В статье рассмотрены и обсуждены работы, посвящённые выделению фосфолипазы Д из различных источников и подробному исследованию её активации на границе раздела фаз при гидролизе лецитина и лизолецитина в присутствии ионов металлов, органических растворителей, детергентов, адсорбентов и продуктов гидролиза, а также в составе мембран митохондрий и микросом в различных условиях проведения реакции. На основании полученных результатов выдвинута гипотеза об активации, функционировании, регуляции и локализации фосфолипазы Д на границе раздела фаз липид – вода, липид – липид и липид – белок при образовании кластера, рафта или микродомена из продуктов гидролиза или их природных и синтетических аналогов в присутствии ионов двухвалентных металлов. Фосфолипаза Д рассматривается как примембранный векторный фермент, латеральный насос или как 2D-принтер, преобразующий субстрат в продукт с одновременным его депонированием с ионами Ca2+ в разрастающийся новый фосфатидатный микродомен. Обсуждены возможные приложения предложенного механизма регуляции липолиза в решении ряда вопросов и проблем мембранологии, биотехнологии, питания и медицины.
В статье рассмотрены и обсуждены работы, посвящённые выделению фосфолипазы Д из различных источников и подробному исследованию её активации на границе раздела фаз при гидролизе лецитина и лизолецитина в присутствии ионов металлов, органических растворителей, детергентов, адсорбентов и продуктов гидролиза, а также в составе мембран митохондрий и микросом в различных условиях проведения реакции. На основании полученных результатов выдвинута гипотеза об активации, функционировании, регуляции и локализации фосфолипазы Д на границе раздела фаз липид – вода, липид – липид и липид – белок при образовании кластера, рафта или микродомена из продуктов гидролиза или их природных и синтетических аналогов в присутствии ионов двухвалентных металлов. Фосфолипаза Д рассматривается как примембранный векторный фермент, латеральный насос или как 2D-принтер, преобразующий субстрат в продукт с одновременным его депонированием с ионами Ca2+ в разрастающийся новый фосфатидатный микродомен. Обсуждены возможные приложения предложенного механизма регуляции липолиза в решении ряда вопросов и проблем мембранологии, биотехнологии, питания и медицины.
Мазкур мақолада фосфолипаза Д ни турли тирик организмлардан ажратиб олишга доир тадқиқотлар, лецитин ва лизолецитин, металл ионлари, органик эритувчилар, детергентлар, адсорбентлар, гидролиз маҳсулотлари ёки уларнинг табиий ва синтетик аналоглари иштирокида ва турли реакция шароитида бу ферментнинг фаоллашуви батафсил ўрганилган ва муҳокама қилинган. Олинган натижалар асосида гидролиз маҳсулотларидан кластер, рафт ёки микродомен ҳосил бўлиши пайтида липид – сув, липид – липид ва липид – оқсил фазалари чегарасида фосфолипаза Д нинг фаоллашиши, ишлаши, регуляцияси ва локализацияси тўғрисида янги гипотеза таклиф қилинган. Фосфолипаза Д га сирт мембранали вектор ферменти, латерал насоси ёки 2D принтери сифатида қаралади. Фосфолипаза Д фосфолипид субстрати Cа2+ ионлари билан бир вақтда ўсиб бораётган микродоменда янги фосфатид маҳсулотига айланади. Шунингдек, мақолада мембранология, биотехнология, овқатланиш ва тиббиётнинг бир қатор масалалари ва муаммоларини ҳал қилишда липолизни тартибга солиш механизмининг мумкин бўлган қўлланилиши муҳокама қилинган.
