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Arm Swapping Autograft Shows Functional Equivalency of Five Arms in Sea Stars

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Arm Swapping Autograft Shows Functional Equivalency of Five Arms in Sea Stars bioRxiv preprint doi: https://doi.org/10.1101/676254; this version posted June 26, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. Title: Arm swapping autograft shows functional equivalency of five arms in sea stars Authors: Daiki Wakita1, Hitoshi Aonuma1,2, Shin Tochinai3 1Graduate School of Life Science; Hokkaido University; Sapporo, Hokkaido, 060-0810; Japan 2Research Institute for Electronic Science; Hokkaido University; Sapporo, Hokkaido, 060-0812; Japan 3Field Science Center for Northern Biosphere, Hokkaido University; Sapporo, Hokkaido, 060-0811; Japan Corresponding author: Daiki Wakita Graduate School of Life Science; Hokkaido University; Sapporo, Hokkaido, 060-0810; Japan. Tel/Fax: +81-11-706-3832, Email: [email protected] Additional keywords: echinoderm, pentaradial symmetry, movement coordination, regeneration, positional information bioRxiv preprint doi: https://doi.org/10.1101/676254; this version posted June 26, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 1 Abstract 2 3 Extant echinoderms show five-part radial symmetry in typical shape. However, we can find 4 some asymmetry in their details, represented by the madreporite position not at the center, 5 different skeletal arrangement in two of the five rays of sea urchins, and a circular cavity 6 formed by two-end closure. We suspect the existence of any difference in hidden information 7 between the five. In our hypothesis, deep equivalency makes no issue in function even after 8 exchanging the position of rays; otherwise, this autograft causes some trouble in behavior or 9 tissue formation. For this attempt, we firstly developed a method to transplant an arm tip to 10 the counterpart of another arm in the sea star Patiria pectinifera. As a result, seven arms were 11 completely implanted—four into the original positions for a control and three into different 12 positions—with underwater surgery where we sutured with nylon thread and physically 13 prevented nearby tube feet extending. Based on our external and internal observation, each 14 grafted arm (i) gradually recovered movement coordination with the proximal body, (ii) 15 regenerated its lost half as in usual distal regeneration, and (iii) formed no irregular 16 intercalation filling any positional gap at the suture, no matter whether two cut arms were 17 swapped. We here suggest a deep symmetry among the five rays of sea stars not only in 18 morphology but also in physiology, representing an evolutionary strategy that has given equal 19 priority to all the radial directions. Moreover, our methodological notes for grafting a mass of 20 body in sea stars would help echinoderm research involving positional information as well as 21 immunology. 2 bioRxiv preprint doi: https://doi.org/10.1101/676254; this version posted June 26, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 22 1. INTRODUCTION 23 24 Extant echinoderms typically show pentaradial symmetry in apparent morphology. We 25 recognize a regular star shape in a sea star at first glance and we collect five pairs of gonads 26 from a sea urchin for food. The five sectors seem to have the same structure and function. 27 Looking into it in detail, however, we become aware of some difference between them. 28 Madreporite is a remarkable one in common echinoderms. On the aboral side of a sea star for 29 instance, we find the madreporite as a button-like structure at a position away from the center. 30 This is because its body has the ring canal opening outward just at one place, which is part of 31 the water vascular system (Nichols, 1971). Thus we can identify each ray in echinoderms 32 based on the madreporite position. One common naming is A to E clockwise in the oral view 33 (Carpenter, 1884), where the madreporite places at the interradius between D and E in the 34 case of general sea stars, and actually the anus also leans a bit toward the C to D region 35 (Hotchkiss, 1995; Moore & Fell, 1966). When limited to sea urchins and some extinct taxa, 36 another asymmetry is found in the alternate pattern of paired plates along each 37 ambulacrum—midline area in each of the five rays. The plate series on the anticlockwise side 38 is in advance of those on the clockwise side in three rays, whereas the other two distant rays 39 show the opposite arrangement (Lovén, 1874). This law has been symbolized by “aabab” 40 (Hotchkiss, 1978, 1995); “a” represents the anticlockwise-advanced ray. In the metamorphic 41 