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Acantharia

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222: 434:: they acquire energy through both heterotrophy and autotrophy. The relationship may make it possible for acantharians to be abundant in low-nutrient regions of the oceans and may also provide extra energy necessary to maintain their elaborate strontium sulfate skeletons. It is hypothesized that the acantharians provide the algae with nutrients (N & P) that they acquire by capturing and digesting prey in return for sugar that the algae produces during photosynthesis. It is not known, however, whether the algal symbionts benefit from the relationship or if they are simply being exploited and then digested by the acantharians. 482: 412: 63: 42: 653: 402:
genes, although the groups are mostly polyphyletic. Holacanthida seems to have evolved first and includes molecular clades A, B, and D. Chaunacanthida evolved second and includes only one molecular clade, clade C. Arthracanthida and Symphacanthida, which have the most complex skeletons, evolved most
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and it is therefore believed that the cysts help acantharians sink into deep water. Genetic data and some imaging suggests that non-cyst-forming acantharians may also sink to deep water to release swarmers. Releasing swarmer cells in deeper water may improve the survival chances of juveniles. Study
494:, and cysts have been observed to release these swarmers. Non-encysted cells have also been seen releasing swarmers in laboratory conditions. Not all life cycle stages have been observed, however, and no one has witnessed the fusion of swarmers to produce a new acantharian. Cysts are often found in 364:
The way that the spines are joined at the center of the cell varies and is one of the primary characteristics by which acantharians are classified. The skeletons are made up of either ten diametric or twenty radial spicules. Diametric spicules cross the center of the cell, whereas radial spicules
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and their internal symbiont community does not necessarily match the relative availability of potential symbionts in the surrounding environment. The mismatch between internal and external symbiont communities suggests that acantharians can be selective in choosing symbionts and probably do not
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Adults are usually multinucleated. Earlier diverging clades are able to shed their spines and form cysts, which are often referred to as reproductive cysts. Reproduction is thought to take place by formation of swarmer cells (formerly referred to as "spores"), which may be
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Febvre, Jean; Febvre-Chevalier, Colette (February 1979). "Ultrastructural study of zooxanthellae of three species of Acantharia (Protozoa: Actinopoda), with details of their taxonomic position in the prymnesiales (Prymnesiophyceae, Hibberd, 1976)".
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terminate at the center of the cell where they either form a tight or flexible junction depending on species. Acantharians with diametric spicules or loosely attached radial spicules are able to rearrange or shed spicules and form cysts.
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in terms of lines of latitude and longitude – the spines lie on the intersections between five of the former, symmetric about an equator, and eight of the latter, spaced uniformly. Each line of longitude carries either two
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Many acantharians, including some in clade B (Holacanthida) and all in clades E & F (Symphiacanthida and Arthracanthida), host single-celled algae within their inner cytoplasm (endoplasm). By participating in this
287:, primarily because strontium sulfate is very scarce in seawater and the crystals dissolve after the acantharians die. The arrangement of the spines is very precise, and is described by what is called the 334:
are maintained in the endoplasm. The ectoplasm consists of cytoplasmic extensions used for prey capture and also contains food vacuoles for prey digestion. The ectoplasm is surrounded by a
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crystal. Celestine is named for the delicate blue colour of its crystals, and is the heaviest mineral in the ocean. The denseness of their celestite ensures acantharian shells function as
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Decelle, Johan; Martin, Patrick; Paborstava, Katsiaryna; Pond, David W.; Tarling, Geraint; Mahé, Frédéric; de Vargas, Colomban; Lampitt, Richard; Not, Fabrice (2013-01-11).
1177:"Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies" 672:"Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies" 545:
Polet, S.; Berney, C.; Fahrni, J.; Pawlowski, J. (2004). "Small-subunit ribosomal RNA gene sequences of Phaeodarea challenge the monophyly of Haeckel's Radiolaria".
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strontium sulfate as the main component of their skeletons. However, unlike other radiolarians whose skeletons are made of silica, acantharian skeletons do not
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of these organisms has been hampered mainly by an inability to "close the lifecycle" and maintain these organisms in culture through successive generations.
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Decelle, Johan; Probert, Ian; Bittner, Lucie; Desdevises, Yves; Colin, SĂ©bastien; Vargas, Colomban de; GalĂ­, MartĂ­; SimĂł, Rafel; Not, Fabrice (2012-10-30).
