太平洋磷虾蛋白酶的生物化学和组织化学研究
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摘要
磷虾，是磷虾科动物的通称，是一个甲壳类大家族，分布广泛于北太平洋。它以浮游植物为食物，位于食物链的初级阶段。磷虾极易自溶，这与虾体内存在高效的酶系统有密切的关系。该系统能够降解各种蛋白质、多糖，并且降解迅速。
目前人们对磷虾的研究工作以南极磷虾(Euphausia superba)居多。在南极磷虾体内具有非常高效的消化系统，这是一个由多种酶和抑制因子相结合的复合酶系统。当磷虾死后，抑制因子失去作用，消化酶迅速将磷虾的蛋白质、多糖等降解。这种高效的蛋白酶活性引起了人们的极大兴趣，现在已经对其中的部分蛋白酶分别进行了纯化和性质研究工作。例如，研究发现，与脊椎动物的胰蛋白酶相比，磷虾酶对纤维蛋白和酪蛋白的降解能力更强；已经确定了一种磷虾外切酶的序列，并且已在实验室合成成功。已从南极磷虾提取了8种蛋白酶：三种丝氨酸类胰蛋白酶，一种丝氨酸类胰凝乳蛋白酶，以及两种羧肽酶A，两种羧肽酶B。磷虾酶作为一种高效的复合酶，目前已被医用，例如对慢性溃疡的清创，清除牙斑等。
目前，对太平洋磷虾(Euphausia pacifica)的研究仅限于对其生态、分布的研究。我们观察到，太平洋磷虾死后同样发生迅速自溶现象，因此推测在其体内可能也存在一个高效的酶系统，而该系统在磷虾死后，抑制因子失去作用，从而迅速地将蛋白质、多糖等降解。由于海上过度捕捞，鱼群数量大幅减少，作为鱼类饵料的磷虾数量大幅增加。目前磷虾的应用主要限于饵料加工，因此研究其体内的蛋白酶，提高磷虾深加工的附加值有重要的意义。
在本文中，进行了下列工作：对太平洋磷虾蛋白酶粗酶的相关生化性质进行探讨；采用不同方法对其类胰蛋白酶进行提纯，并探讨了相关的生化性
太平洋磷虾蛋自酶的生物化学和组织化学研究
质;最后，确定了太平洋磷虾的消化系统结构组成及类胰蛋白酶定位情况。
用0.O5mol/L Tris一HCI(pH7.0)或蒸馏水对太平洋磷虾进行粗提，发现蒸
馏水粗提酶液中的蛋白酶活力普遍比Tris一HCI的高，且稳定性好。粗酶具有
很强的温度稳定性。在长时间(30d)常温20℃保存下，酶活损失小于巧%;
对粗酶液进行热激处理(15min)，温度范围从室温至100℃，结果发现，粗酶
蛋白酶活性减少幅度小，其蛋白酶比活力逐渐提高。这点对于我们今后大规
模的粗酶提取工作有非常大的意义，也就是说，我们可以通过简单的热激方
式，去掉大部分杂蛋白，从而获得粗酶。
通过0%一50%的硫酸钱沉淀比较发现，采用35%浓度的硫酸钱沉淀时(采
用pH 6.8磷酸盐缓冲液溶解沉淀)，沉淀中酶活力损失最少。
粗酶的适宜pH(20℃)较广。在pH 5.0时，粗酶有高酶活力，在pHS一10
也有较高酶活力，推测在粗酶中不仅有酸性蛋白酶存在，而且有多种碱性蛋
白酶存在。
采用DEAE Sepharose Fast Flow离子柱、sephaeryl 5200凝胶过滤柱层析、
p一aminobenzamidine亲和层析和鸡卵类粘蛋白亲和层析等分离方法，获得类胰
蛋白酶。采用12%的分离胶，对不同纯化方法获得的样品电泳后，推算分子
量大约在35kDa。
大平洋磷虾类胰蛋白酶的最适pH有两个，分别为5.0和7.0。最适反应
温度为20oC(pHS.o)。
ca2+对类胰蛋白酶几乎没有激活效果，而Mg2+具有一定的激活作用。鸡
卵类粘蛋白和Benzamidine具有明显的抑制作用，PMSF和TLCK有抑制作用，
而TPCK没有抑制效果，这说明分离出的蛋白酶属于丝氨酸类蛋白酶。
为研究太平洋磷虾类胰蛋白酶的来源，对其消化系统进行了组织学及组
织化学的研究工作。太平洋磷虾的消化系统主要分为消化管和消化腺两部分。
消化管又可分为食道、胃(责门胃和幽门胃)、中肠、后肠。消化腺主要是中
肠腺。显微观察，消化管的组织结构与对虾的基本相同，即由内向外分为四
中国海洋大学硕士研究生毕业论文
层:粘膜层、粘膜下层、肌肉层和外膜。中肠腺发达，为分叶结构，每叶由
许多小的管状腺组成，有薄的结缔组织和肌肉纤维将管状小室隔开。实验发
现，一个分叶的消化酶分泌明显，而另一个则分泌较少，推测每个分叶的功
能可能有所不同。
实验结果表明，在太平洋磷虾的消化道上皮粘膜层广泛存在类胰蛋白酶，
且胃肠道内的类胰蛋白酶含量明显比消化腺高很多。胃壁和中肠腺细胞均分
泌酸性粘多糖，且贡门胃分泌量高于其它消化器官。
Krill belongs to Crustacea, which is widely distributing. The diet of krill contains large numbers of diatoms and copepods, and it locates at the primary phase of food cycle. Krill decomposes very soon by the efficient enzyme system, which can rapidly degrade protein and amylose.
People have been paying much attention in Antarctic krill (Euphausia superbd). Antarctic krill is known to have a very efficient digestive system, which is based on a combination of enzymes controlled by inhibitors. The efficiency of these enzymes is clearly visualized when the krill post mortem is autolyzed, due to the disabling of the inhibitor system. The high proteolytic activity has raised a lot of interest and a number of studies on the characteristics of these enzymes have been performed. For example, krill enzymes have been shown to have a higher activity toward fibrin