Fitoesteroles en la salud humana: ¿amigos o enemigos?
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Resumen
Fitoesteroles son esteroides presentes en organismos fotosintéticos y son moléculas presentes frecuentemente en la dieta humana. Hay evidencia que señala que el consumo de fitoesteroles puede prevenir problemas cardiovasculares, principalmente disminuyendo la absorción del colesterol presente en nuestra dieta. Además, se han encontrado estudios que indican que los fitoesteroles tienen un efecto anticancerígeno. No obstante, la recomendación de incrementar la ingesta de fitoesteroles debería tomarse con cautela y bajo seguimiento médico puesto que se han reportado efectos adversos.
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Amiot, M. J., Knol, D., Cardinault, N., Nowicki, M., Bott, R., Antona, C., Borel, P., Bernard, J.-P., Duchateau, G., & Lairon, D. (2013). Comparable reduction in cholesterol absorption after two different ways of phytosterol administration in humans. European Journal of Nutrition, 52(3), 1215– 1222. https://doi.org/10.1007/s00394-0120432-3.
Assmann, G., Cullen, P., Erbey, J., Ramey, D. R., Kannenberg, F., & Schulte, H. (2006). Plasma sitosterol elevations are associated with an increased incidence of coronary events in men: Results of a nested case-control analysis of the Prospective Cardiovascular Münster (PROCAM) study. Nutrition, Metabolism, and Cardiovascular Diseases: NMCD, 16(1), 13–21. https://doi.org/10.1016/j.numecd.2005.04.001.
Awad, A. B., Chinnam, M., Fink, C. S., & Bradford, P. G. (2007). Beta-Sitosterol activates Fas signaling in human breast cancer cells. Phytomedicine: International Journal of Phytotherapy and Phytopharmacology 14(11), 747–754. https://doi.org/10.1016/j.phymed.2007.01.003.
Bae, H., Park, S., Yang, C., Song, G., & Lim, W. (2021). Disruption of Endoplasmic Reticulum and ROS Production in Human Ovarian Cancer by Campesterol. Antioxidants (Basel, Switzerland), 10(3), 379. https://doi.org/10.3390/antiox10030379.
Brown, A. W., Hang, J., Dussault, P. H., & Carr, T. P. (2010). Plant sterol and stanol substrate specificity of pancreatic cholesterol esterase. The Journal of Nutritional Biochemistry, 21(8), 736-740. https://doi.org/10.1016/j.jnutbio.2009.04.008.
Cabianca, A., Müller, L., Pawlowski, K., & Dahlin, P. (2021). Changes in the Plant β-Sitosterol/ Stigmasterol Ratio Caused by the Plant Parasitic Nematode Meloidogyne incognita. Plants (Basel, Switzerland), 10(2), 292. https://doi.org/10.3390/plants10020292.
Clouse, S. D. (2002). Arabidopsis mutants reveal multiple roles for sterols in plant development. The Plant Cell, 14(9), 1995– 2000. https://doi.org/10.1105/tpc.140930.
Fahy, D. M., O’Callaghan, Y. C., & O’Brien, N. M. (2004). Phytosterols: Lack of cytotoxicity but interference with beta-carotene uptake in Caco-2 cells in culture. Food Additives and Contaminants, 21(1), 42–51. https://doi.org/10.1080/02652030310001636921.
Han, J.-H., Yang, Y.-X., & Feng, M.-Y. (2008). Contents of Phytosterols in Vegetables and Fruits Commonly consumed in China. Biomedical and Environmental Sciences, 21(6), 449–453. https://doi.org/10.1016/S0895-3988(09)60001-5.
Hu, M., Yang, F., Huang, Y., You, X., Liu, D., Sun, S., & Sui, S.-F. (2021). Structural insights into the mechanism of human NPC1L1mediated cholesterol uptake. Science Advances, 7(29), eabg3188. https://doi.org/10.1126/sciadv.abg3188.
Huang, W.-Q., Chen, Y.-M., & Zhang, C.-X. (2017). Association between phytosterol intake and colorectal cancer risk: A case-control study. The British Journal of Nutrition, 117(6), 839–850. https://doi.org/10.1017/S0007114517000617.
Igel, M., Giesa, U., Lutjohann, D., & von Bergmann, K. (2003). Comparison of the intestinal uptake of cholesterol, plant sterols, and stanols in mice. Journal of Lipid Research, 44(3), 533–538. https://doi.org/10.1194/jlr.M200393-JLR200.
