摘 要

每逢季節交替，溫鹽度變化太大或是環境不適時，文蛤Meretrix lusoria會採取閉殼的方式，由於文蛤為底棲性生物，不易由其行為表現來推測其健康狀態，所以本研究探討無氧環境對文蛤血淋巴酸-鹼平衡的影響，對斧足、血淋巴和無氧培育水中無氧代謝物（琥珀酸、丙胺酸和丙酸）含量的影響以及對斧足中丙酮酸激酶（PK）、丙胺酸氨基轉移酶（AlaT）、磷酸烯醇丙酮酸羧化酶（PEPCK）、延胡索酸還原酶（FR）和合成丙酸酵素活性的影響，希望可以作為文蛤活力的指標。
文蛤暴露於23和30℃空氣中1分鐘，其血淋巴中pO2的含量約下降50﹪，而文蛤暴露於15℃空氣中2分鐘，其血淋巴中pO2也顯著地下降75﹪。文蛤在28℃無氧的環境下2小時，血淋巴中pO2下降60﹪，當無氧處理延長為71小時，其血淋巴pH值顯著地下降，然而pCO2則上升。文蛤在含有597 ppm鉀離子的環境下3小時，其血淋巴pO2下降23﹪。而文蛤在含有100 ppm KCN和缺乏鎂離子的環境下10小時，其血淋巴pO2也分別下降39﹪和42﹪。
文蛤在無氧的環境下，其斧足、血淋巴和無氧培育水中主要的無氧代謝物分別為丙胺酸、琥珀酸以及琥珀酸和丙酸。在斧足中的丙胺酸以及血淋巴中的琥珀酸和丙酸隨著無氧時間的延長而顯著地增加，然而無氧培育水中琥珀酸的含量在無氧的後期逐漸地下降。文蛤在30℃無氧的環境下，僅能存活48小時，然而添加5 ppm的氯黴素可以延長文蛤在無氧環境下存活120小時以上，同時發現氯黴素對文蛤無氧代謝物累積量的影響不顯著。文蛤在缺乏鎂離子的環境下，其斧足中丙胺酸和琥珀酸以及血淋巴中丙胺酸、琥珀酸和丙酸的含量隨著培育時間的延長而顯著地增加，然而在斧足中並沒有測到丙酸的含量。
文蛤在15℃無氧的環境下144小時，其斧足中AlaT和PEPCK的活性沒有顯著的變化，然而PK、FR和合成丙酸酵素的活性則分別先增加至控制組的145﹪、124﹪和300﹪，而後分別下降為控制組的119﹪、117﹪和120﹪。文蛤在30℃無氧的環境下48小時，其斧足中AlaT的活性隨著無氧時間的延長而下降，然而PEPCK和FR的活性則增加，而PK和合成丙酸酵素的活性則分別先增加至控制組的152﹪和300﹪，而後下降為控制組的134﹪和200﹪。
由上述結果得知，測定文蛤斧足和血淋巴中琥珀酸的增加量可作為文蛤活力的指標，除此之外，在夏季時，PEPCK和FR的活性亦可作為文蛤活力的指標。