The review elucidates works on isolation of phospholipase D from various sources, a detailed study of its activation at the interface during hydrolysis of lecithin and lysolecithin in the presence of metal ions, organic solvents, detergents, adsorbents and products of hydrolysis, as well as in composition of mitochondrial membranes and microsomes under various reaction conditions. Based on the research findings, a hypothesis is being put forward about activation, functioning, regulation and localization of phospholipase D at the interface of lipid-water, lipidlipid and lipid-protein phases in the course of forming of a cluster, raft or microdomain with products of hydrolysis or their natural and synthetic analogues in the presence of bivalent metal ions. Phospholipase D is considered as a pri-membrane vector enzyme, a lateral pump, or as a 2D printer that converts a substrate into a product while depositing it with Ca2+ ions into a growing new phosphatidate microdomain. Possible applications of the proposed mechanism of lipolysis regulation in solving various issues of membranology, biotechnology, nutrition and medical practice are being discussed.
| № | Author name | position | Name of organisation |
|---|---|---|---|
| 1 | Madyarov S.R. | biologiya fanlar doktori, yetakchi ilmiy xodim | O‘zbekiston Respublikasi Fanlar Akademiyasi Bioorganik kimyo instituti, Mirzo Ulug‘bek nomidagi O‘zbekiston milliy universiteti Biofizika va biokimyo instituti |
| № | Name of reference |
|---|---|
| 1 | Abousalham, A., Nari, J., & Noat, G. (1999). Phospholipase D from plants: purification, enzymology and structural analysis. Plant Biology, 1, 123–132. |
| 2 | Avlyanova, R. R., Akhmedzhanov, R., Rakhimov, M. M., Madyarov, Sh. R., Muratova, U. Z., & Tashmukhamedov, B. A. (1981). Sposob polucheniya adsorbenta dlya gidrofobnoy khromatografii [Method for producing an adsorbent for hydrophobic chromatography]. (In Russian). (USSR Auth. certificate, No. 853990). |
| 3 | Babayev, M. U., Khole, V., Madyarov, Sh. R., Rakhimov, M. M., & Tashmukhamedov, B. A. (1980). Fermentativnyy sintezfosfolipidov s identichnymi zhirnymi kislotami dlya polucheniya liposom [Enzymatic synthesis of phospholipids with identical fatty acids to obtain liposomes]. (In Russian). Proceedings of the AllUnion Symposium “Liposomes. Interaction with cells and tissues”: pp. 20–21. Moscow. |
| 4 | Benzonana, G., & Desnuelle, P. (1965). Kinetic study of the action of pancreatic lipase on emulsified triglycerides. Enzymology assay in heterogeneous medium. Biochim. Biophys. Acta, 105 (1), 121–136. |
| 5 | Bogdanov, M., Umeda, M., & Dowhan, W. (1999). Phospholipid assisted refolding оf an integral membrane protein. Minimum structural features for phosphatidyl ethanolamine to act as a molecular chaperone. Biol. Chem., 274, 12339–12345. |
| 6 | Brokerkhof, Kh., & Dzhensen, R. (1978). Lipoliticheskiye fermenty [Lipolytic enzymes]. (In Russian). Moscow: MIR. |
| 7 | Bruntz, R. C., Lindsley, C. W., & Brown, H. A. (2014). Phospholipase D signaling pathways and phosphatidic acid as therapeutic targets in cancer. Pharmacol Rev., 66, 1033–1079. |
| 8 | Dennis, E. A. (Ed.) (1991). Phospholipases, Methods in Enzymology. N.Y.: Academic Press. |
| 9 | Donaldson, J. G. (2009). Phospholipase D in endocytosis and endosomal recycling pathways. Biochim. Biophys. Acta, 1791 (9), 845–849. |
| 10 | Frieden, C. (1970). Kinetic aspects of regulation of metabolic processes. The hysteretic enzyme concept. Biol. Chem., 245, 5788–5799. |