Jaceldo-Siegl, K., Lütjohann, D., Sirirat, R., Mashchak, A., Fraser, G. E., & Haddad, E. (2017). Variations in dietary intake and plasma concentrations of plant sterols across plant-based diets among North American adults. Molecular Nutrition & Food Research, 61(8). https://doi.org/10.1002/mnfr.201600828.
Le Goff, M., Le Ferrec, E., Mayer, C., Mimouni, V., Lagadic-Gossmann, D., Schoefs, B., & Ulmann, L. (2019). Microalgal carotenoids and phytosterols regulate biochemical mechanisms involved in human health and disease prevention. Biochimie, 167, 106–118. https://doi.org/10.1016/j.biochi.2019.09.012.
Li, K., Yuan, D., Yan, R., Meng, L., Zhang, Y., & Zhu, K. (2018). Stigmasterol exhibits potent antitumor effects in human gastric cancer cells mediated via inhibition of cell migration, cell cycle arrest, mitochondrial mediated apoptosis and inhibition of JAK/STAT signalling pathway. Journal of B.U.ON.: Official Journal of the Balkan Union of Oncology, 23(5), 1420–1425.
Liu, Y., Lei, L., Wang, X., Ma, K. Y., Li, Y. M., Wang, L., Man, S. W., Huang, Y., & Chen, Z. Y. (2015). Plasma cholesterol-raising potency of dietary free cholesterol versus cholesteryl ester and effect of β-sitosterol. Food Chemistry, 169, 277–282. https://doi.org/10.1016/j.foodchem.2014.07.123.
McCann, S. E., Freudenheim, J. L., Marshall, J. R., Brasure, J. R., Swanson, M. K., & Graham, S. (2000). Diet in the epidemiology of endometrial cancer in western New York (United States). Cancer Causes & Control: CCC, 11(10), 965–974. https://doi.org/10.1023/a:1026551309873.
Mendilaharsu, M., De Stefani, E., Deneo-Pellegrini, H., Carzoglio, J., & Ronco, A. (1998). Phytosterols and risk of lung cancer: A case-control study in Uruguay. Lung Cancer (Amsterdam, Netherlands), 21(1), 37–45. https://doi.org/10.1016/s0169-5002(98)00044-0.
Moghadasian, M. H., McManus, B. M., Godin, D. V., Rodrigues, B., & Frohlich, J. J. (1999). Proatherogenic and antiatherogenic effects of probucol and phytosterols in apolipoprotein E-deficient mice: Possible mechanisms of action. Circulation, 99(13), 1733–1739. https://doi.org/10.1161/01.cir.99.13.1733.
Nguyen, T. M., Sawyer, J. K., Kelley, K. L., Davis, M. A., Kent, C. R., & Rudel, L. L. (2012). ACAT2 and ABCG5/G8 are both required for efficient cholesterol absorption in mice: Evidence from thoracic lymph duct cannulation. Journal of Lipid Research, 53(8), 1598–1609. https://doi.org/10.1194/jlr.M026823.
Oh, M.-J., So, H.-J., Hong, E.-S., Shin, J.-A., & Lee, K.-T. (2021). Presence of Cholesterol in Non-Animal Organisms: Identification and Quantification of Cholesterol in Crude Seed Oil from Perilla frutescens and Dehydrated Pyropia tenera. Molecules (Basel, Switzerland), 26(12), 3767. https://doi.org/10.3390/molecules26123767.
Othman, R. A., Myrie, S. B., & Jones, P. J. H. (2013). Non-cholesterol sterols and cholesterol metabolism in sitosterolemia. Atherosclerosis, 231(2), 291–299. https://doi.org/10.1016/j.atherosclerosis.2013.09.038.
Pereira, T. S., Fonseca, F. A. H., Fonseca, M. I. H., Martins, C. M., Fonseca, H. A. R., Fonzar, W. T., Goulart, A. C., Bensenor, I. M., Lotufo, P. A., & Izar, M. C. (2021). Phytosterol consumption and markers of subclinical atherosclerosis: Cross-sectional results from ELSA-Brasil. Nutrition, Metabolism, and Cardiovascular Diseases: NMCD, 31(6), 1756–1766. https://doi.org/10.1016/j.numecd.2021.02.031.