process, echinoderms show a drastic transformation from bilaterally symmetric larva to 42 radially symmetric adults. Here, hydrocoel—anlage of the water vascular system—initially 43 forms a crescent, and then closes the two ends at the interradius between C and D to complete 44 the ring (Gemmill, 1912; Hotchkiss, 1995; Mooi & David, 2008). Based on this closure 45 position, we can postulate the first “a” of the “aabab” pattern in sea urchins is homologous to 46 the ray E in sea stars (Figure 1A) although none has recognized this asymmetrical 47 arrangement from the skeleton of sea stars (Hotchkiss, 1978, 1995). The circumoral 3 bioRxiv preprint doi: https://doi.org/10.1101/676254; this version posted June 26, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 48 (clockwise-anticlockwise) axis developed with exposure to spatially different expression of 49 Hox genes, according to in situ hybridization in metamorphosing sea urchins (Arenas-Mena, 50 Cameron, & Davidson, 2000). In behavioral terms, some studies have revealed a slight 51 preference at the population level in moving and turning-over directions based on repeated 52 observations of sea stars (Ji, Wu, Zhao, Wang, & Lv, 2012; Pollis & Gonor, 1975). Such 53 cracks hidden in pentamerism would reflect an evolutionary background of echinoderms, 54 where their early body had shown obvious asymmetry or even bilateral symmetry (Rozhnov, 55 2014; Sumrall & Wray, 2007). 56 Insights in many aspects give rise to the suspicion of some potential difference 57 among the five apparently equivalent sectors, in terms of functional expression in the adults 58 of extant echinoderms. Our approach for this enigma is to physically exchange two sectors of 59 the five so as to see whether some conflicts arise or not (Figure 1B), which has never been 60 proved. We used the blue bat star Patiria pectinifera, in which we developed a method to 61 transplant an arm to another arm’s position within an individual (Figure 2). In this paper, we 62 firstly introduce autograft technique in exchanging sea stars’ arms, and then probe into any 63 inter-arm differences based on the observation of grafts. We hypothesize the presence of 64 asymmetry in some properties hidden in the deep structure of pentamerism. One possible 65 gradient can be inspired by the “aabab” pattern (Figure 1A) in sea urchins. When the tip of a 66 sea star’s arm potentially with the “a” property is grafted into the base of another arm with “b,” 67 this gap would make some conflict on the suture (Figure 1B). This operation could discover 68 the hidden existence of the “aabab” pattern in extant sea stars. Another simple expectation is 69 that some property differs arm by arm, given the ontogeny where the ring canal originates 70 from a curved linear structure (Figure 1C). In this instance, swapping autograft of two arm 71 tips always brings proximal-distal mismatches, leading to some conflict. For a possible 72 conflict, we suppose (i) impaired coordination across the suture, (ii) abnormal distal 73 regeneration in the graft, or (iii) intercalary regeneration (Iten & Bryant, 1975) at the suture 4 bioRxiv preprint doi: https://doi.org/10.1101/676254; this version posted June 26, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 74 for filling a gap of positional information. The conclusion of our study is, however, contrary 75 to the hypotheses above. We recognized no functional difference among sea stars’ five arms as 76 schematized in Figure 1D, since arm grafts finally behaved as if they saw no exchange. Part 77 of this work has been briefly described in conference proceedings by Wakita and Tochinai 78 (2017), who focused on external behavior without the mention of detailed methods, failure 79 cases, ray identification, regeneration, and internal observation; this paper details the whole. 80 81 82 2. METHODS 83 84 2.1 Animals 85 We collected the blue bat star Patiria pectinifera (MÜLLER & TROSCHEL, 1842) (Asteroidea, 86 Echinodermata; Figure 2A) at Oshoro Bay (Otaru, Hokkaido, Japan) in April to October, 2014. 87 Sea stars were reared in laboratory aquariums in 60 × 30 × 36 cm filled with filtered aerated 88 natural sea water at 24°C and fed with clams, shrimps, and squids once a week. 89 90 2.2 Autograft 91 For autograft experiments, we used 46 individuals which had five arms with almost equal 92 length and had been reared in laboratory for at least a week—individuals #1–6 collected in 93 April; #7–32 in May, and #33–46 in October; surgery conducted in April to November, 2014 94 (Table 1). We applied various ways to graft an arm tip into the counterpart of another arm 95 within an individual (“Method” in Table 1).
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