關鍵詞：文蛤、丙胺酸、琥珀酸、丙酸、無氧代謝酵素、血淋巴酸-鹼平衡

Abstract

Hard clam are closing their shells when changes of season, temperature, salinity and environmental. Hard clam are bottom organisms. This study approaches the effect of anaerobic treatment on the acid-base balance in haemolymph of hard clams, the accumulation of anaerobic metabolites in foot and haemolyph of hard clams as well as in incubated water, and activities of pyruvate kinase (PK), analnine aminotransferase (AlaT), phosphoenopyruvate carboxykinase (PEPCK), fumarate reductase (FR) and propionate-formed enzymes in foot of hard clams.
The pO2 of haemolymph in hard clams exposed to air for 1 minute at 23 and 30℃ is decreased by a half. Furthermore, the remaining of pO2 of haemolymph in hard clams after air exposure for 2 min at 15℃ is 25% of that of control group. Haemolymph pO2 decreases 60% for hard clams incubated in anaerobic condition for 2 hr at 28℃. By extending anaerobic incubation time to 71 hr, haemolymph pH decreases, while pCO2 increases. A 23% decrease in pO2 of haemolymph of hard clams incubated in artificial seawater with 597 ppm K+ for 3 hr is found. Haemolymph pO2 decreases 39% and 42% for hard clams incubated in artificial seawater with 100 ppm of KCN and that lacking Mg2+ for 10 hr, respectively.
In the anaerobic conditions, alanine, succinate as well as succinate and propionate are main anaerobic metabolites in foot, haemolymph, and incubated water, respectively. Alanine in foot, and succinate and propionate in heamolymph and incubation water are accumulated with anaerobic incubation time increased. However, the amount of succinate in incubated water decreases in the late period of anaerobic treatment. Hard clams only survive for 48 hr at 30℃ under anoxia. However they can survive for more than 120 hr by the addition of 5 ppm chloramphenicol at 30℃ under anoxia. The effect of chloramphenicol on the accumulation of anaerobic metabolites is not significant. Alanine and succinate in foot, and alanine, succinate and propionate in haemolymph are accumulated with anoxia time increased, when hard clams are incubated in artificial seawater lacking Mg2+. However, propionate in foot is not detectable.
Changes in the activities of AlaT and PEPCK in foot of hard clams incubated in anaerobic conditions for 144 hr are not significant. However, activities of PK, FR and propionate-formed enzymes in foot first increase to 145%, 124% and 300% of the control group and then thereafter, decrease to 119%, 117% and 120%, respectively. Activities of AlaT, and PEPCK and FR in foot of hard clams incubated in anaerobic condition for 48 hr at 30℃ decrease and increase with anaerobic incubation time increased, respectively. However, the activities of PK and propionate-formed enzymes first increase to 152% and 300% of the control group and then decrease to 134%and 200%, respectively.
The result of the present study was detectable increase amount of succinate in foot and haemolymph of hard clam is indicator of activity. In summer, the activities of PEPCK and FR is indicator of hard clam activity.

Keywords: hard clam, alanine, succinate, propionate, anaerobic metabolism enzymes, haemolymph acid-base balance

目錄
中文摘要 i
英文摘要 iii
第一章 前言
一、本研究的緣由和動機 1
二、二枚貝的簡介 2
三、二枚貝的生長環境 4
四、二枚貝的呼吸作用 7
五、二枚貝的能量代謝 8
第二章 材料與方法
一、 材料 14
（一）實驗動物 15
（二）實驗用耗材、酵素及化學藥品 15
二、方法
（一） 暴露於空氣中、無氧環境或化學物對文蛤血淋
巴酸-鹼平衡和溶氧量的影響 16
（二） 溫度對文蛤在無氧環境下半致死時間的影響 17
（三） 溫度對文蛤無氧代謝物（丙胺酸、琥珀酸和丙
酸）和丙酮酸的影響 18
（四） 人工海水中缺乏鎂離子對文蛤無氧代謝物（丙
胺酸、琥珀酸和丙酸）和丙酮酸的影響 21
（五）溫度對文蛤參與無氧代謝酵素活性的影響 22
第三章 結果
一、暴露於空氣中、無氧環境或化學物對文蛤血淋巴酸-
鹼平衡和溶氧量的影響 29
二、溫度對文蛤在無氧環境下半致死時間的影響 31
三、溫度對文蛤無氧代謝物（丙胺酸、琥珀酸和丙酸）和
丙酮酸的影響 32
四、人工海水中缺乏鎂離子對文蛤無氧代謝物（丙胺酸、
琥珀酸和丙酸）和丙酮酸的影響 36
五、溫度對文蛤參與無氧代謝酵素活性的影響 37
第四章 討論
一、暴露於空氣中、無氧環境或化學物對文蛤血淋巴酸-
鹼平衡和溶氧量的影響 39
二、溫度對文蛤在無氧環境下半致死時間的影響 42
三、溫度對文蛤無氧代謝物（丙胺酸、琥珀酸和丙酸）
和丙酮酸之影響 43
四、人工海水中缺乏鎂離子對文蛤無氧代謝物（丙胺酸、
琥珀酸和丙酸）和丙酮酸的影響 50
五、溫度對文蛤參與無氧代謝酵素活性的影響 50
第五章 結論 55
參考文獻 56
表 76
圖 84

