LEVAMISOLE MODULA A IMUNIDADE MEDIADA POR CÉLULAS DE MATRINXí, Brycon amazonicus
DOI:
https://doi.org/10.20950/1678-2305.2019.45.2.445Palavras-chave:
imunoestimulante, imunidade celular, imunidade inata, leucócitosResumo
A utilização de imunoestimulante é uma alternativa para melhorar o sistema imune de peixes e evitar o uso excessivo de antibióticos. O levamisol é um anti-helmíntico sintético que promove uma potente estimulação imunológica em peixes, aumentando variáveis inatas e adquiridas de defesa. A fim de avaliar os efeitos do levamisol sobre matrinxã, Brycon amazonicus, os peixes foram alimentados com 500 mg kg-1 de levamisol na dieta durante sete dias. Após este período os peixes foram submetidos í colheita de sangue. Variáveis imunes, tais como a atividade respiratória de leucócitos, proteína total sérica, albumina, globulina, índice albumina:globulina e os parí¢metros hematológicos, tais como hematócrito, hemograma completo, hemoglobina, número de eritrócitos e o volume corpuscular médio dos eritrócitos foram avaliados. A utilização do levamisol aumentou a contagem dos leucócitos do sangue indicando que o imunoestimulante modulou a imunidade mediada por células, entretanto, nenhuma alteração foi observada nas demais variáveis. O aumento dos leucócitos do sangue indica que os peixes alimentados com levamisol estão em uma condição corporal melhor, e em um possível surto terão mais resistência í s doenças e potencial para ativação da fagocitose e degranulação de neutrófilos. Além disso, os resultados são importantes para novos projetos sobre a produção de vacinas, e sobre a compreensão mais profunda da imunidade mediada por células em peixes.
Referências
Biller-Takahashi, J.D.; Montassier, H.J.; Takahashi, L.S.; Urbinati, E.C. 2016. Levamisole promotes an adjuvant effect on the immunityof pacu (Piaractus mesopotamicus) when immunized with Aeromonas hydrophila, even when provided in the diet. Animal Feed Science and Technology, 211(1): 164í 173. https://doi.org/10.1016/j.anifeedsci.2015.11.008
Biller-Takahashi, J.D.; Takahashi, L.S.; Mingatto, F.E., Urbinati, E.C. The immune system is limited by oxidative stress: Dietary selenium promotes optimal antioxidative status and greatest immune defense in pacu Piaractus mesopotamicus. Fish and Shellfish Immunology, 47(1): 360-367, 2015. http://dx.doi.org/10.1016/j.fsi.2015.09.022
Biller-Takahashi, J.D.; Takahashi, L.S.; Saita, M.V.; Gimbo, R.Y.; Urbinati, E.C. 2013. Leukocytes respiratory burst activity as indicator of innate immunity of pacu Piaractus mesopotamicus. Brazilian Journal of Biology, 73(2): 425-429. http://dx.doi.org/10.1590/S1519-69842013000200026
Biller-Takahashi, J.D.; Urbinati, E.C. 2014. Fish Immunology. The modification and manipulation of the innate immune system: Brazilian studies. Academia Brasileira de Ciências, 86(3): 75-87. http://dx.doi.org/10.1590/0001-3765201420130159
Costa, D.V.; Ferreira, M.W.; Navarro, R.D.; Rosa, P.V.; Murgas, L.D.S. 2014. Parí¢metros hematológicos de tilápias-do-Nilo (Oreochromis niloticus) alimentadas com diferentes fontes de óleo. Revista Brasileira de Saúde e Produção Animal, 15(3): 754-764.
Cuesta, A.; Esteban, M.A.; Meseguer, J. 2002. Levamisole is a potent enhancer of gilthead sea bream natural cytotoxic activity. Veterinary Immunology and Immunopathology 89(2): 169-174. https://doi.org/10.1016/S0165-2427(02)00220-9
Doumas, B.T.; Watson, W.A.; Biggs, H.G. 1971. Albumin standards and the measurement of serum albumin with bromocresol green. Clinical Chimica Acta, 31(1): 87-96.
Fazio, F. 2019. Fish hematology analysis as an important tool of aquaculture: A review. Aquaculture, 500(1): 237í 242. https://doi.org/10.1016/j.aquaculture.2018.10.030
Faurschou, M.; Borregaard, N. 2003. Neutrophils granules and secretory vesicles in inflamation. Microbes Infection, 5(14): 1317-1327.
