Arctica islandica: The longest lived non colonial animal known to science

Reviews in Fish Biology and Fisheries

Volumn 21, Issue 3, 2011, Pages 297-310

  Ridgway, I D ^a   Richardson, C A ^a  

^a BANGOR UNIVERSITY   (United Kingdom)

Author keywords

Arctica islandica; Bivalves; Model organism; Negligible senescence

Indexed keywords

ABUNDANCE; AGING; ANTIOXIDANT; BIOACCUMULATION; BIVALVE; ENVIRONMENTAL CHANGE; LIFE EXPECTANCY; MOLLUSC; PHYSIOLOGY;

ANIMALIA; ARCTICA ISLANDICA; BIVALVIA; MERCENARIA;

EID: 79961127788     PISSN: 09603166     EISSN: None     Source Type: Journal    
DOI: 10.1007/s11160-010-9171-9     Document Type: Review

References (128)

1. Abele D (2002) Toxic oxygen: the radical life-giver. Nature 420: 27.
2. Abele D, Puntarulo S (2004) Formation of reactive species and induction of antioxidant defense systems in polar and temperate marine invertebrates and fish. Comp Biochem Physiol 138A(4): 405-415.
3. Abele D, Strahl J, Brey T, Philipp EER (2008) Imperceptible senescence: ageing in the ocean quahog Arctica islandica. Free Rad Res 42(5): 474-480.
4. Anisimova AA (2007) Genome sizes of some Bivalvia species of the Peter the Great Bay of the Sea of Japan. Comparative Cytogenetics 1: 63-69.
5. Ansell AD, Lander KF (1967) Studies on the hard-shell clam, Venus mercenaria, in British waters. III. Further observations on the seasonal biochemical cycle and on spawning. J Appl Ecol 4: 425-435.
6. Austad SN (2001) An experimental paradigm for the study of slowly-ageing organisms. Exp Gerontol 36: 599-605.
7. Austad SN (2005) Diverse ageing rates in metazoans: targets for functional genomics. Mech Age Dev 126: 43-49.
8. Baker GT, Sprott RL (1988) Biomarkers of ageing. Exp Gerontol 23: 223-239.
9. Barber BJ (2004) Neoplastic diseases of commercially important marine bivalves. Aquat Living Resour 17: 449-466.
10. Barber BJ, MacCallum GS, Robinson SMC, McGladdery SE (2002) Occurrence and lack of transmissibility of gonadal neoplasia in softshell clams, Mya arenaria, in Maine (USA) and Atlantic Canada. Aquat Living Resour 15: 319-326.
11. Barja G (2002) Rate of generation of oxidative stress-related damage and animal longevity. Free Radic Biol Med 33: 1167-1172.
12. Bauer G (1992) Variation in life span and size of the freshwater pearl mussel. J Anim Ecol 61: 425-436.
13. Bergman MJN, Fonds M, Hup M, Lewis W, van der Puyl A, Sanl A, den Uyl D (1990) Direct effect of beam trawl fishing on benthic fauna in the North Sea-a pilot study. Netherlands Inst Sea Res Rep 8: 33-57.
14. Bert TM, Hesselman DM, Arnold WS, Moore WS, Cruz-Lopez H, Marelli DC (1993) High frequency of gonadal neoplasia in a hard clam (Mercenaria spp.) hybrid zone. Mar Biol 117: 97-104.
15. Bluhm BA, Brey T (2001) Age determination in the Antartic shrimp Notocrangon antarcticus (Crustacea: Decapoda), using the autofluorescent pigment lipofuscin. Mar Biol 138: 247-257.
16. Boore JL, Medina M, Rosenberg LA (2004) Complete sequences of the highly re-arranged molluscan mitochondrial genomes of the scaphopod Graptacme eborea and the bivalve Mytilus edulis. Mol Biol Evol 21(8): 1492-1503.
17. Breton S, Burger G, Stewart DT, Blier PU (2006) Comparative analysis of gender-associated complete mitochondrial genomes in marine mussels (Mytilus spp.). Genetics 172(2): 1107-1119.
18. Brey T, Arntz WE, Pauly D, Rumohr H (1990) Arctica (Cyprina) islandica in Kiel Bay (western Baltic): growth, production and ecological significance. J Exp Mar Biol Ecol 136: 217-235.