參考文獻

山東省水產學校主編，1979。貝類養殖學，衣北出版社，pp55。

巫文隆，1980。台灣重要食用雙枚貝類研究，貝類學報，7：101-114。

何雲達，1988。文蛤底質調查與試驗，Bulletin of Taiwan fisheriers research institute.45, 45-56。

何曉琛，2000。文蛤養殖池水層及底泥層中物質濃度之垂直分怖與季變異，國立台灣大學農業工程學研究所碩士論文。

李龍雄，1982。水產養殖學（中冊），前程出版社，pp221。

李安進、李明貞和賴弘智，2002。文蛤出入水管開關因子之研究，中華生質能源學會會誌，21（1）, 55-62。

林朝榮、吳錫圭、張寬敏、藍子樵、石忠榮、賴景陽、羅進宗和巫隆文，1978。台灣貝類中文名稱定名，貝類學報，5，83。

胡興華，1974。臺灣西南淺海養殖貝類斃死調查研究，臺灣省水產試驗所試驗報告，23, 1-20。

秦宗顯，1993。文蛤Meretrix lusoria Röding受汞影響之急慢性毒性研究，國立台灣大學動物學研究所博士論文。

郭河，1964。臺灣經濟貝類調查，中國農村復興聯合委員會 特刋第三十八號，32-50。

陳建初，1981。水質分析，九大圖書公司，133-143。

陳慧琦，1996。溫度、鹽度及藻類對牡蠣、文蛤濾食影響之研究，國立中山大學海洋資源研究所碩士論文。

許紋華，2002。食物與環境因子對文蛤攝食之影響，國立台灣海洋大
學水產養殖所碩士論文。

曾文陽，1976。臺灣西南沿海養殖貝類大量死亡原因研究，臺灣省水產試驗所試驗報告，26, 1-36。

蔡英亞、張英和魏若飛，1979。貝類學概論pp11。

鄭森雄，1975。臺灣西南部水質污染與養殖貝類大量死亡，臺灣水產學會刊，4, 51-71。

Akberali H.B. & Trueman E.R. (1979) PO2 and pCO2 changes in the mantle cavity of Scrobicularia plana (bivalvia) under normal and stress conditions. Estuarine and Coastal Marine Science 9, 499-507.

Ali F. & Nakamura K. (2000) Metabolic characteristics of the Japanese clam Ruditapes philippinarum（Adams and Reeve）during aerial exposure. Aquaculture Research 31, 157-165.

Alt R.M. (1970) The influence of suspension density and temperature on the filtration rate of Hiatella arctics. Marine Biology 6, 291-302.

Anderson A.E., Felbeck H. & Childress J.J. (1990) Aerobic metabolism is maintained in animal tissue during rapis sulfide oxidation in the symbiont-containing clam Solemya reidi. The Journal of Experimental Zoology 256, 130-134.

Andrews T.R. & Reids R.G.B. (1972) Ornithine cycle and tricolytic enzymes in four bivalve molluscs. Comparative Biochemistry and Physiology 42B, 475-491.

APHA A. & WEF (1995) Standard Methods for the
Examination of Water and Wastewater, 19th ed. American
Public Health Association, Washington, DE, 4-82.

Babarro J.M.F. & de Zwaan A. (2002) Influence of abiotic factors on bacterial proliferation and anoxic survival of the sea mussel Mytilus edulis L. Journal of Experimental Marine Biology and Ecology 273, 33-49.

Bayne B.L., Bayne C.T., Carefoot T.C. & Thomson R.J. (1976) The physiological ecology of Mytilus californianus (Conrad). 2. Adaptation to low oxygen tension and air exposure. Oecologia 22, 229-250.

Bjorn B. (1972) Effects of adaptation to reduced salinity on filtaration activity and growth of mussels (Mytilus edulis L.). Journal of Experimental Marine Biology and Ecology 10, 41-47.

Brooks S.P.J., de Thillart A., van den Cittani G., Cortesi O.P. & Storey K.B. (1991) Differential survival of Venus gallina and Scapharca inaequivalvis during anoxic stress: covalent modification of phosphofructokinase and glycogen phosphorylase during anoxia. Journal of Comparative Physiology 161, 207-212.