and casein compared to trypsin. Also, a serine proteinase from krill referred to as euphauserase has been sequenced, modified, and produced recombinantly. In the present work, proteinases refined from a krill extract used for an enzymatic debrider have been studied. The extract contains eight proteolytic enzymes: three trypsin-like serine proteinases, one chymotrypsin-like serine proteinase, two carboxypeptidase A enzymes, and two carb
oxypeptidase B enzymes. Krill enzymes are now being exploited for medical applications, such as debridement of necrotic wounds and removal of dental plaque.
Euphausia pacifica is the dominant euphausiid species in the North Pacific. The present work is limited in ecology and distribution. We also find the krill post mortem can autolyzed. It can be explained there exists an inhibitor system in krill when it died, for the sake of the system disable, the proteinase degrade protein and amylose. Because of the amount of fishes reduced rapidly, and the krill, which is the diet of fish, rose accordingly. The krill is only used for aquaculture feed now, so studying the proteolytic enzymes will increase the additional value.
In this thesis, the characteristics of Euphausia pacifica crude extract were investigated by different methods, which were used to purify and characterize the trypsin-like enzyme. By histochemistry method, we studied the digestive system and confirmed the site of trypsin-like enzyme in krill.
Compared the two extracting buffer solution of 0.05mol/L Tris-HCl ( pH7.0) and distill water, we found the activity of proteolytic enzyme extracted from distill water is higher than that of Tris-HCl and-with good stability. Crude enzyme has high temperature stability. Keeping at constant temperature(20 ) for a long
time(30d), the activity remains more than 85%. After heat stimulation range from room temperature to 100 for 15min, the activity of crude enzyme reduced little while specific activity raised rapidly, that is to say, we can use simple method such as heat stimulation to get the proteolytic enzymes.
Compared the depositions from 0%~50% (NH4)2SO4, The highest enzyme recovery rate obtained when the concentration of (NH4)2SO4 was 35%.
The effect of pH on crude enzyme was examined. The major protein activity is at pH5.0 and a minor activity is at pH8~10. So it was implied that in crude enzyme there are both acid proteases and more than one kinds of alkaline proteases.
It was found that trypsin-like enzyme isolated as monomeric protein had a molecular weight of 35 000Da.
The enzyme activity was extremely sensitive to ovomucoid and Benzamidine, very sensitive to PMSF, TLCK, however not sensitive to TPCK, indicated that it belongs to the group of serine-type protease.