19. Brousseau DJ (1987) Seasonal aspects of sarcomatous neoplasia in Mya arenaria (soft-shell clam) from Long Island Sound. J Invertebr Pathol 50: 269-276.
20. Brown RS, O'Toole CJ (1978) Histochemical analyses of pigment accumulations in Mercenaria mercenaria L. and Mya arenaria L. Proc Natl Shellfish Ass Md 68: 75-76.
21. Browne RA, Russel-Hunter WD (1978) Reproductive effort in molluscs. Oecologia 37: 23-27.
22. Brunk UT, Terman A (2002) The mitochondrial-lysosomal axis theory of ageing: accumulation of damaged mitochondria as a result of imperfect autophagocytosis. Eur J Biochem 269: 1996-2002.
23. Buffenstein R (2005) The Naked Mole-Rat: a new long-living model for human ageing research. J Gerontol 60A(11): 1369-1377.
24. Byrne PA, O'Halloran J (2001) The role of bivalve molluscs as tools in estuarine sediment toxicity testing: a review. Hydrobiologia 465(1-3): 209-217.
25. Chow DK, Glenn CF, Johnston JL (2006) Sarcopenia in the Caenorhabditis elegans pharynx correlates with muscle contraction rate over lifespan. Exp Gerontol 41: 252-260.
26. Comfort A (1964) Ageing: the biology of senescence. Routledge and Kegan Paul Ltd, London.
27. Cornet M (2006) Primary mantle tissue culture from the bivalve mollusc Mytilus galloprovincialis: investigations on the growth promoting activity of the serum used for medium supplementation. J Biotechnol 123: 78-84.
28. de Magalhães JP (2006) Species selection in comparative studies of ageing and antiageing research. In: Conn PM (ed) Handbook of models for human ageing. Elsevier Academic Press, Burlington, pp 9-20.
29. de Magalhães JP, Cabral JA, Magalhães D (2005) The influence of genes on the ageing process of mice: a statistical assessment of the genetics of ageing. Genetics 169: 265-274.
30. DFO (2007) Assessment of the Ocean Quahog (Arctica islandica) stocks on Sable Bank and St. Mary's Bay, and the Arctic surf clam (Mactromeris polynyma) Stock on Banquereau. DFO Can Sci Advis Sec Sci Advis Rep 2007/034.
31. Doonan R, McElwee JJ, Matthijssens F, Walker GA, Houthoofd K, Back P, Matscheski A, Vanfleteren JR, Gems D (2008) Against the oxidative damage theory of ageing: superoxide dismutases protect against oxidative stress but have little or no effect on life span in Caenorhabditis elegans. Genes Dev 22: 3236-3241.
32. Dorrens J, Rennie MJ (2003) Effects of ageing and human whole body and muscle protein turnover. Scand J Med Sci Sports 13: 26-33.
33. Fajkus J, Simickova M, Malaska J (2002) Tiptoeing to chromosome tips: facts, promises and perils of today's human telomere biology. Philos Trans R Soc Lond B 357: 545-562.
34. Finch CE (1990) Longevity, senescence and the genome. Chicago University Press, Chicago.
35. Fisher AL (2004) Of worms and women: Sarcopenia and its role in disability and mortality. J Am Geriatr Soc 52: 1185-1190.
36. Forster GR (1981) A note on the growth of Arctica islandica. J Mar Biol Assoc UK 61: 817-817.
37. Galinou-Mitsoudi S, Sinis AI (1994) Reproductive cycle and fecundity of the date mussel Lithophaga lithophaga (bivalvia: mytilidae). J Moll Stud 60: 371-385.
38. Garigan D, Hsu AL, Fraser AG, Kamath RS, Ahringer J, Kenyon C (2002) Genetic analysis of tissue ageing in Caenorhabditis elegans: a role for heat-shock factor and bacterial proliferation. Genetics 161: 1101-1112.
39. Gershon H, Gershon D (2002) Review: Caenorhabditis elegans-a paradigm for ageing research: advantages and limitations. Mech Ageing Dev 123: 261-274.
40. Goldberg ED (1975) The mussel watch programme-a first step in global marine monitoring. Mar Poll Bull 6: 111.
41. Gosling E (2003) Bivalve Molluscs: biology, ecology and culture. Wiley, USA.