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Martin, Patrick; Allen, John T.; Cooper, Matthew J.; Johns, David G.; Lampitt, Richard S.; Sanders, Richard; Teagle, Damon A. H. (2010).
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Martin, Patrick; Allen, John T.; Cooper, Matthew J.; Johns, David G.; Lampitt, Richard S.; Sanders, Richard; Teagle, Damon A. H. (2010).
1291: 777: 1226:"Paired high-throughput, in situ imaging and high-throughput sequencing illuminate acantharian abundance and vertical distribution" 1155: 485:
Hypothetical scenario of the life cycle in symbiotic and cyst-forming Acantharia with shallow and deep reproduction, respectively.
2532: 1325: 524: 1320: 481: 345:, but arranged into twenty plates, each with a hole through which one spicule projects. The cortex is linked to the spines by 2485: 1404: 1394: 1377: 458:). Clade E & F acantharians have a more specific symbiosis and primarily host symbionts from the haptophyte genus 62: 2498: 356:
control by allowing the ectoplasm to expand and contract, increasing and decreasing the total volume of the cell.
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continuously digest and recruit new symbionts, and maintain symbionts for extended periods of time instead.
2381: 1284: 704: 2519: 1372: 921:"Intra-host Symbiont Diversity and Extended Symbiont Maintenance in Photosymbiotic Acantharea (Clade F)" 202:
that range in size from about 200 microns in diameter up to several millimeters. Some acantharians have
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surrounding each spicule or spine. Acantharians are unique among marine organisms for their ability to
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Symbiotic Holacanthida acantharians host diverse symbiont assemblages, including several genera of
288: 241: 225: 2259: 2169: 2122: 2031: 1682: 1677: 1458: 369: 1389: 1367: 1355: 1330: 1147: 896: 735: 164: 57: 1519: 1109:"Multiple microalgal partners in symbiosis with the acantharian Acanthochiasma sp. (Radiolaria)" 1561: 1463: 1224:
Brisbin, Margaret Mars; Brunner, Otis Davey; Grossmann, Mary Matilda; Mitarai, Satoshi (2020).
2506: 2069: 1785: 1780: 1740: 1636: 1399: 1362: 1350: 1340: 1277: 1255: 1206: 1139: 1089: 1071: 1032: 1014: 985:"Diversity, Ecology and Biogeochemistry of Cyst-Forming Acantharia (Radiolaria) in the Oceans" 960: 942: 888: 844: 826: 773: 562: 395: 280: 272: 233: 191: 2524: 2347: 2164: 1943: 1700: 1314: 1245: 1196: 1131: 1107:
Decelle, Johan; Siano, Raffaele; Probert, Ian; Poirier, Camille; Not, Fabrice (2012-10-27).
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Mars Brisbin, Margaret; Mesrop, Lisa Y.; Grossmann, Mary M.; Mitarai, Satoshi (2018).
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symbionts. Clade F acantharians simultaneously host multiple species and strains of
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Belcher, Anna; Manno, Clara; Thorpe, Sally; Tarling, Geraint (2018).