In order to confirm the site of trypsin-like enzyme in Euphausia pacifica, we examined the digestive system by using histochemistry method. The digestive system of Euphausia pacifica mainly composed of digestion duct and gland. The digestion duct can be divided by gullet, stomach (cardia stomach and pylorus stomach), mid-gut and hindgut. Mid-gut gland is the main digestive gland. Observed by Olympus microscope, the structure of krill digestive
目前人们对磷虾的研究工作以南极磷虾(Euphausia superba)居多。在南极磷虾体内具有非常高效的消化系统，这是一个由多种酶和抑制因子相结合的复合酶系统。当磷虾死后，抑制因子失去作用，消化酶迅速将磷虾的蛋白质、多糖等降解。这种高效的蛋白酶活性引起了人们的极大兴趣，现在已经对其中的部分蛋白酶分别进行了纯化和性质研究工作。例如，研究发现，与脊椎动物的胰蛋白酶相比，磷虾酶对纤维蛋白和酪蛋白的降解能力更强；已经确定了一种磷虾外切酶的序列，并且已在实验室合成成功。已从南极磷虾提取了8种蛋白酶：三种丝氨酸类胰蛋白酶，一种丝氨酸类胰凝乳蛋白酶，以及两种羧肽酶A，两种羧肽酶B。磷虾酶作为一种高效的复合酶，目前已被医用，例如对慢性溃疡的清创，清除牙斑等。
目前，对太平洋磷虾(Euphausia pacifica)的研究仅限于对其生态、分布的研究。我们观察到，太平洋磷虾死后同样发生迅速自溶现象，因此推测在其体内可能也存在一个高效的酶系统，而该系统在磷虾死后，抑制因子失去作用，从而迅速地将蛋白质、多糖等降解。由于海上过度捕捞，鱼群数量大幅减少，作为鱼类饵料的磷虾数量大幅增加。目前磷虾的应用主要限于饵料加工，因此研究其体内的蛋白酶，提高磷虾深加工的附加值有重要的意义。
在本文中，进行了下列工作：对太平洋磷虾蛋白酶粗酶的相关生化性质进行探讨；采用不同方法对其类胰蛋白酶进行提纯，并探讨了相关的生化性
太平洋磷虾蛋自酶的生物化学和组织化学研究
质;最后，确定了太平洋磷虾的消化系统结构组成及类胰蛋白酶定位情况。
用0.O5mol/L Tris一HCI(pH7.0)或蒸馏水对太平洋磷虾进行粗提，发现蒸
馏水粗提酶液中的蛋白酶活力普遍比Tris一HCI的高，且稳定性好。粗酶具有
很强的温度稳定性。在长时间(30d)常温20℃保存下，酶活损失小于巧%;
对粗酶液进行热激处理(15min)，温度范围从室温至100℃，结果发现，粗酶
蛋白酶活性减少幅度小，其蛋白酶比活力逐渐提高。这点对于我们今后大规
模的粗酶提取工作有非常大的意义，也就是说，我们可以通过简单的热激方
式，去掉大部分杂蛋白，从而获得粗酶。
通过0%一50%的硫酸钱沉淀比较发现，采用35%浓度的硫酸钱沉淀时(采
用pH 6.8磷酸盐缓冲液溶解沉淀)，沉淀中酶活力损失最少。
粗酶的适宜pH(20℃)较广。在pH 5.0时，粗酶有高酶活力，在pHS一10
也有较高酶活力，推测在粗酶中不仅有酸性蛋白酶存在，而且有多种碱性蛋
白酶存在。
采用DEAE Sepharose Fast Flow离子柱、sephaeryl 5200凝胶过滤柱层析、
p一aminobenzamidine亲和层析和鸡卵类粘蛋白亲和层析等分离方法，获得类胰
蛋白酶。采用12%的分离胶，对不同纯化方法获得的样品电泳后，推算分子
量大约在35kDa。
大平洋磷虾类胰蛋白酶的最适pH有两个，分别为5.0和7.0。最适反应
温度为20oC(pHS.o)。
ca2+对类胰蛋白酶几乎没有激活效果，而Mg2+具有一定的激活作用。鸡
卵类粘蛋白和Benzamidine具有明显的抑制作用，PMSF和TLCK有抑制作用，
而TPCK没有抑制效果，这说明分离出的蛋白酶属于丝氨酸类蛋白酶。
为研究太平洋磷虾类胰蛋白酶的来源，对其消化系统进行了组织学及组
织化学的研究工作。太平洋磷虾的消化系统主要分为消化管和消化腺两部分。
消化管又可分为食道、胃(责门胃和幽门胃)、中肠、后肠。消化腺主要是中
肠腺。显微观察，消化管的组织结构与对虾的基本相同，即由内向外分为四
中国海洋大学硕士研究生毕业论文
层:粘膜层、粘膜下层、肌肉层和外膜。中肠腺发达，为分叶结构，每叶由
许多小的管状腺组成，有薄的结缔组织和肌肉纤维将管状小室隔开。实验发
现，一个分叶的消化酶分泌明显，而另一个则分泌较少，推测每个分叶的功
能可能有所不同。
实验结果表明，在太平洋磷虾的消化道上皮粘膜层广泛存在类胰蛋白酶，
且胃肠道内的类胰蛋白酶含量明显比消化腺高很多。胃壁和中肠腺细胞均分
泌酸性粘多糖，且贡门胃分泌量高于其它消化器官。
Krill belongs to Crustacea, which is widely distributing. The diet of krill contains large numbers of diatoms and copepods, and it locates at the primary phase of food cycle. Krill decomposes very soon by the efficient enzyme system, which can rapidly degrade protein and amylose.
