, at the distal-most tip of the new coelomic cavity, they clearly showed intense phenomena of apical secretion (Fig. 2010; Garca-Arrars et al. https://www.thefreedictionary.com/Celomic+epithelium. https://doi.org/10.1002/jez.1402720608, Candia Carnevali MD, Burighel P (2010) Regeneration in echinoderms and ascidians. 2a, b, d). Samples were resolved by liquid chromatography (nanoLC Ultra 2D, Eksigent) on a MicroLC column ChromXP (300m ID 15cm length, 3m particles, 120 pore size, Eksigent) at 5 L/min with a multistep gradient: 02min linear gradient from 2 to 5%, 245min linear gradient from 5 to 30% and, 4546min to 35% of ACN in 0.1% formic acid and 5% dimethyl sulfoxide, and analyzed on a TripleTOF 5600 System (ABSciex). Cell Tissue Res 218:449473. 1998). 2009). Also called: the serous membrane lining the abdominal cavity and investing its viscera. Springer, Netherlands, Dordrecht, pp 165186, Chapter https://doi.org/10.1007/978-0-387-48993-3_1, Naslavsky N, Caplan S (2018) The enigmatic endosome - sorting the ins and outs of endocytic trafficking. Note the presence of coelomocyte clots at the level of the wound (asterisk), the thickening of the perivisceral CE immediately close to the amputation site, and the new thin epithelium covering the wound edges (arrow). This increased internal pressure is generally due to an intense secretion of the epithelium of the organ itself (Tsarouhas et al. According to Righettis protocol (Fasoli et al. Briefly, the deep-frozen CE was placed in a previously chilled Teflon capsule containing 3 stainless steel beads (5mm in diameter). 2011; Ben Khadra et al. The irregular distribution of small cells on the surface and within the coelomic epithelium was demonstrated and the origin of small undifferentiated cells and large agranulocytes from the coelomic epithelium was suggested. https://doi.org/10.1134/S1990519X12020113, Smith LC, Ghosh J, Buckley KM et al (2010) Echinoderm immunity. https://doi.org/10.1007/BF00210107, Wyles JP, McMaster CR, Ridgway ND (2002) Vesicle-associated membrane protein-associated protein-A (VAP-A) interacts with the oxysterol-binding protein to modify export from the endoplasmic reticulum. 2007). The vesicle-associated membrane proteinassociated protein A is localized in the ER/Golgi pathway. The sexually undifferentiated gonadal ridge forms as a thickening of the coelomic or surface epithelium in an anterior/posterior direction on the ventral side of the mesonephros. Genome Biol 19:26. https://doi.org/10.1186/s13059-018-1402-8, Fabriek BO, Dijkstra CD, van den Berg TK (2005) The macrophage scavenger receptor CD163. Circular muscle layer (cm) is housed within the dermis (d) and is surrounded by a proper basal lamina (bl). https://doi.org/10.1007/s00441-017-2589-9, Bhatia VN, Perlman DH, Costello CE, McComb ME (2009) Software tool for researching annotations of proteins: open-source protein annotation software with data visualization. In other animals, such as molluscs, it remains undifferentiated. Peritoneocytes were massively involved in peculiar phenomena of apocrine secretion (Fig. eCollection 2021. Rolling collision was used with a collision energy spread (CES) of 5. https://doi.org/10.1016/j.ydbio.2017.09.035, Franco CF, Santos R, Coelho AV (2011) Proteome characterization of sea star coelomocytesthe innate immune effector cells of echinoderms. Bookshelf aSemithin longitudinal section of M. glacialis regenerating arm tip. Trends Cell Biol 24:352359. Biology 9:234. https://doi.org/10.3390/biology9080234, Article 2021). Re-growing perivisceral coelomic cavity (cc) and radial water canal (rwc) appear to penetrate an area of loose coelomocytes clot (asterisk) at the level of the wound site. Unusual vesicular cells (vc) are detected close to circular myocytes: they project their flagella into intercellular niches. Ultrathin sections were mounted on copper grids and stained with 1% uranyl acetate followed by lead citrate. Browse the use examples 'coelomic epithelium' in the great English corpus. pronouncekiwi Listen to the pronunciation of Coelomic epithelium and learn how to pronounce Coelomic epithelium correctly. Most identified proteins are cytoplasmic (26%) or cytoskeletal (19%) (Fig. 2018). showing peritoneocytes (arrow); longitudinal muscle layer (lml); circular muscle layer (cml) with first signs of rearrangement; vesicular cells (asterisks). The present research was performed according to the national laws; the species used under this study are not endangered and their use does not require any ethical approval. In particular, proteomics is becoming a powerful tool to deepen the problem of cell and tissue functions during both tissue homeostasis and regeneration by depicting the existing protein activities involved in the regrowth processes. A further fact must be considered in deciding this question, which is the discovery of ramifying coelomic tubes, approaching close to, but not entering, the epidermis in the Polychaete Arenicola. Scale bar = 5m. Flagellated vesicular cells are apparently involved. gTEM micrograph of regenerating perivisceral CE (p). 2010), underlining the immune function of these cells. Starfish were anesthetized with 4% (w/v) MgCl2 in sea water and two arm tips (12cm from the arm tip) for each animal were removed with a scalpel. Biol Bull 159:295310. 1e). https://doi.org/10.1093/nar/gku1003, Teixeira FG, Carvalho MM, Panchalingam KM et al (2017) Impact of the secretome of human mesenchymal stem cells on brain structure and animal behavior in a rat model of Parkinsons disease. Proteins identified following the defined acceptance criteria in at least one of the three biological replicates with one of the methods (IDA or SWATH) were considered for posterior analyses. (TIF 9956 KB), Supplementary file3 Online Resource 3 Identified proteins in each biological replicate (A1E1, B1F1 and C1D1) using Information-Dependent Acquisition (IDA) and Sequential Window Acquisition of All Theoretical Mass Spectra (SWATH-MS) methods. 2007). Ana Varela Coelho or Michela Sugni. Considering the molecular function (Fig. 2e), it was most evident in the radial water canal CE, particularly at the level of the terminal podium (Fig. 2014), and has a key regulatory function in the inflammatory response (Bradley 2008). Characterization of Coelomic Fluid Cell Types in the Starfish, 15-04-07798 / (), Alvarado AS, Tsonis PA (2006) Bridging the regeneration gap: genetic insights from diverse animal models. cTEM micrograph of regenerating CE at 24h p.a. Learn the definition of 'coelomic epithelium'. Scattered among these vesicular cytotypes, other cells containing phagosomes and presumptive lipid droplets (Fig. Google Scholar, Ben Khadra Y, Ferrario C, Di Benedetto C et al (2015) Wound repair during arm regeneration in the red starfish Echinaster sepositus. [Ultrastructure of coelomic epithelium and coelomocytes of intact and wounded starfish Asterias rubens L]. They were fed ad libitum with mussels collected at the same site. 2011). Secreted cytoplasmic portion of different shapes and sizes (asterisks) are visible in the perivisceral coelomic lumen. Results Probl Cell Differ 65:285320. 2a, b, d), through which the apical portions of these cells (including cytoplasm and cell membrane) were released into the coelomic lumen. iTEM micrograph of new radial water canal (rwc) CE. . Science 343:12531256. These membrane vesicles (30150nm in size) offer a potent mechanism for communication between nearby or distant cells or tissues in order to change their physiological functions and properties in specific innate and adaptive immune responses, as reported in several marine invertebrates. In particular, from IDA analysis, 338 proteins were identified (Fig. This lipid is a component of cell organelles that regulates a variety of sorting processes ensuring the proper distribution of endocytic cargo that is trafficked through the endosome for degradation or returned to the plasma membrane after recycling (Naslavsky and Caplan 2018). (2015) and, interestingly, in coelomocytes (Franco et al. Overall, proteins specific to hematopoiesis were not found, this result being explainable by the presence in the total protein extract analyzed by LCMS/MS of highly abundant proteins, which hampers the identification of lower abundant proteins. Coelom Definition. However, being a homology-driven proteomics characterization, several of the identified proteins from the starfish CE were homologous to proteins from other