42. Gray DA, Woulfe J (2005) Lipofuscin and aging: a matter of toxic waste. Sci Aging Knowl Environ 2005(5): 1.
43. Hammer C, Braum E (1988) Mini-review: quantification of age pigments (lipofuscin). Comp Biochem 90B: 7-17.
44. Harman D (1956) Ageing: a theory based on free radical and radiation biology. J Gerontol 11: 298-300.
45. Haussmann MF, Vleck CM (2002) Telomere length provides a new technique for ageing animals. Oecologia 130: 325-328.
46. Herndon LA, Schmeissner PJ, Dudaronek JM, Brown PA, Listner KM, Sakano Y, Paupard MC, Hall DH, Driscoll M (2002) Stochastic and genetic factors influence tissue-specific decline in ageing Caenorhabditis elegans. Nature 419: 808-814.
47. Hill KT, Womersley C (1991) Critical aspects of fluorescent age-pigment methodologies: modification for accurate analysis and age assessments in aquatic organisms. Mar Biol 109: 1-11.
48. Holmes DJ, Flückiger R, Austad RN (2001) Comparative biology of birds: an update. Exp Gerontol 36: 883-896.
49. Holmes SP, Witbaard R, van de Meer J (2003) Phenotypic and genotypic population differentiation in the bivalve mollusc Arctica islandica (L.): results from RAPD analysis. Mar Ecol Prog Ser 254: 163-176.
50. Huang T, Carlson E, Gillespie A, Shi Y, Epstein C (2000) Ubiquitous overexpression of Cu-Zn superoxide dismutase does not extend life span in mice. J Gerontol 55: B5-B9.
51. Hughes KA, Reynolds RM (2005) Evolutionary and mechanistic theories of ageing. Ann Rev Entomol 50: 421-445.
52. Ivanina AV, Sokolova IM, Sukhotin AA (2008) Oxidative stress and expression of chaperones in ageing mollusks. Comp Biochem Physiol 150B(1): 53-61.
53. Jolly RD, Palmer DN, Dalefield RR (2002) The analytical approach to the nature of lipofuscin (age pigment). Arch Gerontol Geriatr 34: 205-218.
54. Jones DS (1983) Sclerochronology: reading the record of the molluscan shell. Am Sci 71: 384-391.
55. Kapahi P, Boulton ME, Kirkwood TBL (1999) Positive correlation between mammalian lifespan and cellular resistance to stress. Free Radical Biol Med 26: 495-500.
56. Kennish MJ, Lutz RA (1995) Assessment of the ocean quahog, Arctica islandica (Linnaeus, 1767), in the New Jersey fishery. J Shellfish Res 14: 45-52.
57. Kilada RW, Campana SE, Roddick D (2007) Validated age, growth, and mortality estimates of the ocean quahog (Arctica islandica) in the western Atlantic. ICES J Mar Sci 64: 31-38.
58. Kowald A, Lehrach H, Klipp E (2006) Alternative pathways as mechanism for the negative effects associated with overexpression of superoxide dismutase. J Theor Biol 238: 828-840.
59. Krishnakumar PK, Asokan PK, Pillai VK (1990) Physiological and cellular responses to copper and mercury in the green mussel Perna viridis (Linnaeus). Aquat Toxicol 18: 163-174.
60. Lomovasky BJ, Morriconi E, Brey T, Calvo J (2002) Individual age and connective tissue lipofuscin in the hard clam Eurhomalea exalbida. J Exp Mar Biol Ecol 276: 83-94.
61. Loosanoff VL (1953) Reproductive cycle in Cyprina islandica. Biol Bull 104: 146-155.
62. Lutz RA, Mann R, Goodsell JG, Castagna M (1982) Larval and early post-larval development of Arctica islandica. J Mar Biol Assoc UK 62: 45-769.
63. Mann R (1982) The seasonal cycle of gonadal development in Arctica islandica from the southern New England shelf. Fish Bull US 80: 315-326.
64. Mann R, Wolf C (1983) Swimming behaviour of larvae of the ocean quahog Arctica islandica in response to pressure and temperature. Mar Ecol Prog Ser 13: 211-218.
65. Mathew S, Damodaran R (1997) Lipofuscin as physiological indicator of heavy metal stress in Sunetta scripta (yellow clam) and Perna viridis (green mussel). Indian J Mar Sci 26(1): 64-67.