People have been paying much attention in Antarctic krill (Euphausia superbd). Antarctic krill is known to have a very efficient digestive system, which is based on a combination of enzymes controlled by inhibitors. The efficiency of these enzymes is clearly visualized when the krill post mortem is autolyzed, due to the disabling of the inhibitor system. The high proteolytic activity has raised a lot of interest and a number of studies on the characteristics of these enzymes have been performed. For example, krill enzymes have been shown to have a higher activity toward fibrin and casein compared to trypsin. Also, a serine proteinase from krill referred to as euphauserase has been sequenced, modified, and produced recombinantly. In the present work, proteinases refined from a krill extract used for an enzymatic debrider have been studied. The extract contains eight proteolytic enzymes: three trypsin-like serine proteinases, one chymotrypsin-like serine proteinase, two carboxypeptidase A enzymes, and two carb
oxypeptidase B enzymes. Krill enzymes are now being exploited for medical applications, such as debridement of necrotic wounds and removal of dental plaque.
Euphausia pacifica is the dominant euphausiid species in the North Pacific. The present work is limited in ecology and distribution. We also find the krill post mortem can autolyzed. It can be explained there exists an inhibitor system in krill when it died, for the sake of the system disable, the proteinase degrade protein and amylose. Because of the amount of fishes reduced rapidly, and the krill, which is the diet of fish, rose accordingly. The krill is only used for aquaculture feed now, so studying the proteolytic enzymes will increase the additional value.
In this thesis, the characteristics of Euphausia pacifica crude extract were investigated by different methods, which were used to purify and characterize the trypsin-like enzyme. By histochemistry method, we studied the digestive system and confirmed the site of trypsin-like enzyme in krill.
Compared the two extracting buffer solution of 0.05mol/L Tris-HCl ( pH7.0) and distill water, we found the activity of proteolytic enzyme extracted from distill water is higher than that of Tris-HCl and-with good stability. Crude enzyme has high temperature stability. Keeping at constant temperature(20 ) for a long
time(30d), the activity remains more than 85%. After heat stimulation range from room temperature to 100 for 15min, the activity of crude enzyme reduced little while specific activity raised rapidly, that is to say, we can use simple method such as heat stimulation to get the proteolytic enzymes.
Compared the depositions from 0%~50% (NH4)2SO4, The highest enzyme recovery rate obtained when the concentration of (NH4)2SO4 was 35%.
The effect of pH on crude enzyme was examined. The major protein activity is at pH5.0 and a minor activity is at pH8~10. So it was implied that in crude enzyme there are both acid proteases and more than one kinds of alkaline proteases.
It was found that trypsin-like enzyme isolated as monomeric protein had a molecular weight of 35 000Da.
The enzyme activity was extremely sensitive to ovomucoid and Benzamidine, very sensitive to PMSF, TLCK, however not sensitive to TPCK, indicated that it belongs to the group of serine-type protease.