organisms (i.e., Chordata 28.5%; Arthropoda 18.5%; Mollusca 11.5%; other marine invertebrates 9%; complete list of characterized proteins in Online Resource 4). Zoomorphology 106:4960. This phenomenon perfectly fits with what has been described in other echinoderms. Although also present in the perivisceral coelom (Fig. In non-regenerating samples, we never observed presumptive stem cells within the CE, although the possibility that these are located in other non-investigated part of the CE cannot be excluded. Clipboard, Search History, and several other advanced features are temporarily unavailable. Gene expression of Ras-related protein Rab, a protein from the small GTPases Rab family, is induced by LPS to regulate the replenishing of the plasma membrane with TLRs (Yeo et al. Methods Mol Biol 1120:118. BMC Dev Biol 11:61. https://doi.org/10.1186/1471-213X-11-61, Gillet LC, Navarro P, Tate S et al (2012) Targeted data extraction of the MS/MS spectra generated by data-independent acquisition: a new concept for consistent and accurate proteome analysis. 6b). Scale bar = 50m. 2008; Sharlaimova et al. https://doi.org/10.1002/jez.b.21352, Holm K, Dupont S, Skld H et al (2008) Induced cell proliferation in putative haematopoietic tissues of the sea star, Asterias rubens (L.). Scale bar = 5m. 2009), and the brittle star Amphiura filiformis (Dylus et al. This is in line with the remarkable development, diversification, and complexity of coelom and coelom-derived structures in adult echinoderms, which have no equal among the coelomates. Our aim was to investigate the contribution of coelomic epithelium-derived cells to the intestinal development. Mar Drugs. aSemithin longitudinal section of M. glacialis arm tip at 24h p.a. gTEM micrograph of vesicular cells showing flagella (arrow) projecting into intercellular niches. That article is substantially longer and more informative than this stub. The origin of the observed vesicular layer is uncertain. However, proteomics investigations focused on echinoderm CE are quite limited. 1993). This hypothesis is corroborated by the identification of proteins associated with cell motility that may indicate a migration of cells from CE to the coelomic cavity. Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. Cell Logist 6:e1140615. This study examined the role of coelomocytes outside circulation, named marginal coelomocytes and small undifferentiated cells of the coelomic epithelium in coelomocyte replenishment in Asterias rubens. . National Library of Medicine Vacuum-dried (Scientific Speed-Vacuum, Thermo) tryptic digests re-suspended in 2% (v/v) acetonitrile (ACN) 1% (v/v) formic acid were desalted and concentrated by C18 microcolumns (OMIX tips, Agilent Technologies, Santa Clara, CA, USA) following the manufacturers instructions. There was no direct evidence of active cell release from the CE nor of any presence of morphologically undifferentiated cells throughout the CE. https://doi.org/10.1016/j.dci.2013.01.007, Dolmatov IY (1993) Proliferation of tissues of regenerating aquapharyngeal complex in holoturians. J Pathol 214:149160. In mice, the first morphological sign of the genital ridge formationappearance of the coelomic epithelium thickening and disintegration of the basement membrane underneathis noticeable at 10.3-10.4 dpc (days post coitum), i.e., stage 5-6 ts (tail somite stages) (Hu et al. This hypothesis was also supported by proteomic analyses highlighting partially shared protein profiles between the CE and the circulating coelomocytes (Sharlaimova et al. Both longitudinal and circular muscle layers are not detectable. Zoology. Anal Chem 81:98199823. https://doi.org/10.1007/s00435-009-0095-7, Bonasoro F, Carnevali C, Moss C, Thorndyke M (1998) Epimorphic versus morphallactic mechanism in arm regeneration of crinoids and asteroids: pattern of cell proliferation/differentiation and cell lineage, Bossche JPV, Jangoux M (1976) Epithelial origin of starfish coelomocytes. https://doi.org/10.1007/978-3-319-92486-1_14, Ben Khadra Y, Sugni M, Ferrario C et al (2017) An integrated view of asteroid regeneration: tissues, cells and molecules. 