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Journal of the Marine Biological Association of the United Kingdom
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Material was copied from this source, which is available under a
253: 2379: 1273: 2433: 378:– 20 radial spicules, loosely attached, capable of encystment 394:
The morphological classification system roughly agrees with
319:. The endoplasm, at the core of the cell, contains the main 760:
Decelle, Johan; Not, Fabrice (2015-11-16), "Acantharia",
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have been observed at times in the Iceland Basin and the
797:"An original mode of symbiosis in open ocean plankton" 658:
Creative Commons Attribution 4.0 International License
2388: 2290: 2239: 2178: 2139: 2101: 2090: 2045: 2012: 1994: 1983: 1966: 1921: 1879: 1856: 1849: 1838: 1726: 1691: 1658: 1612: 1601: 1575: 1554: 1547: 1536: 1505: 1472: 1451: 1441: 1428: 403:recently and constitute molecular clades E and F. 801:Proceedings of the National Academy of Sciences 978: 976: 974: 622: 620: 1285: 764:, John Wiley & Sons, Ltd, pp. 1–10, 264:, as much as half of the total gravitational 8: 384:– 20 radial spicules, tight central junction 2376: 2098: 2087: 1991: 1980: 1976: 1963: 1853: 1846: 1835: 1723: 1688: 1655: 1619: 1609: 1598: 1551: 1544: 1533: 1448: 1438: 1425: 1292: 1278: 1270: 390:– 20 radial spines, tight central junction 40: 31: 1249: 1200: 1083: 1026: 1008: 954: 936: 838: 820: 687: 644: 608: 585:"Trace elements in acantharian skeletons" 1056:Communicative & Integrative Biology 508: 446:, Heterocapsa, Scrippsiella, Azadinium 232:Acantharian skeletons are composed of 1170: 1168: 914: 912: 910: 248:, resulting in fast sedimentation to 7: 790: 788: 755: 753: 751: 749: 464:, although they sometimes also host 770:10.1002/9780470015902.a0002102.pub2 627:Le Moigne, FrĂ©dĂ©ric A. C. (2019). 25: 311:is divided into two regions: the 1161:from the original on 2022-10-09. 651: 525:World Register of Marine Species 327:mesh. In symbiotic species, the 190:, distinguished mainly by their 61: 27:Class of single-celled organisms 430:, acantharians are essentially 420:(chlorophyll autofluorescence) 1: 1050:Decelle, Johan (2013-07-30). 418:with symbionts visible in red 1010:10.1371/journal.pone.0053598 194:skeletons. Acantharians are 633:Frontiers in Marine Science 2577: 1230:Limnology and Oceanography 1181:Limnology and Oceanography 676:Limnology and Oceanography 589:Limnology and Oceanography 398:based on the alignment of 2325: 2086: 1979: 1962: 1834: 1722: 1654: 1622: 1597: 1532: 1424: 1308: 1202:10.4319/lo.2010.55.2.0604 1136:10.1007/s13199-012-0195-x 925:Frontiers in Microbiology 885:10.1017/S0025315400046294 732:10.1007/s00227-018-3376-1 689:10.4319/lo.2009.55.2.0604 610:10.4319/lo.1980.25.1.0146 275:crystals are secreted by 240:, in the form of mineral 209:and hence are considered 170: 163: 152: 147: 58:Scientific classification 56: 48: 39: 34: 938:10.3389/fmicb.2018.01998 646:10.3389/fmars.2019.00634 304:spines, in alternation. 822:10.1073/pnas.1212303109 486: 422: 229: 2520:Paleobiology Database 583:Brass, G. W. (1980). 