Byrne R.A. & Mcmahon B.R. (1991) Acid-base and ionic regulation during and following emersion in the freshwater bivalve Anodanta grandis simpsoniana (Bivalvia:Unionidae). Biometric Bulletin 181, 289-297.

Byrne R.A., Shipman B.N., Smatresk N.J., Dietz T.H. & McMahon R.F. (1991) Acid-base balance during emergence in the freshwater bivalve Corbicula fluminae. Physiological Zoology 64, 748-766.

Carpene E.C. & Zurburg W. (1981) Anaerobic energy metabolism in the bivalves Scapharca inaequivalvis (Bruguiere) and Venus gallina L. In Abstracts of the Third ESCPB Conyress 118-119.

Cerrato R.M. (2000) What fish biologists should know about bivalve shells. Fisheries Research 46, 39-49.

Chen J.C. & Chen J.S. (1998) Acid-base balance, ammonia and lactate levels in the haemolymph of Penaeus japonicus during aerial exposure. Comparative Biochemistry and Physiology 121A, 257-262.

Chen Lo-chai (1990) Aquaculture in Taiwan. 223-227.

Cheng W., Liu C.H., Cheng S.Y. & Chen J.C. (2004) Effects of dissolved oxygen on the acid-base balance and ion concentration of Taiwan abalone Haliotis diversicolor supertexta. Aquaculture 231, 573-586.

Cheng W., Liu C.H. & Kuo C.M. (2003) Effects of dissolved oxygen on hemolymph parameters of freshwater giant prawn, Macrobrachium rosenbergii (de Man). Aquaculture 220, 843-856.

Chiancone E., Boffi A., Verzili B., Ascoli F. (1985) Proprieta delle emoglobine del mollusco Scapharca inaequivalvis in rap porto con le condizioni ecologiche del mare Adriatico. Attl simp. biochim.mar.,editoriale grasso.bologna,13-20.

Chiou T.K., Lin J.F. & Shiau C.Y. (1998) Changes in extractive components and glycogen in the edible meat of hard clam Meretrix lusoria during storage at different temperatures. Department of Marine Food Science 64(1), 115-120.

Churchill T.A. & Storey K.B. (1995) Metabolic responses to freezing and anoxia by the periwinkle Littorina littorea. Journal of Thermal Biology 21(1), 57-63.

Daum G., Glatz H. & Paltauf F. (1977) Lipid metabolism in an inositol-deficient yeast, Saccharomyces carlsbergensis: The influence of temperature and anaerobiosis on the cellular lipid composition. Biochimica et Biophysica Acta(BBA)-Lipids and Lipid Metabolism 488(3), 484-492.

Deaton L.E., Derby J.G.S., Subhedar N. & Greenberg M. (1989) Osmorgulation and salinity rolerance in two species of bivalve mollusk Limnoperna fortunei and Mytilopsis leucophaeta. Journal of Experimental Marine Biology and Ecology 133, 67-79.

de Vooys C.G.N. (1980) Anaerobic metabolism in sublittoral living Mytilus galloprovincialis Lam. In Mediterranean-II. Partial adaptation of pyruvate kinase and phosphoenolpyruvate carboxykinase. Comparative Biochemistry and Physiology 65B, 513-518.

de Vooys C.G.N. (1987) Adaptation to anaerobic metabolism in two mussel species Mytilus edulis and Mytilus galloprovincialis from the tidal zone at Arcachon Bay France. Netherlands Journal of Sea Research 21(1), 17-23.

de Zwaan A. (1972) Pyruvate kinase in muscle extracts of the sea mussel Mytilus edulis L. Comparative Biochemistry and Physiology 42B, 7-14.

de Zwaan A. (1977) Anaerobic energy metabolism in bivalve molluscs. Oceanography and Marine Biology 15, 103-187.

de Zwaan A. (1983) Carbohydrate catabolism in bivalves.In the mollusca,Vol1(Edited by Hochachks P. V.),pp. 137-175. Academic Press, New York

de Zwaan A., Babarro J.M.F., Monari M. & Cattani O. (2002) Anoxia survival potential of bivalves:(arte) facts. Comparative Biochemistry and Physiology 131 A, 615-624.

