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A Test for Mucus Removal in the Chiton Lepidochitona Cinerea (Linnaeus, 1767) (Polyplacophora: Chitonida: Ischnochitonidae)

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A Test for Mucus Removal in the Chiton Lepidochitona Cinerea (Linnaeus, 1767) (Polyplacophora: Chitonida: Ischnochitonidae) A test for mucus removal in the chiton Lepidochitona cinerea (Linnaeus, 1767) (Polyplacophora: Chitonida: Ischnochitonidae) Sumner-Rooney, L. H., Cain, S., Brennan, G. P., & Sigwart, J. D. (2014). A test for mucus removal in the chiton Lepidochitona cinerea (Linnaeus, 1767) (Polyplacophora: Chitonida: Ischnochitonidae). Veliger, 51(4), 261-264. Published in: Veliger Document Version: Publisher's PDF, also known as Version of record Queen's University Belfast - Research Portal: Link to publication record in Queen's University Belfast Research Portal Publisher rights © The Authors This is an open access article published under a Creative Commons Attribution-NonCommercial-ShareAlike License (https://creativecommons.org/licenses/by-nc-sa/4.0/), which permits use, distribution and reproduction for non-commercial purposes, provided the author and source are cited and new creations are licensed under the identical terms. General rights Copyright for the publications made accessible via the Queen's University Belfast Research Portal is retained by the author(s) and / or other copyright owners and it is a condition of accessing these publications that users recognise and abide by the legal requirements associated with these rights. Take down policy The Research Portal is Queen's institutional repository that provides access to Queen's research output. Every effort has been made to ensure that content in the Research Portal does not infringe any person's rights, or applicable UK laws. If you discover content in the Research Portal that you believe breaches copyright or violates any law, please contact [email protected]. Download date:26. Sep. 2021 THE VELIGER # The Veliger 51(4):261–264 (Month, 2014) CMS, Inc., 2014 A Test for Mucus Removal in the Chiton Lepidochitona cinerea (Linnaeus, 1767) (Polyplacophora: Chitonida: Ischnochitonidae) LAUREN H. SUMNER-ROONEY Queen’s University Belfast, Marine Laboratory, 12–13 The Strand, Portaferry, Co. Down, Northern Ireland BT22 1PF, and Queen’s University Belfast, School of Biological Sciences, Medical Biology Centre, 97 Lisburn Road, Belfast, Northern Ireland BT9 7BL SHAUN CAIN Eastern Oregon University, La Grande, Oregon 97850, USA GERARD P. BRENNAN Queen’s University Belfast, School of Biological Sciences, Medical Biology Centre, 97 Lisburn Road, Belfast, Northern Ireland BT9 7BL AND JULIA D. SIGWART Queen’s University Belfast, Marine Laboratory, 12–13 The Strand, Portaferry, Co. Down, Northern Ireland BT22 1PF, and Queen’s University Belfast, School of Biological Sciences, Medical Biology Centre, 97 Lisburn Road, Belfast, Northern Ireland BT9 7BL Abstract. Several methods have been proposed to ‘‘clean’’ the soft tissues of molluscs of mucus, so that the surface cilia can be examined microscopically. We report the first empirical test of the effectiveness of methods for removing mucus in the pallial cavity surface of chitons. Three methods were compared, at several time intervals: the enzyme hyaluronidase, the mucolytic agent N-acetyl cysteine (NAC), and seawater washing via the natural action of cilia in excised tissue. Treatment in NAC for 10 min produced the best results, and we recommend this protocol as a starting point for further investigation on mucus removal in a broader suite of taxa. We present the first description of the pallial surface cilia in the chiton Lepidochitona cinerea. During the course of this study, we also determined that these chitons were frequently infested with a ciliate protozoan parasite, Trichodina sp., which have been historically reported from chitons but never studied in detail. The parasites were absent where antimucus treatments were effective, but their abundance and large size (about 30-mm diameter) in less successful treatments obscured the view of the pallial cavity surface. INTRODUCTION (e.g., Nezlin et al., 1994), but there has been no definitive protocol reported for use on adult chitons, nor Traditionally, molluscan systematics has relied largely any comparison of methods in any taxon. To determine on analyses of shell anatomy. More recently, with the the best way forward to visualize cilia in the pallial improvement of visualization techniques such as SEM cavity of chitons, we reviewed published techniques for imaging, shell features have been extended to micro- a variety of invertebrates and directly tested the effect on structural elements in gastropods (e.g., Geiger, 2012), the chiton Lepidochitona cinerea. bivalves (e.g., Turner, 2008) and chitons (e.g., Sigwart The enzyme hyaluronidase has been reported to & Sirenko, 2012). However, another potentially rich digest mucus and similar materials in a variety of taxa source of information that has still not fully exploited including bivalves (Calabro et al., 2005), proso- for systematics is microanatomy of the soft parts of the branchs (Nezlin et al., 1994), caenogastropods (Ron- body. Previous work on the visualization of soft parts kin, 1952), and onychophorans (Mayer & Harzsch, in