66. Medvedev ZA (1990) An attempt at a rational classification of theories of ageing. Biol Rev Camb Philos Soc 65: 375-398.
67. Miller RA (2001) Genetics of increased longevity and retarded ageing in mice. In: Masoro EJ, Austad SN (eds) Handbook of the biology of ageing, 5th ed. Academic Press, New York, pp 369-395.
68. Mockett RJ, Sohal RS, Orr WC (1999) Overexpression of glutathione reductase extends survival in transgenic Drosophila melanogaster under hyperoxia but not normoxia. FASEB J 13: 1733-1742.
69. Mockett RJ, Bayne AC, Kwong LK, Orr WC, Sohal RS (2003) Ectopic expression of catalase in Drosophila mitochondria increases stress resistance but not longevity. Free Radic Biol Med 34: 207-217.
70. Muller FL, Lustgarten MS, Jang Y, Richardson A, van Remmen H (2007) Trends in oxidative ageing theories. Free Rad Biol Med 43: 477-503.
71. Murawski SA, Serchuk FM (1989) Environmental effects of offshore dredge fisheries for bivalves. ICES CM 1989/k: 27, 12 pp.
72. Murawski SA, Ropes JW, Serchuk FM (1982) Growth of the ocean quahog, Arctica islandica, in the Middle Atlantic Bight. Fish Bull US 80: 21-34.
73. Murawski SA, Azarovitz TR, Radosh DJ (1989) Long term biological effects of hypoxic water conditions off New Jersey, USA 1976-1989. ICES/CM 1989 E: 11, 22 pp.
74. NEFSC (1995) 19th northeast regional stock assessment workshop (19th SAW): stock assessment review committee (SARC) consensus summary of assessments. Northeast Fish. Sci. Cent. Ref. Doc. 95-08. 221 pp.
75. Nicol D (1951) Recent species of the veneroid pelecypod Arctica. J Wash Acad Sci 41: 102-106.
76. Oeschger R (1990) Long-term anaerobiosis in sublittoral marine invertebrates from the western Baltic Sea: Halicryptus spinulosus (Priapulida), Astarte borealis and Arctica islandica (Bivalvia). Mar Ecol Prog Ser 59: 133-143.
77. Oeschger R, Storey KB (1993) Impact of anoxia and hydrogen sulphide on the metabolism of Arctica islandica L. (Bivalvia). J Exp Mar Biol Ecol 170: 213-226.
78. Omar BA, Gad NM, Jordan MC, Striplin SP, Russell WJ, Downey JM, McCord JM (1990) Cardioprotection by Cu, Zn-superoxide dismutase is lost at high doses in the reoxygenated heart. Free Radic Biol Med 9: 465-471.
79. Peters EC, Yevich PP, Harshbarger JC, Zaroogian GE (1994) Comparative histology of gonadal neoplasms in marine bivalve molluscs. Dis Aquat Org 20: 59-76.
80. Peterson CH (1986) Quantitaitive allometry of gamete production by Mercenaria mercenaria into old age. Mar Ecol Prog Ser 29: 93-97.
81. Philipp E, Brey T, Pörtner HO, Abele D (2005) Chronological and physiological ageing in a polar and a temperate mud clam. Mech Ageing Dev 126: 589-609.
82. Philipp E, Heilmayer O, Brey T, Abele D, Pörtner HO (2006) Physiological ageing in a polar and a temperate swimming scallop. Mar Ecol Prog Ser 307: 187-198.
83. Plohl M, Prats E, Marti{dotless}nez-Lage A, Gonzalez-Tizon A, Mendez J, Cornudella L (2002) Telomeric localization of the vertebrate-type hexamer repeat, (TTAGGG)n, in the Wedgeshell Clam Donax trunculus and other marine invertebrate genomes. J Biol Chem 277(22): 19839-19846.
84. Poder M, Auffret M (1986) Sarcomatous lesion in the cockle Cerastoderma edule I. Morphology and population survey in Brittany, France. Aquaculture 58: 1-8.
85. Powell E, Mann R (2005) Evidence of recent recruitment in the ocean quahog Arctica islandica in the Mid-Atlantic bight. J Shellfish Res 24(2): 517-530.