Ferraz, F.B.; Gomes, L.C. 2009. Social relationship as inducer of immunological and stress responses in matrinxã (Brycon amazonicus). Comparative Biochemistry and Physiology Part A, 153(3): 293-296. https://doi.org/10.1016/j.cbpa.2009.03.002
Ishikawa, N.M.; Ranzani-Paiva, M.J.T.; Lombardi, J.V. 2008. Metodologia para quantificação de leucócitos totais em peixes, Oreochromis niloticus. Archives of Veterinary Science, 13(1): 54-63. http://dx.doi.org/10.5380/avs.v13i1.11560
Ispir, U.; Yonar, M.E. 2007. The effects of levamisole on phagocytic activity of rainbow trout (Oncorhynchus mykiss W.). Acta Veterinária BRNO, 76(3): 493-497. https://doi.org/10.2754/avb200776030493
Ispir, U.; Dorucu, M. 2005. A study on the effects of levamisole on the immune system of rainbow trout (Oncorhynchus mykiss, Walbaum). Turkish Journal of Veterinary and Animal Science, 29(1): 1169-1176.
Jeney, G.; Anderson, D.P. 1993. Enhanced immune response and protection in rainbow trout to Aeromonas salmonicida bacterin following prior immersion in immunostimulants. Fish and Shellfish Immunology, 3(1): 51-58. https://doi.org/10.1006/fsim.1993.1005
Kajita, Y.; Sakai, M.; Atsuta, S.; Kobayashi, M. 1990. The immunomodulatory effects of levamisole on rainbow trout, Oncorhynchus mykiss. Fish Pathology, 25(1): 93-98. https://doi.org/10.3147/jsfp.25.93
Korytá, T.; Thi, D.; Takizawa, H.; Kí¶llner, F. .2013. Multicolour flow cytometry identifying defined leukocyte subsets of rainbow trout (Oncorhynchus mykiss). Fish and Shellfish Immunology, 35(6): 2017-2019. https://doi.org/10.1016/j.fsi.2013.09.025.
Li, P.; Li, P.; Wang, X.; Gatliniii, D.M. 2006. Evaluation of levamisole as a feed additive for growth and health management of hybrid striped bass (Morone chrysopsí"”Morone saxatilis). Aquaculture, 251(2-4): 201-209. https://doi.org/10.1016/j.aquaculture.2005.11.015
Maqsood, S.; Samoon, M.H.; Singh, P. 2009. Immunomodulatory and growth promoting effect of dietary levamisole in Cyprinus carpio fingerlings against the challenge of Aeromonas hydrophila. Turkish Journal of Fisheries and Aquatic Sciences, 9(1): 111-120.
Misra, C.K.; Das, B.K.; Mukherjee, S.C. 2009. Immune response, growth and survival of Labeo rohita fingerlings fed with levamisole supplemented diets for longer duration. Aquaculture Nutrition, 15(4): 356-365. https://doi.org/10.1111/j.1365-2095.2008.00600.x
Morrison, R.N; Nowak, B.F.; Carson, J. 2001. The histopathological effects of a levamisole-adjuvanted Vibrio anguillarum vaccine on Atlantic salmon (Salmo salar L.). Aquaculture, 195(1-2): 23-33. https://doi.org/10.1016/S0044-8486(00)00546-9
Morrison, R.N; Fast, M.D.; Muise, D.M.; Easy, R.E.; Ross, N.W.; Johnson, S.C. 2000. The Effect of Levamisole as an Adjuvant on the Humoral Immune Response of Atlantic Salmon (Salmo solar L.). Bulletin of the European Association of Fish Pathologists, 20(3): 101-105. https://doi.org/10.1016/j.fsi.2005.11.010
Mulero, M.; Esteban, A.; Meseguer, J. 1998. In vitro levamisole fails to increase sea bream (Sparus aurata L.) phagocyte functions. Fish and Shellfish Immunology, 8(4): 315-318.
Ranzani-Paiva, M.J.T.; Silva-Souza, E.A.T. 2004. Hematologia de peixes brasileiros. In: Ranzani-Paiva M.J.T. et al. (Ed.). Sanidade de organismos aquáticos. São Paulo: Editora Varela, cap.4, 89-120.