de Zwaan A., Cortesi P., van den Thillart G., Roos J. & Storey K.B. (1991) Differential sensitivities to hypoxia by two anoxia-tolerant marine molluscs: a biochemical analysis. Marine Biology 111, 343-351.

de Zwaan A. & de Bont A.M.T. (1975) Phosphoenolpyruvate carboxykinase from adductor muscle tissue of the sea mussel Mytilus edulis L. Journal of Comparative Physiology 96, 85-94.

de Zwaan A. & Holwerda D. A. (1972) The effect of phosphoenolpyruvate, fructose 1,6 diphosphate and pH on allosteric pyruvate kinase in muscle tissue of the bivalve Mytilus edulis L. Biochimica et Biophysica Acta 276, 430-433.

de Zwaan A., Isani G., Cattani O. & Cortrsi P. (1995) Long-term anaerobic metabolism of erythrocytes of the arcid clam Scapharca inaequivalvis. Journal of Experimental Marine Biology and Ecology 187, 27-37.

de Zwaan A. & Putzer V. (1985) Metabolic adaptations of intertidal invertebrates to environmental hypoxia (a comparison of environmental anoxia to exercise anoxia). In Physiological Adaptations of Marine Animals 39, 33-62.

de Zwaan A., Schaub B.E.M. & Babarro J.M.F. (2001) Anoxic survival of Macoma balthica: the effect of antibiotics, molybdate and sulphide. Journal of Experimental Marine Biology and Ecology 256, 241-251.

de Zwaan A. & van Marrewijk W.J.A. (1973) Anaerobic glucose degradation in the sea mussel Mytilus edulis L. Comparative Biochemistry and Physiology 44B, 429-439.

de Zwaan A. & Zwandee D.I. (1971) Body distribution and seasonal changes in the glycogen content of the common sea mussel Mytilus edulis L. Comparative Biochemistry and Physiology 43A, 53-58.

Downie J.A. & Garland P.B. (1973) An antimycin A- and
cyanide-resistant variant of Candida utilis arising during copper-
limited growth. Biochemistry 134, 1051-1061.

Ebberink R.H.M & de Zwaan A. (1980) Control of glycolysis in the posterior adductor muscle of the sea mussel Myrilus edulis L. Journal of Comparative Physiology 137B, 165-171.

Eberlee J.C., Storey J.M. & Storey K.B. (1983) Anaerobiosis recovery from anoxia and the role of strombine and alanopine in the oyster Crassostrea rirginica. Canadian Journal of Zoology 61, 2682-2687.

Ellington W.R. (1983) The recovery from anaerobic metabolism in invertebrates. Journal of Experimental Zoology 228, 431-444.

Fisher S.W., Stromberg P., Bruner K.A. & Boulet L.D. (1991) Moulluscicidal activity of potassium to the zebra mussel Dreissena polymorphia: toxicity and mode of action. Aquatic Toxicology 20, 219-234.

Föll R.L., Pleyers A., Lewandovski G.J., Wermter C., Hegemann V. & Paul R.J. (1999) Anaerobiosis in the nematode Caenorhabditis elegans. Comparative Biochemistry and Physiology 124B, 269-280.

Gäde G. (1983) Energy metabolism of arthropods and mollusks during environmental and functional anaerobiosis. The Journal of Experimental Zoology 228, 415-429.

Gäde G. (1988) Energy metabolism during anoxia and recovery in shell adductor and foot muscle of the gastrogod mollusk Haliotis lamellose: Formation of the novel anaerobic and product tauropine. Biometric Bulletin 175, 122-131.

Greenway C.S. & Storey B.K. (1999) The effect of prolonged on enzyme activities in oysters (Crassostrea virginica) at different seasons. Journal of Experimental Marine Biology and Ecology 242, 259-272.

Greenway C.S. & Storey B.K. (2000) Seasonal change and prolonged anoxia affect the kinetic properties of phosphofructokinase and pyruvate in oysters. Journal of Comparative Physiology 170, 285-293.

Heagaret H, Wikfors G.H., Soudant P., Depaporte M., Alix J.H., Smith B.C., Dixon M.S., Quere C., Lecoz J.R., Paillard C., Moal J. & Samain J.F. (2004) Immunological competence of western oysters Crassostrea virginica, fed different microalgal diets and challenged with a temperature elevation. Aquaculture 234, 541-560.