86. Rees EIS, Nicholaidou A, Laskaridou P (1977) The effects of storms on the dynamics of shallow water associations. In: Keegan BF, Ceidigh PO, Boaden PJS (eds) Biology of benthic organisms. Pergamon, New York, pp 465-474.
87. Reid RGB, Fankboner PV, Brand DG (1984) Studies of the physiology of the giant clam Tridacna gigas Linne: 2. Kidney function. Comp Biochem Physiol 78A(1): 103-108.
88. Richardson CA (2001) Molluscs as archives of environmental change. Oceanogr Mar Biol Ann Rev 39: 103-164.
89. Rinkevich B (1999) Marine invertebrate cell cultures: new millennium trends. Mar Biotechnol 7: 429-439.
90. Ropes JW (1985) Modern methods used to age oceanic bivalves. Nautilus 99: 53-57.
91. Ropes JW (1988) Ocean quahog Arctica islandica: p. 129-132. In: Penttila J, Dery LM (eds) Age determination methods for Northwest Atlantic Species. NOAA technical report NMFS 72, 136 pp.
92. Ropes JW, Murawski SA (1983) Maximum shell length and longevity in ocean quahogs, Arctica islandica Linné. ICES C. M. 1983/K: 32. 8 p.
93. Roubenoff R, Hughes VA (2000) REVIEW ARTICLE: Sarcopenia: current concepts. J Gerontol 55A(12): M716-M724.
94. Rumohr H, Krost P (1991) Experimental-evidence of damage to benthos by bottom trawling with special reference to Arctica Islandica. Meer Rep Mar Res 33: 340-345.
95. Sakai M, Okumura S, Yamamori K (2005) Telomere analysis of pacific abalone Haliotis discus hannai chromosomes by fluorescence in situ hybridization. J Shellfish Res 24(4): 1149-1151.
96. Sarasquete MC, Gonzales de Canales ML, Gimeno S (1992) Comparative histopathological alterations in the digestive gland of marine bivalves exposed to Cu and Cd. Eur J Histochem 36(2): 223-232.
97. Schafer W (1972) Ecology and Palaeoecology of marine environments. University of Chicago Press, Chicoago 568 pp.
98. Schöne BR, Fiebig J, Pfeiffer M, Gleh R, Hickson J, Johnson ALA, Dreyer W, Oschmann W (2005) Climate records from a bivalved Methuselah (Arctica islandica, Mollusca; Iceland). Palaeogeogr Palaeoclimatol Palaeoecol 228: 130-148.
99. Scourse J, Richardson C, Forsythe G, Harris I, Heinemeier J, Fraser N, Briffa K, Jones P (2006) First cross-matched floating chronology from the marine fossil record: data from growth lines of the long-lived bivalve mollusc Arctica Islandica. Holocene 16: 967-974.
100. Sheehy MRJ (1996) Quantitative comparison of in situ lipofuscin concentration with soluble autofluorescence intensity in the crustacean brain. Exp Gerontol 31: 421-432.
101. Sheehy MRJ (2002) A flow-cytometric method for neurolipofuscin quantification and comparison with existing histological and biochemical approaches. Arch Gerontol Geriatr 34: 233-248.
102. Sheehy MRJ, Roberts BE (1991) An alternative explanation for anomalies in "soluble lipofuscin" fluorescence data from insects, crustaceans, and other aquatic species. Exp Gerontol 26: 495-509.
103. Sheehy MRJ, Bannister RCA, Wickins JF, Shelton PMJ (1999) New perspectives on the growth and longevity of the European lobster (Homarus gammarus). Can J Fish Aquat Sci 56: 1904-1915.
104. Sohal RS (1981) In: Sohal RS (ed) Age pigments. Elsevier, Amsterdam, p 394.
105. Sohal RS, Brunk UT (1989) Lipofuscin as an indicator of oxidative stress and ageing. Adv Exp Med Biol 266: 17-26.
106. Sohal RS, Weindruch R (1996) Oxidative stress caloric restriction, ageing. Science 273: 59-63.
107. Speakman JR (2005) Body size, energy metabolism and lifespan. J Exp Biol 208: 1717-1730.
108. Strahl J, Philipp E, Brey T, Broeg K, Abele D (2007) Physiological ageing in the Icelandic population of the ocean quahog Arctica islandica. Aquat Biol 1: 77-83.