In order to confirm the site of trypsin-like enzyme in Euphausia pacifica, we examined the digestive system by using histochemistry method. The digestive system of Euphausia pacifica mainly composed of digestion duct and gland. The digestion duct can be divided by gullet, stomach (cardia stomach and pylorus stomach), mid-gut and hindgut. Mid-gut gland is the main digestive gland. Observed by Olympus microscope, the structure of krill digestive
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许实荣等：维生素B1和B62对中国对虾消化酶的影响。海洋科学，1987(4)：34～37
杨德渐、王永良编著：中国北部海洋无脊椎动物。北京：高等教育出版社，1996：324～325
于书坤，张树荣：虾类及甲壳动物消化酶研究的现状。海洋科学，1986，10(2)：60～63
于书坤：中国对虾消化酶的研究Ⅰ：消化酶的活力测定及性质的研究。海洋科学集刊，第28集，1987：85～90
张志峰，姜明：对中国对虾幼体中肠腺的结构和细胞化学研究。中国水产科学，2000，7(3)：11～15
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Al～Mohanna S Y, Nott JA, Lane D J W. M-"midget"cells in the hepatopancreas of the shrimp Penaeus semisulcatus 1844(Decapoda, Natantia) Crustaceana, 1985b, 48:260～268
Al-Mohanna S.Y., Nott J. A., B-cell and digestion in the hepatopancreas of the shrimp Penaeus semisulcatus(Crustacea, Decapoda). Mar. Biol. Ass U.K. 1986, 66:403～414
Al-Mohanna S.Y., Nott J. A., Lane D. J. W. Midgut cells in the hepatopancreas of the shrimp Penaeus semisulcatus(Crustacea, Deccapoda) Mar.Biol. Ass U.K, 1985,48:260～268
Al-Mohanna S.Y., Nott J. A. M-cell and moult cycle in Penaeus semisulcatus (Crustacea, Decapoda), Mar. Biol. Ass. U. K.,1987,61:803～813
Al-Mohanna S.Y., Nott J. A. R-cell and digestive cycle in Penaeus semisulcatus(Crustacea, Decapoda) Mar. Biol. Ass. U.K.,1987,95:129～137;
Al-Mohanna S.Y., Nott J.A., Lane D.J.W. Mitotic E-and secretary F-cell in the hepatopancreas of the shrimp Penaeus semisulcatus(Crustacea, Decapoda)Mar. Biol. Ass U.K.,1985,65:901-910
Anheller J. E. Hellgren L., Karlstam B., Vincent J. Biochemical and biological profile of a new enzyme preparation from Antarctic krill(E, superba) suitable for debridement of ulcerative lesions, Arch. Dermatol. Res. 1989, 281:105～110
Barker P.L., Gibson R., Observation on the structure of the mouthparts, histology of the alimentary tract and digestive physiology of the mud crab Scylla serrata (Decapoda, Portunidae) Exp. Mar. Biol. Ecol.,1978,32:177～196
Barker P.L.,Gibson R. Observation on the feeding mechanism, structure of the gut, and digestive physiology of the European lobster Homarus gammarus(Decapoda, Nephropidae) Exp. Mar. Biol. Ecol., 1977,26:297～324
Barrett A J,McDonld JK.Mammalian proteases.A glossary and bibliography. Vol.1.Endoproteinases. Toronto:Academic Press, 1980
Berg CH, Kalfas S, Maimasten M, Arnebrant T. Proteolytic degradation of oral biofilms in vitro and in vivio: potential of proteases originating from Euphausia superba for plaque control. Eur J Oral Sci 2001, 109(5):316～24
Brinton, E.O. Population biology of Euphausia pacifica off southern California. Fisheries Bulletin 1976, 74: 733～762.