| 11 | Frohman, M. A. (2015). The phospholipase D superfamily as therapeutic targets. Trends Pharmacol. Sci., 36 (3), 137–144. doi:10.1016/j.tips.2015.01.001 |
| 12 | Harauma, A., Yasuda, H., Hatanaka, E., Nakamura, M. T., Salem N. Jr., & Moriguchi, T. (2017). Prostaglandins, Leukotrienes. Essenttial Fatty Acids, 116, 9–18. |
| 13 | Huang, H., & Frohman, M. A. (2009). Lipid Signaling on the Mitochondrial Surface. Biochim. Biophys. Acta, 1791 (9), 839–844. |
| 14 | Huang, P., Jiang, Z., Teng, S., Wong, Y. C., Frohman, M. A., Chung, S. K, & Chung, S. S. (2006). Synergism between phospholipase D2 and sorbitol accumulation in diabetic cataract formation through modulation of Na, K-ATPase activity and osmotic stress. Exp. Eye Res., 83 (4), 939–948. |
| 15 | Ito, T., & Ohnishi, S-I. (1974). Ca2+-induced lateral phase separation in РА – РС membranes. Biochim. Biophys. Acta, 352 (1), 29–37. |
| 16 | Jain, M. K., & Gelb, M.H. (1991). Phospholipase A2 catalyzed hydrolysis of vesicles: uses of interfacial catalysis in the scooting mode. Methods Enzymol., 197, 112–125. |
| 17 | Kagava, Y. (1985). Biomembrany [Biomembranes]. (In Russian). A. A. Selishchev (Trans.). Moscow: Vysshaya shkola. |
| 18 | Khole, V., & Madyarov, Sh. R. (1983). On interaction of cobra venom toxic polypeptides with different substrate systems. Indian. J. Biochem. Biophys., 20, 177–179. |
| 19 | Kravtsov, A. V., & Alekseyenko, I. R. (1990). Mekhanizm regulyatsii vektornykh fermentov biomembran [The mechanism of regulation of vector enzymes of biomembranes]. (In Russian). Kyiv: Naukova Dumka. |
| 20 | Lee, Y., & Chan, S. I. (1977). Effect of lysolecithin on the structure and permeability of lecithin bilayer vesicles. Biochemistry, 16 (7), 1303–1309. |
| 21 | Leiros, I., Secundo, F., Zambonelli, C., Servi, S., & Hough, E. (2000). The first crystal structure of phospholipase D. Structure, 8, 655–667. |
| 22 | Liscovitch, M., Gzarny, M., Fiucci, J., & Tang, X. (2000). Phospholipase D: molecular and cell biology of a novel gene family. Biochem. J., 345, 401–415. |
| 23 | Luzikov, V.N., Rakhimov, M. M., & Berezin, I. V. (1969). Effect of substrates on reconstitution of the mitochondrial respiratory chain under various conditions. Biochim. Biophys. Acta, 180 (3), 429–438. |
| 24 | Madyarov Sh. R., Ziyavitdinov, Sh. S., & Rakhimov, M. M. (1975). Aktivatsiya fosfolipazy D na granitse razdela faz [Activation of phospholipase D at the interface]. (In Russian). Proceedings of the symposium “Chemistry and biochemistry of proteins and peptides”: pp. 59–60. Tashkent. |
| 25 | Madyarov, Sh. R. (1976). Vydeleniye fosfolipazy D iz semyan khlopchatnika [Isolation of phospholipase D from cotton seed]. (In Russian). Biochemistry, 41, 255–259. |
| 26 | Madyarov, Sh. R. (1977). Izucheniye deystviya fosfolipazy D na granitse razdela faz [Study of the action of phospholipase D at the interface]. (In Russian). [PhD thesis]. Tashkent: TashSU. |
| 27 | Madyarov, Sh. R. (1985). Immobilizatsiya fosfolipaz na fibroine shelka [Immobilization of phospholipases on silk fibroin]. (In Russian). Proceedings of the I All-Union Сonference “Catalysis and catalytic processes in chemical pharmaceutical production”: Part II, p. 242–243. Moscow. |
| 28 | Madyarov, Sh. R. (1985). Immobilizatsiya fosfolipaz na fibroine shelka [Immobilization of phospholipases on silk fibroin]. (In Russian). Proceedings of the I All-Union Сonference “Catalysis and catalytic processes in chemical pharmaceutical production”: pp. 78–79. Tashkent. |