Reich, R.; Mulvaney, P.; Robinson-Bostom, L. 2014. Antihelminthic Drugs. Side Effects of Drugs Annual, 36(1): 457-464.
Reinhold, J.G. 1953. Manual determination of serum total protein, albumin and globulin fractions by Biuret method. In: Standard methods in clinical chemistry (Reiner, M. ed), pp.88-90. New York: Academic Press.
Renoux, G. 1980. The general immunopharmacology of levamisole. Drugs, 20(2): 89-99.
Rieger, A.M.; Barreda, D.R. 2011. Antimicrobial mechanism of fish leukocytes. Developmental and Comparative Immunology, 35(12): 1238-1245. https://doi.org/10.1016/j.dci.2011.03.009
Rocha, R.L.; Ferreira, J.M.; Miranda Lopes, T.O.; Gomes, L.C. 2018. Morphometric, bromatological, and sensory analysis of nile tilapia vaccinated against Streptococcus agalactiae. Boletim do Instituto de Pesca, 44(1): 100-104. 0. http://dx.doi.org/10.20950/1678-2305.2018.281
Rosenfeld, G. 1947. Corante pancrômico para hematologia e citologia clínica. Nova combinação dos componentes do May-Grünwald e do Giemsa num só corante de emprego rápido. Memórias do Instituto Butantan, 20(1): 329-334.
Sado, R.Y.; Bicudo, A.J.A.; Cyrino, J.E.P. 2010. Dietary levamisole influenced hematological variables of juvenile pacu, Piaractus mesopotamicus (Holmberg 1887). Journal of the World Aquaculture Society, 41(1): 66-75. https://doi.org/10.1111/j.1749-7345.2009.00334.x
Sahoo, P.K.; Mukherjee, S.C. 2002. The effect of dietary immunomodulation upon Edwardsiella tarda vaccination in healthy and immunocompromised Indian major carp (Labeo rohita). Fish and Shellfish Immunology, 12(1): 1-16. https://doi.org/10.1111/j.1749-7345.2009.00334.x
Sakai, M. 1999. Current research status of fish immunostimulants. Aquaculture, 72(1-2): 63- 92. https://doi.org/10.1016/S0044-8486(98)00436-0
Schell, L.M.; Blumberg, B.S. 1977. The genetics of human serum albumin, function an uses. Pergamon Press. Oxford and New York, pp. 113-141.
Siwicki, A.K. 1989. Immunostimulating influence of levamisole on nonspecific immunity in carp (Cyprinus carpio). Developmental and Comparative Immunology, 13(1): 87-91. https://doi.org/10.1016/0145-305X(89)90021-9
Stuart, L.M.; Ezekowitz, R.A. 2005. Phagocytosis: elegant complexity. Immunity, 22(5): 539-550. https://doi.org/10.1016/j.immuni.2005.05.002
Tavares-Dias, M.; Affonso, E.G.; Oliveira, S.R.; Marcon, J.L.; Egami, M.I. 2008. Comparative study on hematological variables of farmed matrinxã, Brycon amazonicus Spix and Agassiz, 1829 (Characidae: Bryconinae) with others Bryconinae species. Acta Amazonica, 38(4): 799-806. http://dx.doi.org/10.1590/S0044-59672008000400026
Tavares-Dias, M., Moraes, F.R., Imoto, M.E. 2008. Haematological parameters in two neotropical freshwater teleost, Leporinus macrocephalus (Anostomidae) and Prochilodus lineatus (Prochilodontidae). Bioscience Journal, 24(3), 96í 101.
Urbinati, E.C.; Gonçalves, F.D.; Takahashi, L.S. 2010. Pacu (Piaractus mesopotamicus). In Espécies nativas para piscicultura no Brasil (Baldisseroto, B. and Gomes, L. C. Eds), Santa Maria, Santa Catarina: Editora UFSM, 2nd ed. pp. 225-246.
Utke, K.; Kock, H.; Schuetze, H.; Bergmann, S.M.; Lorenzen, N.; Einer-Jensen, K.; Koller, B.; Dalmo, R.A.; Velsely, T.; Ototake, M.; Fischer, U. 2008. Cell-mediated immune response in rainbow trout after DNA immunization against the viral hemorrhagic septicemia virus. Developmental and Comparative Immunology, 32(3): 239-252. https://doi.org/10.1016/j.dci.2007.05.010
Wintrobe, M.M. 1934. Variations on the size and haemoglobin content of erythrocytes in the blood various vertebrates. Folia Haematologica, 51(1): 32-49.