2ad), typical junctional complexes between adjacent peritoneocytes being recognizable apically (Fig. 2021 Jun 30;12:688106. doi: 10.3389/fimmu.2021.688106. The vesicular cells displayed larger phagosomes if compared to the previous stage, often including rests of disorganized contractile apparatus of myocytes (Fig. 2014). https://doi.org/10.1126/science.1248228, Han J, Lee JD, Bibbs L, Ulevitch RJ (1994) A MAP kinase targeted by endotoxin and hyperosmolarity in mammalian cells. HHS Vulnerability Disclosure, Help Two micrograms of recombinant green fluorescent protein (MBP-GFP) was added to each protein sample as an internal standard for retention time adjustment and intensity normalization. Note the apocrine secretion(s) of peritoneocyte apical portions. https://doi.org/10.1379/csc-288.1, Piovani L, Czarkwiani A, Ferrario C et al (2021) Ultrastructural and molecular analysis of the origin and differentiation of cells mediating brittle star skeletal regeneration. 2011), the deep-frozen powdered sample was re-suspended in PBS buffer (20mM phosphate-buffered saline, pH 7.2, containing 5mM ethylenediaminetetraacetic acid (EDTA) and 1mM dithiothreitol), supplemented with a protease inhibitor cocktail 0.2 (SigmaFast, Sigma). Wound Repair Regen 23(4):623634 2010; Sharlaimova et al. 2016). Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. fSemithin longitudinal section of radial water canal CE: a massive apical secretion is shown. The heart is a special case because it is continuous with . 3a, b). The Arp2/3 is a complex involved in the regulation of the actin cytoskeleton and in the formation of protrusive processes, like pseudopodia and filopodia, that is a crucial event for phagocytosis, a central process in immunity and, most of all, in autophagy and debris clearance (May et al. 16, Muoz-Chpuli R, Carmona R, Guadix JA et al (2005) The origin of the endothelial cells: an evo-devo approach for the invertebrate/vertebrate transition of the circulatory system. https://doi.org/10.1146/annurev.physiol.66.071702.145229, ONeil PT, Machen AJ, Deatherage BC et al (2018) The Chaperonin GroEL: a versatile tool for applied biotechnology platforms. Upload it here to share it with the entire community. 2g). However, we cannot answer medical or research questions or give advice. Log in to Pronouncekiwi Note the absence of the circular muscle layer. Google Scholar, Ariza L, Carmona R, Caete A et al (2016) Coelomic epithelium-derived cells in visceral morphogenesis. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. Appearance of the ovary depends widely on the age / menopausal state: Newborn ovaries are elongated and approximately 1.3 cm in greatest dimension, Ovaries enlarge during infancy and childhood and reach adult size and shape by the time of puberty, Neonatal ovaries often have cysts, which resolve spontaneously (, 3 - 5 cm long and weigh 5 - 8 g; size and weight depend on the amount of follicular derivatives (cysts and corpora albicantia / lutea), Pink-white exterior is initially smooth but gradually becomes more convoluted, Cystic follicles and corpora lutea may be visible from outside, Cut section may exhibit 3 zones: cortex, medulla and hilus, with follicular derivatives usually in the cortex and medulla, Firm consistency with solid, pale cut surface, Occasional cysts measuring several millimeters in diameter (inclusion cysts) may be discernible within the cortex, Small white scars (corpora albicantia) are typically present within the medulla, Thick walled blood vessels may be appreciable within the medulla and the hilus, Arterial supply: approximately 10 arterial branches from anastomotic arcade of ovarian artery (branch of aorta) and ovarian branch of uterine artery penetrate hilus into medulla and cortex, Venous drainage: left ovarian vein drains to left renal vein, right ovarian vein drains to inferior vena cava, Lymphatic drainage: originates predominantly from theca layer of follicles, exiting through the hilus, to the mesovarium, along the infundibulopelvic ligament, into upper paraaortic lymph nodes; may bypass to internal iliac, external iliac, common iliac, sacral, obturator, pelvic, retroperitoneal or inguinal nodes, Enlarged corpus luteum cyst can trigger an intraoperative consultation; the cyst lining is yellow and has a convoluted appearance, Luteoma