Pinedo, S., Vissers, M. N., von Bergmann, K., Elharchaoui, K., Lütjohann, D., Luben, R., Wareham, N. J., Kastelein, J. J. P., Khaw, K.-T., & Boekholdt, S. M. (2007). Plasma levels of plant sterols and the risk of coronary artery disease: The prospective EPIC-Norfolk Population Study. Journal of Lipid Research, 48(1), 139–144. https://doi.org/10.1194/jlr.M600371-JLR200.
Rajaratnam, R. A., Gylling, H., & Miettinen, T. A. (2000). Independent association of serum squalene and noncholesterol sterols with coronary artery disease in postmenopausal women. Journal of the American College of Cardiology, 35(5), 1185–1191. https://doi.org/10.1016/s0735-1097(00)00527-1.
Ronco, A., De Stefani, E., Boffetta, P., Deneo-Pellegrini, H., Mendilaharsu, M., & Leborgne, F. (1999). Vegetables, fruits, and related nutrients and risk of breast cancer: A case-control study in Uruguay. Nutrition and Cancer, 35(2), 111–119. https://doi.org/10.1207/S15327914NC352_3.
San Mauro-Martín, I., Collado-Yurrita, L., Blumenfeld-Olivares, J. A., Cuadrado-Cenzual, M. Á., Calle-Purón, M. E., Hernández-Cabria, M., Garicano-Vilar, E., Pérez-Arruche, E., Arce-Delgado, E., & CiudadCabañas, M. J. (2016). Efecto de esteroles vegetales en la reducción del colesterol plasmático: Ensayo clínico, controlado, aleatorizado, cruzado y doble ciego. Nutricion Hospitalaria, 33(3), 279. https://doi.org/10.20960/nh.279.
Sonawane, P. D., Pollier, J., Panda, S., Szymanski, J., Massalha, H., Yona, M., Unger, T., Malitsky, S., Arendt, P., Pauwels, L., Almekias-Siegl, E., Rogachev, I., Meir, S., Cárdenas, P. D., A. A., Kamble, A., … Aharoni, A. (2016). Plant cholesterol biosynthetic pathway overlaps with phytosterol metabolism. Nature Plants, 3, 16205. https://doi.org/10.1038/nplants.2016.205.
Sundarraj, S., Thangam, R., Sreevani, V., Kaveri, K., Gunasekaran, P., Achiraman, S., & Kannan, S. (2012). γ-Sitosterol from Acacia nilotica L. induces G2/M cell cycle arrest and apoptosis through c-Myc suppression in MCF-7 and A549 cells. Journal of Ethnopharmacology, 141(3), 803–809. https://doi.org/10.1016/j.jep.2012.03.014.
Tavares, A. K. M. M., Ribas, S. A., Paravidino, V. B., Sgambato, M. R., Rodrigues, R. da R. M., da Rocha, C. M. M., Sichieri, R., & Cunha, D. B. (2021). Effect of phytosterol capsule supplementation associated with the National Cholesterol Education Program Step 2 diet on low-density lipoprotein in children and adolescents with dyslipidemia: A double-blind crossover trial. Nutrition (Burbank, Los Angeles County, Calif.), 82, 111051. https://doi.org/10.1016/j.nut.2020.111051.
Volger, O. L., Mensink, R. P., Plat, J., Hornstra, G., Havekes, L. M., & Princen, H. M. (2001). Dietary vegetable oil and wood derived plant stanol esters reduce atherosclerotic lesion size and severity in apoE*3-Leiden transgenic mice. Atherosclerosis, 157(2), 375–381. https://doi.org/10.1016/s00219150(00)00750-4.
Wang, M., Huang, W., Hu, Y., Zhang, L., Shao, Y., Wang, M., Zhang, F., Zhao, Z., Mei, X., Li, T., Wang, D., Liang, Y., Li, J., Huang, Y., Zhang, L., Xu, T., Song, H., Zhong, Y., & Lu, B. (2018). Phytosterol Profiles of Common Foods and Estimated Natural Intake of Different Structures and Forms in China. Journal of Agricultural and Food Chemistry, 66(11), 2669–2676. https://doi.org/10.1021/acs.jafc.7b05009.