559:10.1078/1434461000164 484: 416:A clade F acantharian 414: 224: 2338:organization type: 1242:2020LimOc..65.2953M 1193:2010LimOc..55..604M 1128:2012Symbi..58..233D 1001:2013PLoSO...853598D 877:1979JMBUK..59..215F 813:2012PNAS..10918000D 807:(44): 18000–18005. 724:2018MarBi.165..117B 601:1980LimOc..25..146B 250:bathypelagic depths 1972:    1968:    487: 423: 396:phylogenetic trees 352:, which assist in 341:, also made up of 230: 2548: 2547: 2507:Open Tree of Life 2382:Taxon identifiers 2373: 2372: 2356:heliozoan amoebae 2348:amoeboflagellates 2312:Labyrinthomyxidae 2286: 2285: 2282: 2281: 2278: 2277: 2274: 2273: 2231:Globigerinitoidea 2082: 2081: 2078: 2077: 1958: 1957: 1954: 1953: 1917: 1916: 1830: 1829: 1826: 1825: 1822: 1821: 1818: 1817: 1814: 1813: 1810: 1809: 1806: 1805: 1741:Phaeogymnocellida 1718: 1717: 1714: 1713: 1650: 1649: 1637:Paracercomonadida 1593: 1592: 1528: 1527: 1251:10.1002/lno.11567 1068:10.4161/cib.24560 273:strontium sulfate 234:strontium sulfate 192:strontium sulfate 177: 176: 143: 16:(Redirected from 2568: 2541: 2540: 2528: 2527: 2515: 2514: 2502: 2501: 2489: 2488: 2476: 2475: 2463: 2462: 2450: 2449: 2437: 2436: 2424: 2423: 2422: 2409: 2408: 2407: 2377: 2165:Silicoloculinida 2099: 2095: 2088: 1992: 1988: 1981: 1977: 1973: 1964: 1944:Plasmodiophorida 1854: 1847: 1843: 1836: 1724: 1701:Thaumatomonadida 1689: 1656: 1620: 1610: 1606: 1599: 1552: 1545: 1541: 1534: 1449: 1439: 1435: 1426: 1337: 1317: 1294: 1287: 1280: 1271: 1264: 1263: 1253: 1221: 1215: 1214: 1204: 1172: 1163: 1162: 1160: 1122:(1–3): 233–244. 1113: 1104: 1098: 1097: 1087: 1047: 1041: 1040: 1030: 1012: 980: 969: 968: 958: 940: 916: 905: 904: 859: 853: 852: 842: 824: 792: 783: 782: 757: 744: 743: 709: 700: 694: 693: 691: 667: 661: 655: 650: 648: 624: 615: 614: 612: 580: 571: 570: 542: 536: 535: 533: 531: 513: 466:Chrysochromulina 455:Chrysochromulina 252:. High settling 156:Acanthophractida 134: 66: 65: 44: 32: 21: 2576: 2575: 2571: 2570: 2569: 2567: 2566: 2565: 2551: 2550: 2549: 2544: 2536: 2531: 2523: 2518: 2510: 2505: 2497: 2492: 2484: 2479: 2471: 2466: 2458: 2453: 2445: 2440: 2432: 2427: 2418: 2417: 2412: 2403: 2402: 2397: 2384: 2374: 2369: 2352:testate amoebae 2321: 2270: 2235: 2214:Globotruncanida 2174: 2135: 2091: 2074: 2041: 2037:Symphyacanthida 2008: 1984: 1967: 1950: 1913: 1889:Claustrosporida 1875: 1839: 1802: 1710: 1687: 1646: 1602: 1589: 1571: 1537: 1524: 1501: 1468: 1429: 1420: 1419: 1335: 1313: 1304: 1298: 1268: 1267: 1223: 1222: 1218: 1174: 1173: 1166: 1158: 1111: 1106: 1105: 1101: 1049: 1048: 1044: 982: 981: 972: 918: 917: 908: 861: 860: 856: 794: 793: 786: 780: 759: 758: 747: 707: 702: 701: 697: 669: 668: 664: 626: 625: 618: 582: 581: 574: 544: 543: 539: 529: 527: 515: 514: 510: 505: 479: 439:dinoflagellates 421: 419: 417: 409: 382:Symphiacanthida 362: 256:of acantharian 246:mineral ballast 239: 219: 183:are a group of 142:Mikrjukov, 2000 133: 60: 28: 23: 22: 15: 12: 11: 5: 2574: 2572: 2564: 2563: 2553: 2552: 2546: 2545: 2543: 2542: 2529: 2516: 2503: 2490: 2477: 2464: 2451: 2438: 2425: 2410: 2394: 2392: 2386: 2385: 2380: 2371: 2370: 2368: 2367: 2360:sporozoan-like 2336: 2333: 2326: 2323: 2322: 2320: 2319: 2317:Rhizoplasmidae 2314: 2309: 2307:Gymnosphaerida 2304: 2296: 2294: 2292:Incertae sedis 2288: 2287: 2284: 2283: 2280: 2279: 