Zanuzzo, F.S.; Zaiden, S.F.; Senhorini, J.A.; Marzocchi-Machado, C.M.; Urbinati, E.C. 2015. Aloe vera bathing improved physical and humoral protection in breeding stock after induced spawning in matrinxã (Brycon amazonicus). Fish and Shellfish Immunology, 45(1): 132-140. https://doi.org/10.1016/j.fsi.2015.02.017
Zanuzzo, F.S.; Biller-Takahashi, J.D.; Urbinati, E.C. 2012. Effect of Aloe vera extract on the improvement of the respiratory activity of leukocytes of matrinxã during the transport stress. Revista Brasileira de Zootecnia, 41(10): 2299-2302. http://dx.doi.org/10.1590/S1516-35982012001000023
Biller-Takahashi, J.D.; Takahashi, L.S.; Mingatto, F.E., Urbinati, E.C. The immune system is limited by oxidative stress: Dietary selenium promotes optimal antioxidative status and greatest immune defense in pacu Piaractus mesopotamicus. Fish and Shellfish Immunology, 47(1): 360-367, 2015. http://dx.doi.org/10.1016/j.fsi.2015.09.022
Biller-Takahashi, J.D.; Takahashi, L.S.; Saita, M.V.; Gimbo, R.Y.; Urbinati, E.C. 2013. Leukocytes respiratory burst activity as indicator of innate immunity of pacu Piaractus mesopotamicus. Brazilian Journal of Biology, 73(2): 425-429. http://dx.doi.org/10.1590/S1519-69842013000200026
Biller-Takahashi, J.D.; Urbinati, E.C. 2014. Fish Immunology. The modification and manipulation of the innate immune system: Brazilian studies. Academia Brasileira de Ciências, 86(3): 75-87. http://dx.doi.org/10.1590/0001-3765201420130159
Costa, D.V.; Ferreira, M.W.; Navarro, R.D.; Rosa, P.V.; Murgas, L.D.S. 2014. Parí¢metros hematológicos de tilápias-do-Nilo (Oreochromis niloticus) alimentadas com diferentes fontes de óleo. Revista Brasileira de Saúde e Produção Animal, 15(3): 754-764.
Cuesta, A.; Esteban, M.A.; Meseguer, J. 2002. Levamisole is a potent enhancer of gilthead sea bream natural cytotoxic activity. Veterinary Immunology and Immunopathology 89(2): 169-174. https://doi.org/10.1016/S0165-2427(02)00220-9
Doumas, B.T.; Watson, W.A.; Biggs, H.G. 1971. Albumin standards and the measurement of serum albumin with bromocresol green. Clinical Chimica Acta, 31(1): 87-96.
Fazio, F. 2019. Fish hematology analysis as an important tool of aquaculture: A review. Aquaculture, 500(1): 237í 242. https://doi.org/10.1016/j.aquaculture.2018.10.030
Faurschou, M.; Borregaard, N. 2003. Neutrophils granules and secretory vesicles in inflamation. Microbes Infection, 5(14): 1317-1327.
Ferraz, F.B.; Gomes, L.C. 2009. Social relationship as inducer of immunological and stress responses in matrinxã (Brycon amazonicus). Comparative Biochemistry and Physiology Part A, 153(3): 293-296. https://doi.org/10.1016/j.cbpa.2009.03.002
Ishikawa, N.M.; Ranzani-Paiva, M.J.T.; Lombardi, J.V. 2008. Metodologia para quantificação de leucócitos totais em peixes, Oreochromis niloticus. Archives of Veterinary Science, 13(1): 54-63. http://dx.doi.org/10.5380/avs.v13i1.11560
Ispir, U.; Yonar, M.E. 2007. The effects of levamisole on phagocytic activity of rainbow trout (Oncorhynchus mykiss W.). Acta Veterinária BRNO, 76(3): 493-497. https://doi.org/10.2754/avb200776030493
Ispir, U.; Dorucu, M. 2005. A study on the effects of levamisole on the immune system of rainbow trout (Oncorhynchus mykiss, Walbaum). Turkish Journal of Veterinary and Animal Science, 29(1): 1169-1176.