Windler, E., Zyriax, B.-C., Kuipers, F., Linseisen, J., & Boeing, H. (2009). Association of plasma coronary heart disease Data from the CORA study, a case-control study of coronary artery disease in women. Atherosclerosis, 203(1), 284-290. https://doi.org/10.1016/j.atherosclerosis.2008.06.014.
Zhang, X., Lin, K., & Li, Y. (2020). Highlights to phytosterols accumulation and equilibrium in plants: Biosynthetic pathway and feedback regulation. Plant Physiology and Biochemistry: PPB, 155, 637–649. https://doi.org/10.1016/j.plaphy.2020.08.021.
Assmann, G., Cullen, P., Erbey, J., Ramey, D. R., Kannenberg, F., & Schulte, H. (2006). Plasma sitosterol elevations are associated with an increased incidence of coronary events in men: Results of a nested case-control analysis of the Prospective Cardiovascular Münster (PROCAM) study. Nutrition, Metabolism, and Cardiovascular Diseases: NMCD, 16(1), 13–21. https://doi.org/10.1016/j.numecd.2005.04.001.
Awad, A. B., Chinnam, M., Fink, C. S., & Bradford, P. G. (2007). Beta-Sitosterol activates Fas signaling in human breast cancer cells. Phytomedicine: International Journal of Phytotherapy and Phytopharmacology 14(11), 747–754. https://doi.org/10.1016/j.phymed.2007.01.003.
Bae, H., Park, S., Yang, C., Song, G., & Lim, W. (2021). Disruption of Endoplasmic Reticulum and ROS Production in Human Ovarian Cancer by Campesterol. Antioxidants (Basel, Switzerland), 10(3), 379. https://doi.org/10.3390/antiox10030379.
Brown, A. W., Hang, J., Dussault, P. H., & Carr, T. P. (2010). Plant sterol and stanol substrate specificity of pancreatic cholesterol esterase. The Journal of Nutritional Biochemistry, 21(8), 736-740. https://doi.org/10.1016/j.jnutbio.2009.04.008.
Cabianca, A., Müller, L., Pawlowski, K., & Dahlin, P. (2021). Changes in the Plant β-Sitosterol/ Stigmasterol Ratio Caused by the Plant Parasitic Nematode Meloidogyne incognita. Plants (Basel, Switzerland), 10(2), 292. https://doi.org/10.3390/plants10020292.
Clouse, S. D. (2002). Arabidopsis mutants reveal multiple roles for sterols in plant development. The Plant Cell, 14(9), 1995– 2000. https://doi.org/10.1105/tpc.140930.
Fahy, D. M., O’Callaghan, Y. C., & O’Brien, N. M. (2004). Phytosterols: Lack of cytotoxicity but interference with beta-carotene uptake in Caco-2 cells in culture. Food Additives and Contaminants, 21(1), 42–51. https://doi.org/10.1080/02652030310001636921.
Han, J.-H., Yang, Y.-X., & Feng, M.-Y. (2008). Contents of Phytosterols in Vegetables and Fruits Commonly consumed in China. Biomedical and Environmental Sciences, 21(6), 449–453. https://doi.org/10.1016/S0895-3988(09)60001-5.
Hu, M., Yang, F., Huang, Y., You, X., Liu, D., Sun, S., & Sui, S.-F. (2021). Structural insights into the mechanism of human NPC1L1mediated cholesterol uptake. Science Advances, 7(29), eabg3188. https://doi.org/10.1126/sciadv.abg3188.
Huang, W.-Q., Chen, Y.-M., & Zhang, C.-X. (2017). Association between phytosterol intake and colorectal cancer risk: A case-control study. The British Journal of Nutrition, 117(6), 839–850. https://doi.org/10.1017/S0007114517000617.
Igel, M., Giesa, U., Lutjohann, D., & von Bergmann, K. (2003). Comparison of the intestinal uptake of cholesterol, plant sterols, and stanols in mice. Journal of Lipid Research, 44(3), 533–538. https://doi.org/10.1194/jlr.M200393-JLR200.
Jaceldo-Siegl, K., Lütjohann, D., Sirirat, R., Mashchak, A., Fraser, G. E., & Haddad, E. (2017). Variations in dietary intake and plasma concentrations of plant sterols across plant-based diets among North American adults. Molecular Nutrition & Food Research, 61(8). https://doi.org/10.1002/mnfr.201600828.