Brocherhoff H, Hoyle R. J. Digestive enzyme of the american lobster. J Fish Res Board of
Canada, 1970,27(8) : 1357-1370
Bucht A, Karlstam, B. Isolation and immunological characterization of three highly purified serine proteinases from Antarctic krill (Euphausia superba), Polar Biol. 1991,11:495-500
Bustos R. O., Romo C. R., Healy M. G. Purification of trypsin-like enzymes from Antarctic krill processing wasterwater. Process Biochemistry.1999, 35:327-333
Cacei T. Ultrastructure of the hepatopancreas of the Pacific white shrimp, Penaeus vannamei (Crustacea, Decapoda) Mar. Biol. Ass. U.K., 1983,68:323-327
Chen C. S., Yan T. R., Chen H.Y. Purification and properties of trypsin-like enzymes and a carboxypeptidase A from Euphausia superba, J. Food Biochem. 1979, 2:349-366
Dall W 著，陈楠生等译。对虾生物学。青岛：青岛海洋大学出版社，1992
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Endo, Y. Ecological Studies on the Euphausiids Occurring in the Sanriku Waters with Special Reference to Their Life History and Aggregated Distribution. Sendai: Tohoku University. (in Japanese), 1981:166
Friedrich Buchholz, Reinhard Saborowski, Metabolic and enzymatic adaptations in northern krill, Meganyctiphanes norvegica, and Antarctic krill, Euphausia superba, Can. J. Fish. Aquat. Sci.,2000, 57(Suppl. 3) : 115-129
Glenner BC, Coben LA. Histochemical demonstration of a species-specific tryptase-like enzyme in mast cells. Nature,1960, 185(3) :846
Gunther J, Augustin M, Vanscheidt W. Krill-Enzyme: Ein viel versprechendes Therapieprinzip in der Behandlung chronischer Wunden. Ars Medici 2002,7: 310-313．
Haig-Brown, A. Going for the krill. In National Fisherman, 1994, 5: 18-19．
Heath, W.A. The Ecology and Harvesting of Euphausiids in the Strait of Georgia. Vancouver: University of British Columbia. 1977:187
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Hellgren L., Karlstam B., Mohr V, Vincent J. Krill enzymes. A new concept for efficient debridement of necrotic ulcers. Int J Dermatol 1991,30(2) : 102-103
Hirche H.J., Anger K. Digestive enzyme activities during larval development of Hyasaraneus, Comp. Biochem. Physiol., 1987,87B(2) :297-302
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许实荣等：维生素B1和B62对中国对虾消化酶的影响。海洋科学，1987(4)：34～37
杨德渐、王永良编著：中国北部海洋无脊椎动物。北京：高等教育出版社，1996：324～325
于书坤，张树荣：虾类及甲壳动物消化酶研究的现状。海洋科学，1986，10(2)：60～63
于书坤：中国对虾消化酶的研究Ⅰ：消化酶的活力测定及性质的研究。海洋科学集刊，第28集，1987：85～90
张志峰，姜明：对中国对虾幼体中肠腺的结构和细胞化学研究。中国水产科学，2000，7(3)：11～15
Allahpichay, I, Shimizu, C. Separation of growth promoting factors from non-muscle krill meal of Euphausia superba. Bull. Japanese Soc. Sci. Fish, 1985, 51:945～951
Al～Mohanna S Y, Nott JA, Lane D J W. M-"midget"cells in the hepatopancreas of the shrimp Penaeus semisulcatus 1844(Decapoda, Natantia) Crustaceana, 1985b, 48:260～268
Al-Mohanna S.Y., Nott J. A., B-cell and digestion in the hepatopancreas of the shrimp Penaeus semisulcatus(Crustacea, Decapoda). Mar. Biol. Ass U.K. 1986, 66:403～414
Al-Mohanna S.Y., Nott J. A., Lane D. J. W. Midgut cells in the hepatopancreas of the shrimp Penaeus semisulcatus(Crustacea, Deccapoda) Mar.Biol. Ass U.K, 1985,48:260～268
Al-Mohanna S.Y., Nott J. A. M-cell and moult cycle in Penaeus semisulcatus (Crustacea, Decapoda), Mar. Biol. Ass. U. K.,1987,61:803～813
Al-Mohanna S.Y., Nott J. A. R-cell and digestive cycle in Penaeus semisulcatus(Crustacea, Decapoda) Mar. Biol. Ass. U.K.,1987,95:129～137;
Al-Mohanna S.Y., Nott J.A., Lane D.J.W. Mitotic E-and secretary F-cell in the hepatopancreas of the shrimp Penaeus semisulcatus(Crustacea, Decapoda)Mar. Biol. Ass U.K.,1985,65:901-910
Anheller J. E. Hellgren L., Karlstam B., Vincent J. Biochemical and biological profile of a new enzyme preparation from Antarctic krill(E, superba) suitable for debridement of ulcerative lesions, Arch. Dermatol. Res. 1989, 281:105～110
Barker P.L., Gibson R., Observation on the structure of the mouthparts, histology of the alimentary tract and digestive physiology of the mud crab Scylla serrata (Decapoda, Portunidae) Exp. Mar. Biol. Ecol.,1978,32:177～196
Barker P.L.,Gibson R. Observation on the feeding mechanism, structure of the gut, and digestive physiology of the European lobster Homarus gammarus(Decapoda, Nephropidae) Exp. Mar. Biol. Ecol., 1977,26:297～324
Barrett A J,McDonld JK.Mammalian proteases.A glossary and bibliography. Vol.1.Endoproteinases. Toronto:Academic Press, 1980
Berg CH, Kalfas S, Maimasten M, Arnebrant T. Proteolytic degradation of oral biofilms in vitro and in vivio: potential of proteases originating from Euphausia superba for plaque control. Eur J Oral Sci 2001, 109(5):316～24
Brinton, E.O. Population biology of Euphausia pacifica off southern California. Fisheries Bulletin 1976, 74: 733～762.