other molluscan taxa using a scanning electron 2007). However, the efficacy of proteolytic enzymes microscope (SEM) has required the removal of mucus has been refuted (e.g., Stamm & Docter, 1965), and The Veliger veli-51-04-09.3d 25/7/14 16:43:39 261 Page 262 The Veliger, Vol. 51, No. 4 hyaluronidase has not been widely used. N-acetyl 10 min, or 30 min, and also at temperatures of either cysteine (NAC) has also been reported to function as 10uC or room temperature (six treatment regimes, two a mucolytic agent in both vertebrate and invertebrate specimens each). Four L. cinerea were decapitated and animals, including a marine bivalve (Moraes & Lopes, kept in seawater at 5uC for either 1 hr or for 24 hr before 2003), a sacoglossan (Pierce et al., 1996), and fixation (two treatment regimes, two specimens each), a planarians (Bocchinfuso et al., 2012), but its primary method to induce ‘‘natural’’ clearing of the tissue by the use has been in mammals (e.g., Stamm & Docter, action of still-active cilia, which has given good results 1965). In the absence of a chemical treatment, motor in Tritonia diomedea (SC, personal observation). cilia may continue to beat in excised tissue. So by Finally, one specimen of L. cinerea was fixed without excising tissue blocks and maintaining them in cold mucus removal treatment, as a control. seawater, the mucus could clear ‘‘naturally’’ (SC, personal observation). This is also analogous to washing in seawater or phosphate-buffered saline, RESULTS AND DISCUSSION which is another treatment suggested previously (e.g., The development of SEM techniques has been a crucial Buckland-Nicks, 1993). In order to test and optimize step forward in the study of microanatomy across all a treatment effective for chitons, we experimentally taxa, yet recent attempts to characterize the pallial compared these three approaches. cavity of another chiton, Leptochiton rugatus via SEM were hampered by excessive mucus obscuring the surface epithelium (JS, SC, personal observation). MATERIALS AND METHODS The SEM images here (Figure 1) show that the pallial Fourteen treatment regimes were tested for their cavity of Lepidochitona cinerea is carpeted with efficacy in mucus removal. These were all variants on microvilli, with large tufts of cilia (6 mmindiameter) the three techniques outlined above (hyaluronidase, of approximately 15–20 mm in length at regular NAC, and surgical separation) using different time and intervals (approximately 25 mm apart, Figure 1A, B). temperature intervals. Live specimens of Lepidochitona This pattern persisted throughout the pallial cavity and cinerea collected in Strangford Lough (County Down, into the gill row with no visible change in density or Northern Ireland) were used for each treatment. All frequency. These are the first images to illustrate the animals were held in aquaria flow-through seawater at surface features of the pallial cavity in L. cinerea. the Queen’s University Marine Laboratory (QML), The ‘‘success’’ of our treatments was assessed qualita- where one of the authors (JS) has successfully kept tively by visualization under SEM, to determine whether long-term captive specimens. All experiments were cilia were clearly visible, or whether tissue appeared badly performed on site in laboratory facilities at QML. dehydrated, or covered in mucus sheets obscuring some Following treated incubation with gentle agitation, or all of the cilia in the pallial cavity. Specimens were specimens were flushed with a pipette. All specimens examined whole; that is, the entire pallial cavity on both were fixed in 4% glutaraldehyde in cacodylate buffer sides was visually examined under SEM although only (pH 7.4) and post-fixed in 1% osmium tetroxide in small portions are illustrated herein (Figure 1). Temper- cacodylate buffer in preparation for SEM visualization. ature did not have any substantive influence on tissue Tissues were then dehydrated in acetone and prepared quality, and we conclude that performing these chemical for histology as described by Ruthensteiner (2008). treatments at room temperature does not decrease quality. Specimens were dried overnight using hexamethyldisi- The different treatments for mucus removal had lazane (HMDS, Sigma-Aldrich), sputter-coated with varying rates of success, with NAC generally producing gold using a Polaron E5100 SEM Coating System the best results. Incubation of specimens with hyal- (Quorum Technologies) and visualized in a FEI Quanta uronidase did not effectively remove mucus from the 200 scanning electron microscope operating at 10 kV. pallial cavity when incubated for less than an hour Bovine hyaluronidase (Sigma-Aldrich) was applied (Figure 1C, D), but after longer incubations both cilia at 4.46 e23 mM in low-calcium artificial seawater and microvilli were visible and only small spheres of (420 mM NaCl, 10 mM KCl, 50 mM MgCl2,10mM mucus were present. NAC preserved cilia and microvilli Trizma, 10 mM MnCl). Specimens were incubated for in good condition, and left little mucus when applied periods of 5 min, 10 min, or 1 hr, and at temperatures of for 10 min or more (Figure 1A, B). In addition, the use either 10uC or room temperature (six treatment regimes, of hyaluronidase appears to have disrupted the tissue, two specimens each).
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