| 29 | Madyarov, Sh. R., & Rakhimov, M. M. (1989). Nekotoryye osobennosti funktsionirovaniya fosfolipazy D v razlichnykh substratnykh sistemakh [Some features of the functioning of phospholipase D in various substrate systems]. (In Russian). Biochemistry, 54, 1780–1789. |
| 30 | Madyarov, Sh. R., & Sadykova, N. (1989). Fosfolipaza D iz soi – preparat dlya gidroliza i sinteza fosfolipidov [Phospholipase D from soybean - a drug for the hydrolysis and synthesis of phospholipids]. (In Russian). Proceedings of the II All-Union Сonference “Catalysis and catalytic processes in chemical pharmaceutical production”: Part II, p. 235. Moscow. |
| 31 | Madyarov, Sh. R., Syunyaeva, S. T., & Chung, B.H. (2000). Effect of micellar substrates on phospholipase D reaction. Proceedings of the International Sympozium and Spring Meeting “Microorganisms and Health”: p. 228. Seoul, Korea. |
| 32 | Madyarov, Sh. R., & Chung, B.H. (2001). Sensorial properties of phospholipase D in reflection of physicochemical states of membrane-like structures. Proceedings of the 4th Int. Congress on Biol. Phys., ICBP. Kyoto, Japan. |
| 33 | Madyarov, Sh. R. (2002). Phospholipase D as vector controlling a membrane anisotropy. Proceedings of the 43rd ICBL (Graz, Austria). Chem. Phys. Lipids, 118 (1-2), 76. |
| 34 | Madyarov, Sh. R. (2002). Vector property of phospholipase D and membrane Regulation. Proceedings of the International Conference “Lipids and Biomembranes: New Technologies”: p. 80. Davos, Switzerland. |
| 35 | Madyarov, S. R. (2002). New trends in silk science and biotechnology. From bullet prove clothes to “magic bullet” concept. Chem. Phys. Lipids., 118 (1–2), 56–57. |
| 36 | Madyarov, S. R. (2004). Phospholipids clustering mode of phospholipase D action. Proceedings of the 5th Int. Conf. on Biol. Physics: B05-206, p. 101. Gothenburg, Sweden. |
| 37 | Madyarov, Sh. R. (2010). Biotekhnologicheskiye podkhody v shelkovodstve i tekhnologii shelka [Biotechnological approaches in sericulture and silk technology]. (In Russian). [Abstract of Doctoral Dissertation]. Tashkent: Institute of Microbiology. |
| 38 | Madyarov, Sh. R. (2010). Klasterobrazovaniye produkta reaktsii i vektornyye cvoystva fosfolipazy D [Clustering of the reaction product and vector properties of phospholipase D]. (In Russian). Biological Journal of Uzbekistan, spec. iss., 55–59. |
| 39 | Madyarov, Sh. R. (2012). Vydeleniye, ochistka, stabilizatsiya i kharakteristika preparatov fosfolipazy D iz Raphanus sativus longipinnatus [Isolation, purification, stabilization and characterization of phospholipase D preparations from Raphanus Sativus Longipinnatus]. (In Russian). Chemistry of Natural Compounds, 4, 499–501. |
| 40 | Madyarov, Sh. R. (2012). Vydeleniye, ochistka, kharakteristika i prolongirovannyye formy fosfolipazy D iz red’ki daikon [Isolation, purification, characterization and prolonged forms of phospholipase D from daikon radish]. (In Russian). Bulletin “TINBO”, 48–52. Tashkent: Chinor. |
| 41 | Madyarov, Sh. R. (2014). Deystviye detergentov, tripsina i ionov dvukhvalentnykh metallov na mezhfaznuyu aktivatsiyu i funktsionirovaniye fosfolipazy D [The effect of detergents, trypsin and divalent metal ions on the interphase activation and functioning of phospholipase D]. (In Russian). Biochemistry, 79 (7), 864–872. |
| 42 | Madyarov, Sh. (2015). Interfacial refolding and tuning of lipolytic enzymes at membrane like structures. Proceedings of the Int. Conf. on Protein Engineering (Chicago. USA). J. Proteomics Bioinform., 8, 10. doi:10.4172/0974-276X.C1.080 |