109. Strehler BL (1986) Genetic instability as the primary cause of human ageing. Exp Gerontol 21: 283-319.
110. Sukhotin AA, Abele D, Pörtner HO (2002) Growth, metabolism and lipid peroxidation in Mytilus edulis L.: age and size effects. Mar Ecol Prog Ser 226: 223-234.
111. Taylor AC (1976) Burrowing behavior and anaerobiosis in the bivalve Arctica islandica (L.). J Mar Biol Assoc UK 56: 95-109.
112. Thompson I, Jones DS, Deribelbis D (1980a) Annual internal growth banding and life history of the ocean quahog Arctica islandica (Mollusca: Bivalvia). Mar Biol 57: 25-34.
113. Thompson I, Jones DS, Ropes JW (1980b) Advanced age for sexual maturity in the ocean quahog Arctica islandica (Mollusca: Bivalvia). Mar Biol 57: 35-39.
114. Thórarinsdóttir GG (2000) Annual gametogenic cycle in ocean quahog, Arctica islandica from north-western Iceland. J Mar Biol Assoc UK 80: 661-666.
115. Thórarinsdóttir GG, Einarsson ST (1996) Distribution, abundance, population structure and meat yield of the ocean quahog, Arctica islandica, in Icelandic waters. J Mar Biol Assoc UK 76: 1107-1114.
116. Vaupel JW, Baudischa A, Döllinga M, Roach DA, Gampe J (2004) The case for negative senescence. Theor Pop Biol 65(4): 339-351.
117. Viarengo A, Canesi L, Pertica M, Poli G, Moore MN, Orunesu M (1990) Heavy metal effects on lipid peroxidation in the tissues of Mytilus galloprovincialis L. Comp Biochem Physiol 97C(1): 37-42.
118. Vleck CM, Haussmann MF, Vleck D (2003) The natural history of telomeres: tools for ageing animals and exploring the ageing process. Exp Gerontol 38: 791-795.
119. Wanamaker AD Jr, Heinemeier J, Scourse JD, Richardson CA, Butler PG, Eiríksson J, Knudsen KL (2008) Very long-lived molluscs confirm 17th century AD tephra-based radiocarbon reservoir ages for north Icelandic shelf waters. Radiocarbon 50(3): 1-14.
120. Weigelt M (1991) Short-and long-term changes in the benthic community of the deeper parts of Kiel Bay (Western Baltic) due to oxygen depletion and eutrophication. Meerestorsch 33: 197-224.
121. Weinert BT, Timiras PS (2003) Invited review: theories of ageing. J Appl Physiol 95: 1706-1716.
122. Wilson DS, Tracy CR, Tracy CR (2003) Estimating age of turtles from growth rings: a critical evaluation of the technique. Herpetologica 59(2): 178-194.
123. Winter JE (1969) Über den Einfluß der Nahrungskonzentration und anderer Faktoren auf Filtrierleistung und Nahrungsausnutzung der Muscheln Arctica islandica und Modiolus modiolus. Mar Biol 4: 87-135 (In German; English abstract).
124. Witbaard R, Jenness MI, van der Borg K, Ganssen G (1994) Verification of annual growth increments in Arctica islandica L. from the North Sea by means of oxygen and carbon isotopes. Neth J Sea Res 33: 91-101.
125. Witbaard R, Duineveld GCA, De Wilde PAWJ (1997) A long-term growth record derived from A. islandica (Mollusca, Bivalvia) from the Fladen ground (northern North Sea). J Mar Biol Assoc UK 77: 801-816.
126. Witbaard R, Duineveld GCA, de Wilde PAWJ (1999) Geographical differences in growth rates of Arctica islandica (Mollusca: Bivalvia) from the North Sea and adjacent waters. J Mar Biol Assoc UK 907-915.
127. Zatsepin VI, Filatova ZA (1961) The bivalve mollusc Cyprina islandica (L.), its geographical distribution and role in communities of benthic fauna. Trans Inst Ocean Acad Sci USSR 46: 201-216.
128. Ziuganov V, Miguel ES, Neves RJ, Longa A, Fernandez C, Amaro R, Beletsky V, Popkovitch E, Kaliuzhin C, Johnson T (2000) Life span variation of the freshwater pearl shell: a model species for testing longevity mechanisms in animals. Ambio J Hum Environ 29: 102-105.