Le Goff, M., Le Ferrec, E., Mayer, C., Mimouni, V., Lagadic-Gossmann, D., Schoefs, B., & Ulmann, L. (2019). Microalgal carotenoids and phytosterols regulate biochemical mechanisms involved in human health and disease prevention. Biochimie, 167, 106–118. https://doi.org/10.1016/j.biochi.2019.09.012.
Li, K., Yuan, D., Yan, R., Meng, L., Zhang, Y., & Zhu, K. (2018). Stigmasterol exhibits potent antitumor effects in human gastric cancer cells mediated via inhibition of cell migration, cell cycle arrest, mitochondrial mediated apoptosis and inhibition of JAK/STAT signalling pathway. Journal of B.U.ON.: Official Journal of the Balkan Union of Oncology, 23(5), 1420–1425.
Liu, Y., Lei, L., Wang, X., Ma, K. Y., Li, Y. M., Wang, L., Man, S. W., Huang, Y., & Chen, Z. Y. (2015). Plasma cholesterol-raising potency of dietary free cholesterol versus cholesteryl ester and effect of β-sitosterol. Food Chemistry, 169, 277–282. https://doi.org/10.1016/j.foodchem.2014.07.123.
McCann, S. E., Freudenheim, J. L., Marshall, J. R., Brasure, J. R., Swanson, M. K., & Graham, S. (2000). Diet in the epidemiology of endometrial cancer in western New York (United States). Cancer Causes & Control: CCC, 11(10), 965–974. https://doi.org/10.1023/a:1026551309873.
Mendilaharsu, M., De Stefani, E., Deneo-Pellegrini, H., Carzoglio, J., & Ronco, A. (1998). Phytosterols and risk of lung cancer: A case-control study in Uruguay. Lung Cancer (Amsterdam, Netherlands), 21(1), 37–45. https://doi.org/10.1016/s0169-5002(98)00044-0.
Moghadasian, M. H., McManus, B. M., Godin, D. V., Rodrigues, B., & Frohlich, J. J. (1999). Proatherogenic and antiatherogenic effects of probucol and phytosterols in apolipoprotein E-deficient mice: Possible mechanisms of action. Circulation, 99(13), 1733–1739. https://doi.org/10.1161/01.cir.99.13.1733.
Nguyen, T. M., Sawyer, J. K., Kelley, K. L., Davis, M. A., Kent, C. R., & Rudel, L. L. (2012). ACAT2 and ABCG5/G8 are both required for efficient cholesterol absorption in mice: Evidence from thoracic lymph duct cannulation. Journal of Lipid Research, 53(8), 1598–1609. https://doi.org/10.1194/jlr.M026823.
Oh, M.-J., So, H.-J., Hong, E.-S., Shin, J.-A., & Lee, K.-T. (2021). Presence of Cholesterol in Non-Animal Organisms: Identification and Quantification of Cholesterol in Crude Seed Oil from Perilla frutescens and Dehydrated Pyropia tenera. Molecules (Basel, Switzerland), 26(12), 3767. https://doi.org/10.3390/molecules26123767.
Othman, R. A., Myrie, S. B., & Jones, P. J. H. (2013). Non-cholesterol sterols and cholesterol metabolism in sitosterolemia. Atherosclerosis, 231(2), 291–299. https://doi.org/10.1016/j.atherosclerosis.2013.09.038.
Pereira, T. S., Fonseca, F. A. H., Fonseca, M. I. H., Martins, C. M., Fonseca, H. A. R., Fonzar, W. T., Goulart, A. C., Bensenor, I. M., Lotufo, P. A., & Izar, M. C. (2021). Phytosterol consumption and markers of subclinical atherosclerosis: Cross-sectional results from ELSA-Brasil. Nutrition, Metabolism, and Cardiovascular Diseases: NMCD, 31(6), 1756–1766. https://doi.org/10.1016/j.numecd.2021.02.031.
Pinedo, S., Vissers, M. N., von Bergmann, K., Elharchaoui, K., Lütjohann, D., Luben, R., Wareham, N. J., Kastelein, J. J. P., Khaw, K.-T., & Boekholdt, S. M. (2007). Plasma levels of plant sterols and the risk of coronary artery disease: The prospective EPIC-Norfolk Population Study. Journal of Lipid Research, 48(1), 139–144. https://doi.org/10.1194/jlr.M600371-JLR200.