Jeney, G.; Anderson, D.P. 1993. Enhanced immune response and protection in rainbow trout to Aeromonas salmonicida bacterin following prior immersion in immunostimulants. Fish and Shellfish Immunology, 3(1): 51-58. https://doi.org/10.1006/fsim.1993.1005
Kajita, Y.; Sakai, M.; Atsuta, S.; Kobayashi, M. 1990. The immunomodulatory effects of levamisole on rainbow trout, Oncorhynchus mykiss. Fish Pathology, 25(1): 93-98. https://doi.org/10.3147/jsfp.25.93
Korytá, T.; Thi, D.; Takizawa, H.; Kí¶llner, F. .2013. Multicolour flow cytometry identifying defined leukocyte subsets of rainbow trout (Oncorhynchus mykiss). Fish and Shellfish Immunology, 35(6): 2017-2019. https://doi.org/10.1016/j.fsi.2013.09.025.
Li, P.; Li, P.; Wang, X.; Gatliniii, D.M. 2006. Evaluation of levamisole as a feed additive for growth and health management of hybrid striped bass (Morone chrysopsí"”Morone saxatilis). Aquaculture, 251(2-4): 201-209. https://doi.org/10.1016/j.aquaculture.2005.11.015
Maqsood, S.; Samoon, M.H.; Singh, P. 2009. Immunomodulatory and growth promoting effect of dietary levamisole in Cyprinus carpio fingerlings against the challenge of Aeromonas hydrophila. Turkish Journal of Fisheries and Aquatic Sciences, 9(1): 111-120.
Misra, C.K.; Das, B.K.; Mukherjee, S.C. 2009. Immune response, growth and survival of Labeo rohita fingerlings fed with levamisole supplemented diets for longer duration. Aquaculture Nutrition, 15(4): 356-365. https://doi.org/10.1111/j.1365-2095.2008.00600.x
Morrison, R.N; Nowak, B.F.; Carson, J. 2001. The histopathological effects of a levamisole-adjuvanted Vibrio anguillarum vaccine on Atlantic salmon (Salmo salar L.). Aquaculture, 195(1-2): 23-33. https://doi.org/10.1016/S0044-8486(00)00546-9
Morrison, R.N; Fast, M.D.; Muise, D.M.; Easy, R.E.; Ross, N.W.; Johnson, S.C. 2000. The Effect of Levamisole as an Adjuvant on the Humoral Immune Response of Atlantic Salmon (Salmo solar L.). Bulletin of the European Association of Fish Pathologists, 20(3): 101-105. https://doi.org/10.1016/j.fsi.2005.11.010
Mulero, M.; Esteban, A.; Meseguer, J. 1998. In vitro levamisole fails to increase sea bream (Sparus aurata L.) phagocyte functions. Fish and Shellfish Immunology, 8(4): 315-318.
Ranzani-Paiva, M.J.T.; Silva-Souza, E.A.T. 2004. Hematologia de peixes brasileiros. In: Ranzani-Paiva M.J.T. et al. (Ed.). Sanidade de organismos aquáticos. São Paulo: Editora Varela, cap.4, 89-120.
Reich, R.; Mulvaney, P.; Robinson-Bostom, L. 2014. Antihelminthic Drugs. Side Effects of Drugs Annual, 36(1): 457-464.
Reinhold, J.G. 1953. Manual determination of serum total protein, albumin and globulin fractions by Biuret method. In: Standard methods in clinical chemistry (Reiner, M. ed), pp.88-90. New York: Academic Press.
Renoux, G. 1980. The general immunopharmacology of levamisole. Drugs, 20(2): 89-99.
Rieger, A.M.; Barreda, D.R. 2011. Antimicrobial mechanism of fish leukocytes. Developmental and Comparative Immunology, 35(12): 1238-1245. https://doi.org/10.1016/j.dci.2011.03.009
Rocha, R.L.; Ferreira, J.M.; Miranda Lopes, T.O.; Gomes, L.C. 2018. Morphometric, bromatological, and sensory analysis of nile tilapia vaccinated against Streptococcus agalactiae. Boletim do Instituto de Pesca, 44(1): 100-104. 0. http://dx.doi.org/10.20950/1678-2305.2018.281
Rosenfeld, G. 1947. Corante pancrômico para hematologia e citologia clínica. Nova combinação dos componentes do May-Grünwald e do Giemsa num só corante de emprego rápido. Memórias do Instituto Butantan, 20(1): 329-334.