2276: 2275: 2272: 2271: 2269: 2268: 2263: 2256: 2251: 2245: 2243: 2241:Incertae sedis 2237: 2236: 2234: 2233: 2228: 2226:Guembelitriida 2222: 2220:Heterohelicida 2216: 2210: 2205: 2200: 2195: 2190: 2184: 2182: 2176: 2175: 2173: 2172: 2167: 2162: 2157: 2152: 2145: 2143: 2137: 2136: 2134: 2133: 2126: 2120: 2118:Xenophyophorea 2115: 2108: 2106: 2096: 2084: 2083: 2080: 2079: 2076: 2075: 2073: 2072: 2067: 2062: 2057: 2051: 2049: 2043: 2042: 2040: 2039: 2034: 2029: 2027:Chaunocanthida 2024: 2022:Arthracanthida 2018: 2016: 2010: 2009: 2007: 2006: 2000: 1998: 1989: 1974: 1960: 1959: 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1418: 1417: 1416: 1415: 1414: 1413: 1410:Mesomycetozoea 1407: 1402: 1392: 1382: 1381: 1380: 1375: 1370: 1365: 1360: 1359: 1358: 1346:Diaphoretickes 1343: 1338: 1333: 1328: 1323: 1318: 1310: 1309: 1306: 1305: 1303:classification 1299: 1297: 1296: 1289: 1282: 1274: 1266: 1265: 1216: 1187:(2): 604–614. 1164: 1099: 1042: 970: 906: 871:(1): 215–226. 854: 784: 778: 745: 712:Marine Biology 695: 682:(2): 604–614. 662: 616: 595:(1): 146–149. 572: 537: 507: 506: 504: 501: 496:sediment traps 478: 475: 428:photosymbiosis 415: 408: 405: 392: 391: 388:Arthracanthida 385: 379: 376:Chaunacanthida 373: 361: 358: 296:spines or one 266:organic carbon 262:Southern Ocean 237: 218: 215: 204:photosynthetic 175: 174: 168: 167: 161: 160: 159: 158: 150: 149: 145: 144: 128: 124: 123: 118: 114: 113: 108: 104: 103: 98: 91: 90: 88:Diaphoretickes 85: 78: 77: 72: 68: 67: 54: 53: 46: 45: 37: 36: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 2573: 2562: 2559: 2558: 2556: 2539: 2534: 2530: 2526: 2521: 2517: 2513: 2508: 2504: 2500: 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588: 553:(1): 53–63. 550: 546: 540: 528:. Retrieved 520: 517:"Acantharia" 511: 488: 469: 465: 459: 453: 444:Pelagodinium 442: 436: 424: 393: 370:Holacanthida 363: 343:microfibrils 306: 301: 297: 293: 270: 231: 180: 178: 171: 139: 130: 94: 81: 49: 29: 2455:iNaturalist 2414:Wikispecies 2364:fungus-like 2340:flagellates 2203:Carterinida 2198:Robertinida 2065:Spumellaria 2060:Nassellaria 1939:Phagomyxida 1922:Proteomyxia 1899:Mikrocytida 1497:Limnofilida 1452:Skiomonadea 470:Phaeocystis 461:Phaeocystis 347:contractile 336:periplasmic 325:microfibril 185:radiolarian 172:Acantharea 117:Subphylum: 35:Acantharia 18:Acantharian 2561:Acantharea 2420:Acantharea 2390:Acantharia 2266:Komokiacea 2150:Fusulinida 2055:Collodaria 2014:Acantharea 2004:Taxopodida 1986:Radiolaria 1934:Phytomyxea 1909:Paramyxida 1904:Paradinida 1736:Phaeodarea 1706:Euglyphida 1660:Imbricatea 1632:Katabiidae 1520:Minorisida 718:(7): 117. 503:References 492:flagellate 477:Life cycle 450:haptophyte 432:mixotrophs 321:organelles 298:equatorial 217:Morphology 211:mixotrophs 181:Acantharia 131:Acantharia 121:Radiolaria 50:Acantharea 2193:Rotaliida 2155:Miliolida 1858:Gromiidea 1850:Marimyxia 1772:Eothecia 1668:Krakenida 1585:Helkesida 1562:Metopiida 1464:Tremulida 1390:Amoebozoa 1368:Alveolata 1356:Cryptista 1331:Eukaryota 1260:1939-5590 1211:1939-5590 1144:0334-5114 1116:Symbiosis 1076:1942-0889 1019:1932-6203 947:1664-302X 893:1469-7769 831:0027-8424 530:13 August 407:Symbiosis 332:symbionts 317:ectoplasm 313:endoplasm 309:cytoplasm 307:The cell 285:fossilize 242:celestine 226:Celestine 75:Eukaryota 2555:Category 2399:Wikidata 2335:†extinct 2254:Lagenida 2070:Orodaria 1866:Gromiida 1841:Endomyxa 1786:Matazida 1781:Ebriacea 1577:Helkesea 1548:Eoglissa 1432:Cercozoa 1385:Amorphea 1363:Rhizaria 1351:Hacrobia 1341:Excavata 1326:Bacteria 1301:Rhizaria 1156:Archived 1094:23986805 1037:23326463 989:PLOS ONE 965:30210473 931:: 1998. 