Rajaratnam, R. A., Gylling, H., & Miettinen, T. A. (2000). Independent association of serum squalene and noncholesterol sterols with coronary artery disease in postmenopausal women. Journal of the American College of Cardiology, 35(5), 1185–1191. https://doi.org/10.1016/s0735-1097(00)00527-1.
Ronco, A., De Stefani, E., Boffetta, P., Deneo-Pellegrini, H., Mendilaharsu, M., & Leborgne, F. (1999). Vegetables, fruits, and related nutrients and risk of breast cancer: A case-control study in Uruguay. Nutrition and Cancer, 35(2), 111–119. https://doi.org/10.1207/S15327914NC352_3.
San Mauro-Martín, I., Collado-Yurrita, L., Blumenfeld-Olivares, J. A., Cuadrado-Cenzual, M. Á., Calle-Purón, M. E., Hernández-Cabria, M., Garicano-Vilar, E., Pérez-Arruche, E., Arce-Delgado, E., & CiudadCabañas, M. J. (2016). Efecto de esteroles vegetales en la reducción del colesterol plasmático: Ensayo clínico, controlado, aleatorizado, cruzado y doble ciego. Nutricion Hospitalaria, 33(3), 279. https://doi.org/10.20960/nh.279.
Sonawane, P. D., Pollier, J., Panda, S., Szymanski, J., Massalha, H., Yona, M., Unger, T., Malitsky, S., Arendt, P., Pauwels, L., Almekias-Siegl, E., Rogachev, I., Meir, S., Cárdenas, P. D., A. A., Kamble, A., … Aharoni, A. (2016). Plant cholesterol biosynthetic pathway overlaps with phytosterol metabolism. Nature Plants, 3, 16205. https://doi.org/10.1038/nplants.2016.205.
Sundarraj, S., Thangam, R., Sreevani, V., Kaveri, K., Gunasekaran, P., Achiraman, S., & Kannan, S. (2012). γ-Sitosterol from Acacia nilotica L. induces G2/M cell cycle arrest and apoptosis through c-Myc suppression in MCF-7 and A549 cells. Journal of Ethnopharmacology, 141(3), 803–809. https://doi.org/10.1016/j.jep.2012.03.014.
Tavares, A. K. M. M., Ribas, S. A., Paravidino, V. B., Sgambato, M. R., Rodrigues, R. da R. M., da Rocha, C. M. M., Sichieri, R., & Cunha, D. B. (2021). Effect of phytosterol capsule supplementation associated with the National Cholesterol Education Program Step 2 diet on low-density lipoprotein in children and adolescents with dyslipidemia: A double-blind crossover trial. Nutrition (Burbank, Los Angeles County, Calif.), 82, 111051. https://doi.org/10.1016/j.nut.2020.111051.
Volger, O. L., Mensink, R. P., Plat, J., Hornstra, G., Havekes, L. M., & Princen, H. M. (2001). Dietary vegetable oil and wood derived plant stanol esters reduce atherosclerotic lesion size and severity in apoE*3-Leiden transgenic mice. Atherosclerosis, 157(2), 375–381. https://doi.org/10.1016/s00219150(00)00750-4.
Wang, M., Huang, W., Hu, Y., Zhang, L., Shao, Y., Wang, M., Zhang, F., Zhao, Z., Mei, X., Li, T., Wang, D., Liang, Y., Li, J., Huang, Y., Zhang, L., Xu, T., Song, H., Zhong, Y., & Lu, B. (2018). Phytosterol Profiles of Common Foods and Estimated Natural Intake of Different Structures and Forms in China. Journal of Agricultural and Food Chemistry, 66(11), 2669–2676. https://doi.org/10.1021/acs.jafc.7b05009.
Windler, E., Zyriax, B.-C., Kuipers, F., Linseisen, J., & Boeing, H. (2009). Association of plasma coronary heart disease Data from the CORA study, a case-control study of coronary artery disease in women. Atherosclerosis, 203(1), 284-290. https://doi.org/10.1016/j.atherosclerosis.2008.06.014.
Zhang, X., Lin, K., & Li, Y. (2020). Highlights to phytosterols accumulation and equilibrium in plants: Biosynthetic pathway and feedback regulation. Plant Physiology and Biochemistry: PPB, 155, 637–649. https://doi.org/10.1016/j.plaphy.2020.08.021.