| 43 | Majd, Sh., Yusko, E. C., Yang, J., Sept, D., & Mayer, M. (2013). A Model for the Interfacial Kinetics of Phospholipase D Activity on Long-Chain Lipids. Biophys., 105 (7), 146–153. |
| 44 | Mokhova, Ye.N., Starkov, A. A., & Bobyleva, V. A. (1993). Razobshcheniye okislitel’nogo fosforilirovaniya zhirnymi kislotami v mitokhondriyakh pecheni i myshts [Uncoupling of oxidative phosphorylation by fatty acids in liver and muscle mitochondria]. (In Russian). Biochemistry, 58, 1513–1522. |
| 45 | Mouritsen, O. G. (2011). Lipids, curvature, and nano-medicine. Eur. J. LipidSci. Technol., 13 (10), 1174–1187. |
| 46 | Nicolson, G. L. (2014). The Fluid Mosaic Model of Membrane Structure: Stillrelevant to understanding the structure, function and dynamics of biological membranes after more than 40 years: Review. Biochim. Biophys. Acta, 1838 (6), 1451–1466. |
| 47 | Oliveira, T. G., & Di, P. G. (2010). Phospholipase D in brain function and Alzheimer’s desease. Biochim. Biophys. Acta, 1801 (8), 799–805. doi:10.1016/j.bbalip.2010.04.004 |
| 48 | Op den Kamp, J. A., De Gier, J., & Van Deenen, L. L. M. (1974). Hydrolysis of phosphatidilcholine liposomes by pancreatic phospholipase A2 at the transitiontemperature. Biochim. Biophys. Acta, 345, 253–256. |
| 49 | Panaiotov, I., & Verger, R. (2000). Enzymatic reaction at interface: Interfacial and temporal organization of enzymatic lipolysis. In Physical chemistry of biological interface (A. Baszkin, & W. Norde, Eds., pp. 359–400). Marcel Dekker, Inc. NY-Basel. |
| 50 | Peng, X., & Frohman, M. A. (2012). Mammalian Phospholipase D Physiological and Pathological Roles. Acta Physiol (Oxf), 204 (2), 219–226. doi:10.1111/j.1748-1716.2011.02298.x |
| 51 | Pieterson, W. A., Volwerk, J. J., & de Haas, G. H. (1974). Interaction of phospholipaseA2 and its zymogen with divalent metal ions. Biochemistry, 13, 1439–1445. |
| 52 | Quarles, R. H., & Dawson, R. M. C. (1969). The hydrolysis of monolayers of Рhosphatidyl-(Me-14C) choline by phospholipase D. Biochem., 113 (5), 697–705. |
| 53 | Rakhimov, M. M., Madyarov, Sh. R., Ziyavitdinov, Sh. S., & Abdumalikov, A. Kh. (1976). Immobilizovannaya fosfolipaza D [Immobilized phospholipase D]. (In Russian). Biochemistry, 41, 569–573. |
| 54 | Rakhimov, M. M., Madyarov, Sh. R., Ziyavitdinov Sh. S., & Abdumalikov A. Kh. (1976). Aktivatsiya rastitel’noy fosfolipazy D silikagelem [Activation of plant phospholipase D by silica gel]. (In Russian). Biological Journal of Uzbekistan, 3, 3–6. |
| 55 | Rakhimov, M. M., Madyarov, Sh. R., & Abdumalikov, A. Kh. (1976). Ob aktivatorakh fosfolipazy D [About phospholipase D activators]. (In Russian). Applied Biochemistry and Microbiology, 12 (1), 30–32. |
| 56 | Rakhimov, M. M., Madyarov, Sh. R., & Abdumalikov, A. Kh. (1977). Sposob polucheniya fosfolipazy D [Method for producing phospholipase D]. (In Russian). (USSR Auth. certificate, No. 560614). |
| 57 | Rakhimov, M. M., Madyarov, Sh. R., Ziyavitdinov, Sh. S., & Abdumalikov, A. Kh. (1977). Obnaruzheniye i svoystva dvukh form fosfolipazy D rasteniy [Detection and properties of two forms of plant phospholipase D]. (In Russian). Biochemistry, 42 (5), 788–798. |
| 58 | Rakhimov, M. M., & Madyarov, Sh. R. (1977). Sostoyaniye fosfolipazy D v rastvore i yeyo kataliticheskaya aktivnost’ [The state of phospholipase D in solution and its catalytic activity]. (In Russian). Biochemistry, 42, 622–635. |