Sado, R.Y.; Bicudo, A.J.A.; Cyrino, J.E.P. 2010. Dietary levamisole influenced hematological variables of juvenile pacu, Piaractus mesopotamicus (Holmberg 1887). Journal of the World Aquaculture Society, 41(1): 66-75. https://doi.org/10.1111/j.1749-7345.2009.00334.x
Sahoo, P.K.; Mukherjee, S.C. 2002. The effect of dietary immunomodulation upon Edwardsiella tarda vaccination in healthy and immunocompromised Indian major carp (Labeo rohita). Fish and Shellfish Immunology, 12(1): 1-16. https://doi.org/10.1111/j.1749-7345.2009.00334.x
Sakai, M. 1999. Current research status of fish immunostimulants. Aquaculture, 72(1-2): 63- 92. https://doi.org/10.1016/S0044-8486(98)00436-0
Schell, L.M.; Blumberg, B.S. 1977. The genetics of human serum albumin, function an uses. Pergamon Press. Oxford and New York, pp. 113-141.
Siwicki, A.K. 1989. Immunostimulating influence of levamisole on nonspecific immunity in carp (Cyprinus carpio). Developmental and Comparative Immunology, 13(1): 87-91. https://doi.org/10.1016/0145-305X(89)90021-9
Stuart, L.M.; Ezekowitz, R.A. 2005. Phagocytosis: elegant complexity. Immunity, 22(5): 539-550. https://doi.org/10.1016/j.immuni.2005.05.002
Tavares-Dias, M.; Affonso, E.G.; Oliveira, S.R.; Marcon, J.L.; Egami, M.I. 2008. Comparative study on hematological variables of farmed matrinxã, Brycon amazonicus Spix and Agassiz, 1829 (Characidae: Bryconinae) with others Bryconinae species. Acta Amazonica, 38(4): 799-806. http://dx.doi.org/10.1590/S0044-59672008000400026
Tavares-Dias, M., Moraes, F.R., Imoto, M.E. 2008. Haematological parameters in two neotropical freshwater teleost, Leporinus macrocephalus (Anostomidae) and Prochilodus lineatus (Prochilodontidae). Bioscience Journal, 24(3), 96í 101.
Urbinati, E.C.; Gonçalves, F.D.; Takahashi, L.S. 2010. Pacu (Piaractus mesopotamicus). In Espécies nativas para piscicultura no Brasil (Baldisseroto, B. and Gomes, L. C. Eds), Santa Maria, Santa Catarina: Editora UFSM, 2nd ed. pp. 225-246.
Utke, K.; Kock, H.; Schuetze, H.; Bergmann, S.M.; Lorenzen, N.; Einer-Jensen, K.; Koller, B.; Dalmo, R.A.; Velsely, T.; Ototake, M.; Fischer, U. 2008. Cell-mediated immune response in rainbow trout after DNA immunization against the viral hemorrhagic septicemia virus. Developmental and Comparative Immunology, 32(3): 239-252. https://doi.org/10.1016/j.dci.2007.05.010
Wintrobe, M.M. 1934. Variations on the size and haemoglobin content of erythrocytes in the blood various vertebrates. Folia Haematologica, 51(1): 32-49.
Zanuzzo, F.S.; Zaiden, S.F.; Senhorini, J.A.; Marzocchi-Machado, C.M.; Urbinati, E.C. 2015. Aloe vera bathing improved physical and humoral protection in breeding stock after induced spawning in matrinxã (Brycon amazonicus). Fish and Shellfish Immunology, 45(1): 132-140. https://doi.org/10.1016/j.fsi.2015.02.017
Zanuzzo, F.S.; Biller-Takahashi, J.D.; Urbinati, E.C. 2012. Effect of Aloe vera extract on the improvement of the respiratory activity of leukocytes of matrinxã during the transport stress. Revista Brasileira de Zootecnia, 41(10): 2299-2302. http://dx.doi.org/10.1590/S1516-35982012001000023
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2019-03-26
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Nota científica (Short Communication)