Hochachka P.W. & Musrafa T. (1972) Invertebrate facultative anaerobiosis. Science 178, 1056-1060.

Holwerda D.A. & de Zwaan A. (1980) On the role of fumarate reductase in anaerobic carbohydrate catabolism of Mytilus edulis L. Comparative Biochemistry and Physiology 67B, 447-453.

Ho M.S. & Zubkoff L.P. (1982) Anaerobic metabolism of the ribbed mussel Geukensia demissa. Comparative Biochemistry and Physiology 73B(4), 931-936.

Incze L.S., Lutz R.A. & Watling L. (1980) Relationships between effects of environmental temperature and seston on growth and mortality of Mytilus edulis in a temperate northern estuary. Marine Biology 57, 147-156.

Isani G., Cattani O., Carpene E., Tacconi S. & Cortesi P. (1989) Energy metabolism during anaerobiosis and recovery in the posterior adductor muscle of the bivalve Scapharca inaequivalvie (bruguiere). Comparative Biochemistry and Physiology 93B(1), 193-200.

Isani G., Cattani O., Zurzolo M., Pagnucco C. & Cortesi P. (1995) Energy metabolism of the mussel, Mytilus galloprovincialis, during long-term anoxia. Comparative Biochemistry and Physiology 110, 103-113.

Jeng S.S. & Tyan Y.M. (1982) Growth of the hard clam, Meretrix lusoria in Taiwan. Aquaculture 27, 19-28.

Jorgensen C.B. (1981) Feeding and clearning mechanisms in the suspension feeding bivalve Mytilus edulis. Marine Biology 65, 159-163.

Kita K., Hirawake H., Miyadera H., Amino H. & Takeo S. (2002) Role of complex II in anaerobic respiration of the parasite mitochondria from Ascaris suum and Plasmodium falciparum. Biochimica et Biophysica Acta 1553, 123-139.

Kluytmans J.H., de Bont A.M.T., Janus J. & Wusman T.C.M. (1977) Time dependent changes and tissue specificities in the accumulation of anaerobic fermentation products in the sea mussel Mytilus edulis L. Comparative Biochemistry and Physiology 58B, 81-87.

Kluytmans J.H., de Bont A.M.T., Kruttwagen E.C.J., Ravestein H.J.L. & Veenhof P.R. (1983) Anaerobic capacities and anaerbic energy production of some mediterranean bivalves. Comparative Biochemistry and Physiology 75B, 171-179.

Kluytmans J.H., van Graft M., Janus J. & Pieters H. (1978) Production and excretion of volatile fatty acids in the sea mussel Mytilus edulis L. Journal of Comparative Physiology 123, 163-167.

Kluytmans J.H., Veenhof P.R. & de Zwaan A. (1975) Anaerobic production of volatile fatty acids in the sea mussel Mytilus edulis L. Journal of Comparative Physiology 104, 71-78.

Konosu S. & Shibota M.Y. (1967) Hashimoto:Concentration of organic acids in shellfifh with particular reference to succinate acid. Eiyo To Shokuryo 20, 186-189.

Laudien J., Schiedek D., Brey T., Portner H.O. & Arntz W.E. (2002) Survivorship of juvenile surf clams Dpmax serra (Bivalvia, Donacidae) exposed to severe hypoxia and hydrogen sulphide. Journal of Experimental Marine Biology and Ecology 271, 9-23.

Lazou A., Michaelidis B. & Beis I. (1989) Evidence for glycolytic enzyme binding during anaerobiosis of the foot muscle of Patella caerulea (L.) . Journal of Comparative Physiology 158B, 771-777.

Lee A.C., Lee M.C., Chen S.M. & Chin T.S. (2005) A study on the effect of environmental parameters on the oxygen consumption of the bivalve, Meretrix lusoria. Aquaculture Research (submitted)

Lee A.C., Lin C.R. & Chen S.Y. (2004) The effect of temperature and components of artificial seawater on digging ability of hard clam (Meretrix lusoria). Journal of The Fisheries Society of Taiwan 31(4), 251-262.

Levitt M.J & Arp J.A. (1991) The effects of sulfide on the anaerobic metabolism of two congrneric species of mudflat clams. Comparative Biochemistry and Physiology 98B, 339-347.