of pregnancy can be discovered incidentally during cesarean section or preoperatively on imaging study mimicking an ovarian mass (. The total protein concentration of each replicate was determined using the QuantiPro BCA Assay Kit according with the manufacturers protocol. The biological processes attributed to the majority of the identified proteins in the CE (Fig. The origin of cells involved in regeneration in echinoderms remains an open question. Nat Cell Biol 2:246248. Curr Pharm Des 16:942955, Garca-Arrars JE, Valentn-Tirado G, Flores JE et al (2011) Cell dedifferentiation and epithelial to mesenchymal transitions during intestinal regeneration in H. glaberrima. https://doi.org/10.1016/j.jprot.2010.10.009, Ferrario C, Ben Khadra Y, Czarkwiani A et al (2018) Fundamental aspects of arm repair phase in two echinoderm models. Russ J Dev Biol 24:7281, Dolmatov IY, Ginanova TT (2001) Muscle regeneration in holothurians. https://doi.org/10.1134/S1990519X10030107, Sharlaimova N, Shabelnikov S, Bobkov D et al (2021) Coelomocyte replenishment in adult Asterias rubens: the possible ways. Each arm was opened and the perivisceral CE was removed from the inner oral and aboral sides of the arm with tweezers. In echinoderms, this protein is expressed in coelomocytes following physical stress (Ben Khadra et al. Overall, our work underlines that echinoderm CE is not a simple epithelial lining and that instead it plays multiple functions which span from immunity-related roles as well as being a source of regeneration-competent cells for arm growth/regrowth. Silvia Guatelli was financed by an Erasmus Grant (European Commission). Several proteins found in M. glacialis CE proteome are proper of the immune response (Table 1). Adv Wound Care (new Rochelle) 3:445464. 2017). The Rho-related protein (RacB) and cell division cycle 42 (Cdc42) protein, both found in our analyses, are required for the activation of the Arp2/3 complex to form protrusive structures during phagocytosis, whereas Rho controls actin assembly at phagosomes (Olazabal et al. Nucleic Acids Res 43:D447-452. 2005). https://doi.org/10.1016/j.imbio.2005.05.010, Fasoli E, DAmato A, Kravchuk AV et al (2011) Popeye strikes again: the deep proteome of spinach leaves. 2008; Gorshkov et al. At 24h p.a., the perivisceral CE close to the wound area was apparently thicker than that in non-regenerating conditions (Fig. PubMed Central 2021). Replenishment of circulatory coelomocytes-cells of the coelomic cavity in starfish-is an example of physiological regeneration. 4f) basally hosting a highly euchromatic nucleus and apically joined to each other by typical junctional complexes. These cells presented large euchromatic nuclei and high nucleus/cytoplasm ratios. In echinoderms, it was regarded as ancient multifunctional organogenetic tissue (Candia Carnevali et al. You can contribute this audio pronunciation of coelomic to HowToPronounce dictionary. (2014, 2021) identified a sub-population of small undifferentiated cells in the CE context and suggested these elements as putative progenitors and source of new circulating coelomocytes. Cell Tissue Res 390, 207227 (2022). 2010) and our observations confirm this morphological analogy, thus providing further support to the hypothesis of a coelothelial origin of this cell sub-population. Evol Dev 7:351358. Their cytoplasm was characterized by several small electron-lucent or finely granular roundish vesicles, different in shapes and sizes, occasional phagosomes, myelin figures, and presumptive lipid droplets (Fig. Scale bar = 1m. Adv Exp Med Biol 708:260301. 6c), proteins having a binding function and a catalytic activity were highly represented (46% and 37%, respectively). 2k) were occasionally detected as well, together with neuronal or presumptive neurosecretory processes. A qualitative and quantitative comparison of circulatory and marginal coelomocytes, as well as changes of circulatory coelomocyte concentrations in response to injury at different physiological statuses, was analysed. Miller-Keane Encyclopedia and Dictionary of Medicine, Nursing, and Allied Health, Seventh Edition . Sorting nexins belong to a family of peripheral proteins that contain a phosphoinositide-binding domain.