Brocherhoff H, Hoyle R. J. Digestive enzyme of the american lobster. J Fish Res Board of
Canada, 1970,27(8) : 1357-1370
Bucht A, Karlstam, B. Isolation and immunological characterization of three highly purified serine proteinases from Antarctic krill (Euphausia superba), Polar Biol. 1991,11:495-500
Bustos R. O., Romo C. R., Healy M. G. Purification of trypsin-like enzymes from Antarctic krill processing wasterwater. Process Biochemistry.1999, 35:327-333
Cacei T. Ultrastructure of the hepatopancreas of the Pacific white shrimp, Penaeus vannamei (Crustacea, Decapoda) Mar. Biol. Ass. U.K., 1983,68:323-327
Chen C. S., Yan T. R., Chen H.Y. Purification and properties of trypsin-like enzymes and a carboxypeptidase A from Euphausia superba, J. Food Biochem. 1979, 2:349-366
Dall W 著，陈楠生等译。对虾生物学。青岛：青岛海洋大学出版社，1992
Ellingsen T.E., Mohr V. Lipids in Antarctic krill. Composition and post mortem changes. Proc. 11th Scandinavian Symp. On Lipids (Edited by Marcusr R.),SIK, Gothenborg, Sweden. Ellingsen T.E. Biochemical studies on Antarctic krill (Ph.D. thesis). Department of Biochemistry, The Norwegian Institute of Technology, The University of Trondheim, N-7034 Trondheim-NTH, Norway. 1982: 382
Ellingsen TE, Mohr V., Biochemistry of the autolytic processes in Antarctic krill post mortem. Autoproteolysis. Biochem J., 1987 , 246(2) :295-305
Endo, Y. Ecological Studies on the Euphausiids Occurring in the Sanriku Waters with Special Reference to Their Life History and Aggregated Distribution. Sendai: Tohoku University. (in Japanese), 1981:166
Friedrich Buchholz, Reinhard Saborowski, Metabolic and enzymatic adaptations in northern krill, Meganyctiphanes norvegica, and Antarctic krill, Euphausia superba, Can. J. Fish. Aquat. Sci.,2000, 57(Suppl. 3) : 115-129
Glenner BC, Coben LA. Histochemical demonstration of a species-specific tryptase-like enzyme in mast cells. Nature,1960, 185(3) :846
Gunther J, Augustin M, Vanscheidt W. Krill-Enzyme: Ein viel versprechendes Therapieprinzip in der Behandlung chronischer Wunden. Ars Medici 2002,7: 310-313．
Haig-Brown, A. Going for the krill. In National Fisherman, 1994, 5: 18-19．
Heath, W.A. The Ecology and Harvesting of Euphausiids in the Strait of Georgia. Vancouver: University of British Columbia. 1977:187
Hellgren L, Mohr V, Vincent J., Proteases of Antarctic krill-a new system for effective enzymatic debridement of necrotic ulcerations. Experientia. 1986, 42(4) : 403-404
Hellgren L., Karlstam B., Mohr V, Vincent J. Krill enzymes. A new concept for efficient debridement of necrotic ulcers. Int J Dermatol 1991,30(2) : 102-103
Hirche H.J., Anger K. Digestive enzyme activities during larval development of Hyasaraneus, Comp. Biochem. Physiol., 1987,87B(2) :297-302
Iguchi, N., Ikeda, T, Imamura A. Growth and life cycle of a euphausiid crustacean (Euphausia pacifica Hansen) in Toyama Bay, southern Japan Sea. Bull. Japan Sea Nat. Fish. Res. Inst. 1993, 43: 69-81． (in Japanese with English abstract)
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