Livingstone D.R. (1983) Invertebrate and vertebrate pathways of anaerobic metabolism: evolutionary considerations. Journal of the Geological Society 140, 27-37.

Luquet C.M. & Ansaldo M. (1997) Acid-base balance and ionic regulation during emersion in the estuarine intertidal crab Chasmagnathus granulata dana (decapoda grapsidae). Comparative Biochemistry and Physiology 117A(3), 407-410.

Lushchak V.I., Bahnjukova T.V. & Spichenkov A.V. (1997) Modification of pyruvate kinase and lactate dehydrogenase in foot muscle of the sea mussel Mytilus galloprovincialis under anaerobiosis and recovery. Brazilian Journal of Medical and Biological Research 30(3), 381-385.

Macy J. M., Ljungdahl L.G. & Gottschalk G. (1978) Pathway of succinate and propionate formation in Bacteroides fragilis. Journal of Bacteriology 134, 81-84.

Meinardus G. & Gäde G. (1981). Anaerobic metabolism of the common cockle Cardium edule-IV. Time dependent changes of metaboites on the foot and gill tissue induced by anoxia and electrical stimulation. Comparative Biochemistry and Physiology 70B, 271-277.

Moss W.D. (1984a) Methods of Enzymatic Analysis. Volume VII. Metabolites2 : Tri- and Dicarboxylic Acid, Purines, Pyrimidines and Derivatives, Coenzymes, Inorganic Compounds. 25-34.

Moss W.D. (1984b) Methods of Enzymatic Analysis. Volume VI. Metabolites 1 : Carbohydrates. 570-577.

Moss W.D. (1984c) Methods of Enzymatic Analysis. Volume VIII. Metabolites3 : Lipids, Amino Acids and Related Compounds. 345-349.

Moss W.D. (1984d) Methods of Enzymatic Analysis. Volume III. Enzymes 1 : Oxidoreductases, Transferases. 444-456.

Moon T.W. & Pritchard A.W. (1970) Metabolic adaptations in vertically-separated populations of Mytilus californianus Conrad. Journal of Experimental Marine Biology and Ecology 5, 35-46.

Newsholme E.A. (1978) Substrate cycles:their metabolic, energetic and thermic consequences in man. Biochemical Society Symposia 43, 183-205.

O’Donnell J.M., Durand M.E., Robitaille P.M.L., Fifher S.W. & Stromberg P.C. (1996) P31-NMR analysis of lethal and sublethal lesions produced by KCl-intoxication in the zebra mussel Dreissena polymorpha. Journal of Experimental Zoology 276, 53-62.

Oeschger R. (1990) Long-term anaerobiosis in sublittoral marine invertebrates from the Western Baltic Sea: Halicryptus spimulosus(Priapulida) Astarte borealis and Arctica islandica (Bivalvia). Marine Ecology Progress Series 59, 133-143.

Paparo A.A. & Dean R.C. (1984) Activity of the lateral cilia of the oyster Creassostrea virginica: response to changes in salinity and to changes in potassium and magnesium concentration. Marine Behaviour and Physiology 11, 111-130.

Perez C.A., Beiras R. & Albentosa M. (1994). Effects of algal food concentration and body size on the ingestion rate of Ruditapes decussates (Bivalvia) veliger larvae. Marine Ecology Progress Series 115, 87-92.

Portner H.O., Grieshaber M.K. & Heisler N. (1984) Anaerobiosis and acid-base status in marine invertebrates: effect of environmental hypoxia on extracellular and intracellular pH in Sipunculus nudus L. Journal of Comparative Physiology 155B, 13-20.

Powell M.A. & Somero G.N. (1986) Hydrogen sulfide oxidation is coupled to oxidative phosphorylation in mitochondria of Solemys reidi. Marine Biology Research Division 233, 563-566.

Rice M.A. (1999) Uptake of dissolved free amino acids by northern quahogs, Mercenaria mercenaria and its relative importance to organic nitrogen deposition in Narragansett Bay, Rhode Island. Journal of Shellfish Research 18(2), 547-553.

Riisgard H.U. & Randor A. (1981) Energy budgets, growth and filtration rate in Mytilus edulis at different algal concentrations. Marine Biology 61, 227-234.