| 59 | Rakhimov, M. M., Madyarov, Sh. R., Kholodkova, T. P., Babayev, M. U., Rashidova S. Sh., Kalendareva, T. I., Almatov, K. T., Mirsalikhova, N. M., & Mirkhodzhayev, U. Z. (1978). Vliyaniye ionov kal’tsiya na fermentativnyy gidroliz fosfolipidov v zavisimosti ot fizicheskogo sostoyaniya substrata. The influence of calcium ions on the enzymatic hydrolysis of phospholipids depending on the physical state of the substrate]. (In Russian). Biochemistry, 43 (2), 433–445. |
| 60 | Rakhimov, M. M., Madyarov, Sh. R., & Babayev, M. U. (1979). Fermentativnyy sintez fosfolipidov na poverkhnosti razdela [Enzymatic synthesis of phospholipids at interfaces]. (In Russian). Biological Journal of Uzbekistan, 3, 6–10. |
| 61 | Rakhimov, M. M., Akhmedzhanov, P., Babayev, M. U., Khole, V., & Madyarov, Sh. R. (1981). Svoystva fosfolipazy D iz Raphanus Sativu [Properties of phospholipase D from Raphanus Sativu]. (In Russian). Biochemistry, 46 (2), 240–249. |
| 62 | Rakhimov, M. M., & Madyarov, Sh. R. (1982). Rol’ fazovykh sostoyaniy fosfolipidov v regulirovanii kataliticheskikh effektov fosfolipaz v iskusstvennykh i membrannykh sistemakh [The role of phase states of phospholipids in the regulation of the catalytic effects of phospholipases in artificial and membrane systems, in collection]. (In Russian). In B. A. Tashmukhamedov (Ed.), Transport of ions through biological membranes and the mechanism of action of physiologically active substances (pp. 186–230). Tashkent: FAN. |
| 63 | Rakhimov, M. M., Akhmedzhanov, R., Madyarov, Sh. R., & Tashmukhamedov, B.A. (1982). Razdeleniye i svoystva molekulyarnykh form fosfolipazy D iz rasteniy [Separation and properties of molecular forms of phospholipase D from plants]. (In Russian). Biochemistry, 47 (9), 1454–1465. |
| 64 | Rakhimov, M. M., Kalendareva, T. I., Rashidova, S. Sh., & Madyarov, Sh. R. (1982). Rol’ ionov kal’tsiya v razvitii kataliticheskoy aktivnosti fosfolipazy D [The role of calcium ions in the development of the catalytic activity of phospholipase D]. (In Russian). Biochemistry, 47 (10), 1649–1662. |
| 65 | Reichert, A., Ringsdorf, H., & Wagenknecht, A. (1992). Spontaneous domain formation оf phospholipase A2 at interfaces: fluorescence microscopy of the interaction of phospholipase A2 with mixed monolayers of lecithin, lysolecithin and fatty acid. Biochim. Biophys. Acta, 1106 (1), 178–188. |
| 66 | Rudge, S. A., & Wakelam, M. J. (2009). Inter-regulatory dynamics of PLD and the actin cytoskeleton. Biochim. Biophys. Acta, 1791 (9), 856–861. |
| 67 | Stegner, D., Thielmann, I., Kraft, P., Frohman, M.A., Stoll, G., & Nieswandt, B. (2013). Pharmacological inhibition of phospholipase D protects mice from occlusive thrombus formation and ischemic stroke-brief report. Arterioscler. Thromb. Vasc. Biol., 33 (9), 2212–2217. doi:10.1161/ATVBAHA.113.302030. |
| 68 | Taniguchi, Y., Ohba, T., Miyata, H., & Ohki, K. (2006). Rapid phase change of lipid microdomains in giant vesicles induced by conversion of sphingomyelin to ceramide. Biochim. Biophys. Acta, 1758, 145–153. |
| 69 | Tinker, D. O., & Wei, J. (1979). Heterogeneous catalysis of phospholipase A2: formulation of a kinetic description of surface effects. Can. J. Biochem., 57, 97–106. |
| 70 | Wells, M. A. (1974). The mechanism of interfacial activation of phospholipase A2. Biochemistry, 13, 2248–2257. |
| 71 | Wells, M. A. (1974). А phospholipase A2 model system. Calcium enhancement of the аmine catalyzed methanolysis of phosphatidylcholine. Biochemistry, 13, 2258–2264. |