901:86040570 849:23071304 740:90349921 567:15144058 448:) and a 360:Taxonomy 354:buoyancy 350:myonemes 315:and the 300:and two 294:tropical 277:vacuoles 188:protozoa 165:Synonyms 138:, 1881, 107:Phylum: 71:Domain: 52:species 2405:Q135584 2344:amoebae 1970:Retaria 1400:Animals 1321:Archaea 1238:Bibcode 1189:Bibcode 1152:6142181 1124:Bibcode 1085:3742057 1028:3543462 997:Bibcode 956:6120437 873:Bibcode 840:3497740 809:Bibcode 720:Bibcode 597:Bibcode 547:Protist 228:crystal 198:marine 136:Haeckel 127:Class: 111:Retaria 2538:586732 2460:424296 1434:  1430:  1378:Plants 1315:Domain 1258:  1209:  1150:  1142:  1092:  1082:  1074:  1035:  1025:  1017:  963:  953:  945:  899:  891:  847:  837:  829:  776:  738:  565:  339:cortex 268:flux. 254:fluxes 236:, SrSO 148:Order 140:emend. 2533:WoRMS 2512:45752 2499:65574 2486:46107 2468:IRMNG 1405:Fungi 1159:(PDF) 1148:S2CID 1112:(PDF) 897:S2CID 736:S2CID 708:(PDF) 521:WoRMS 329:algal 302:polar 258:cysts 95:Clade 82:Clade 2494:NCBI 2481:ITIS 2473:1050 2442:GBIF 2434:4876 1256:ISSN 1207:ISSN 1140:ISSN 1090:PMID 1072:ISSN 1033:PMID 1015:ISSN 961:PMID 943:ISSN 889:ISSN 845:PMID 827:ISSN 774:ISBN 563:PMID 532:2024 271:The 179:The 2525:717 2447:234 2429:EoL 1246:doi 1197:doi 1132:doi 1080:PMC 1064:doi 1023:PMC 1005:doi 951:PMC 933:doi 881:doi 835:PMC 817:doi 805:109 766:doi 762:eLS 728:doi 716:165 684:doi 641:doi 605:doi 555:doi 551:155 101:SAR 2557:: 2535:: 2522:: 2509:: 2496:: 2483:: 2470:: 2457:: 2444:: 2431:: 2416:: 2401:: 2362:; 2358:; 2354:; 2350:; 2346:; 2342:; 1254:. 1244:. 1234:65 1232:. 1228:. 1205:. 1195:. 1185:55 1183:. 1179:. 1167:^ 1154:. 1146:. 1138:. 1130:. 1120:58 1118:. 1114:. 1088:. 1078:. 1070:. 1058:. 1054:. 1031:. 1021:. 1013:. 1003:. 991:. 987:. 973:^ 959:. 949:. 941:. 927:. 923:. 909:^ 895:. 887:. 879:. 869:59 867:. 843:. 833:. 825:. 815:. 803:. 799:. 787:^ 772:, 748:^ 734:. 726:. 714:. 710:. 680:55 678:. 674:. 639:. 635:. 631:. 619:^ 603:. 593:25 591:. 587:. 575:^ 561:. 549:. 523:. 519:. 213:. 97:: 84:: 2366:. 2329:* 2224:† 2218:† 2212:† 2148:† 2125:* 2114:* 2105:* 1616:* 1445:* 1412:) 1293:e 1286:t 1279:v 1262:. 1248:: 1240:: 1213:. 1199:: 1191:: 1134:: 1126:: 1096:. 1066:: 1060:6 1039:. 1007:: 999:: 993:8 967:. 935:: 929:9 903:. 883:: 875:: 851:. 819:: 811:: 768:: 742:. 730:: 722:: 692:. 686:: 660:. 649:. 643:: 637:6 613:. 607:: 599:: 569:. 557:: 534:. 452:( 441:( 238:4 20:)

Index

Acantharian

Scientific classification
Edit this classification
Eukaryota
Diaphoretickes
SAR
Retaria
Radiolaria
Acantharia
Haeckel
Acanthophractida
Synonyms
radiolarian
protozoa
strontium sulfate
heterotrophic
microplankton
photosynthetic
endosymbionts
mixotrophs

Celestine
strontium sulfate
celestine
mineral ballast
bathypelagic depths
fluxes
cysts
Southern Ocean

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