Saleuddin A.S.M. & Wilbur K. (1983) The mollusca. Volume2: Environmental biochemistry and physiology.

Schroff G. & Schottler U. (1977) Anaerobic reduction of fumarate in the body wall musculature of the lugworm Arenicola marine. Journal of Comparative Physiology 116, 325-336.

Schulz T.K.F., Kluytmans J.H. & Zandee D.I. (1982) In vitro production of propionate by mantle mitochondria of the sea mussel Mytilus edulis L: overall mechanism. Comparative Biochemistry and Physiology 73B(3), 673-680.

Shick J.M., Gnaiger E., Widdows J., Bayne B.L. & de Zwaan A. (1986) Activity and metabolism in the mussel Mytilus edulis L. during intertidal hypoxia and aerobic recovery. Physiological Zoology 59, 627-642.

Silrerman H., Steffens W.L. & Dietz T.H. (1983) Calaium concretions in the gills of a freshwater mussel serve as a calcium reservoir during period of hypoxia. Journal of Experimental Zoology 227, 177-189.

Spicer J.I., Taylor A.C. & Hill A.D. (1988) Acid-base status in the sea urchins Psammechinus miliaris and Echinus esculentus (Echinodermata:Echinoidea) during emersion. Marine Biology 99, 527-534.

Thachuck R.D., Saz H.J., Weinstein P.P., Finnegan K. & Mueller J.F. (1977) The presence and pissible function of methylmonyl CoA muease and propionyl CoA carboxylase in Spirometra mansonoides. Journal of Parasitology 63, 769-774.

Tyler D. D. ( 1992) In the mitochondrion in Health&Disease (Tyler,D.D.,ed) VCH Publishers Inc., New York, 269-351.

van den Thillart G. & de Vries I. (1985) Excretion of volatile fatty acids by anoxic Mytilus edulis and Anodonta cygnea. Comparative Biochemistry and Physiology 80B, 299-301.

Vial M.V., Simpfendorfer R.W., Lopez D.A., Gonzalez M.L. & Oelckers K. (1992) Metabolic responses of the intertidal mussel Perumytilus purpuratus (Lamarck) in emersion and immersion. Journal of Experimental Marine Biology and Ecology 159, 191-201.

Wadsworth W.G. & Riddle D.L. (1989) Developmental regulation of energy metabolism in Caenorhabditis elegans. Development Biology 132, 167-173.

Waller T.A. (1980) Scannind electron microscopy of shell and mantle in the order Arcoida(Mollusca:Bivalvia). Smithsonian Contribution to Zoolfy 313. Smithsonian Institution Press, Washington, D. C.58pp

Ward J.E. & Targett N.M. (1989) Influence of marine microalgal metabolites on the feeding behavior of the blue mussel Mytilus edulis. Marine Biology 101, 313-321.

Widdows J., Bayne B.L., Livingstone D.R., Newell R.I.E. & Donlcin P. (1979) Physiological and biochemical responses of bivalve mollusks to exposure to air. Comparative Biochemistry and Physiology 62A, 301-308.

Zammit V.A. & Newsholme E.A. (1978) Properties of pyruvate kinase and phosphoenolpyruvate carboxykinase in relation to the direction and regulation of phosphoenolpyruvate metabolism in muscles of the frog and marine invertebrates. Biochemical Journal 174, 979-987.

Zs.-Nagy I. (1977) Cytosome (yellow pigment granules) of mollusks as cell organelles of anoxic energy production. International Review of Cytology 49, 331-377.

Zurburg D.I., Kluytmans J.H., Zurburg W. & Pieters H. (1980) Seasonal variations in biochemical composition of Mytilus edulis with reference to energy metabolism and gametogenesis . Netherlands Journal of Sea Research 14, 1-29.

Zurburg W. & de Zwaan A. (1981) The role of amino acids in anaerobiosis and osmoregulation in bivalves. Journal of Experimental Zoology 215, 315-325.

Zurbuge W. & Kluytmans J.H. (1980) Organ specific changes in energy metabolism due to anaerobiosis in the sea mussel Mytilus edulis L. Comparative Biochemistry and Physiology 67B, 317-322.