Estudio anatómico-histológico de las flores del endemismo Lamottea diania (Asteraceae) y de los efectos del ozono troposférico sobre su desarrollo.

"Estudio anatómico-histológico de las flores del endemismo Lamottea diania (Asteraceae) y de los efectos del ozono troposférico sobre su desarrollo". Lilloa 53 (2). Plantas del endemismo Lamottea diania se expusieron en cámaras OTC (Open Top Chambers) a aire ambiente filtrado y a aire ambiente no filtrado más 30 ppb de ozono para observar el efecto de este contaminante sobre el desarrollo de sus flores, particularmente sobre su androceo. Hemos comprobado, mediante estudios de microscopía óptica y electrónica, que el ozono afecta el proceso de desarrollo y maduración de los estambres y del polen. Las anteras se ven afectadas, en unos casos al abortar algunos estambres en su desarrollo y, en otros, impidiéndose el desarrollo correcto de los sacos polínicos en el interior de las mismas. Asimismo, el ozono impide la formación correcta del polen encontrando numerosos granos de polen sin desarrollar o desarrollados y madurados de forma anómala en el interior de los sacos polínicos. Los resultados indican que el ozono es el responsable del desarrollo anormal del androceo y del polen en plantas de L. diania. Anatomical and histological study of endemism flowers of Lamottea diania (Asteraceae) and the effects of tropospheric ozone on their development'. Lilloa 53 (2). Plants of endemism Lamottea diania were exposed in cameras OTC (Open Top Chambers) to filtered ambient air and ambient air unfiltered over 30 ppb ozone to observe the effect of this pollutant on the development of its flowers, particularly on its androecium. We have found, through studies of optical and electron microscopy, that ozone affects the process of development and maturation of stamens and pollen. The anthers are affected, in some cases by aborting some stamens in their development and in other, prevented the proper development of the pollen sacs within them. In addition, ozone prevents proper formationand maturation of pollen found many undeveloped pollen grains or developed abnormally inside the pollen sacs. Results indicate that ozone was responsible for the abnormal development of androecium and pollen in L. diania

Antimicrobial activity of xanthatin from Xanthium spinosum L

[EN] Dichloromethane extracts from Xanthium spinosum L. were fractionated and the fractions tested for their bactericidal and fungicidal activity. From the active fraction, a compound was isolated and identified as xanthatin (I). Xanthatin was active against Colletotrichum gloesporoides, Trichothecium roseum, Bacillus cereus and Staphylococcus aureus.Ginesta Peris, E.; García-Breijo, F.; Primo Yúfera, E. (1994). Antimicrobial activity of xanthatin from Xanthium spinosum L. Letters in Applied Microbiology. 18(4):206-208. doi:10.1111/j.1472-765X.1994.tb00848.xS20620818

Ozone-induced reductions in below-ground biomass: an anatomical approach in potato

[EN] Potato plants were grown in open-top chambers under three ozone concentrations during two complete cropping seasons (93 and 77 d in 2004 and 2005, respectively). The effects of chronic exposure to ozone on leaf anatomy, cell ultrastructure and crop yield were studied. Severe cell damage was found, even at ambient ozone levels, mainly affecting the spongy parenchyma and areas near the stomata. Damage to the cell wall caused loss of cell contact, and loss of turgor pressure due to tonoplast disintegration, contributed to cell collapse. Phloem sieve plates were obstructed by callose accumulation, and damaged mesophyll cells increased their starch stores. Tuber yield fell sharply (24–44%), due to the biggest tubers becoming smaller, which affected commercial yield. These anatomical findings show the mechanisms of ozone effect on assimilate partitioning, and thus crop yield decrease, in potato. Further implications of ozone causing reductions in belowground biomass are also discussed.The authors thank Prof Secundino del Valle (Valencia University, Spain) for his helpful comments. We are also grateful to Mr Duncan Gates for revising the English style of the text. AAF was supported by a grant from the Generalitat Valenciana's FPI programme (Government of Valencia, Spain).Asensi-Fabado, A.; García-Breijo, F.; Reig Armiñana, J. (2010). Ozone-induced reductions in below-ground biomass: an anatomical approach in potato. Plant, Cell and Environment. 33(7):1070-1083. doi:10.1111/j.1365-3040.2010.02128.xS1070108333

Development of flexible zein-wax composite and zein-fatty acid blend films for controlled release of lysozyme

The aim of this study was controlled release of lysozyme by modification of hydrophobicity and morphology of zein films using composite and blend film making methods. The incorporation of beeswax, carnauba or candelilla wax into films at 5% (w/w) of zein gave composite films containing amorphous wax particles, while incorporation of oleic acid into film at 5% of zein caused formation of blend films containing many spherical zein capsules within their matrix. The lysozyme release rates of composites reduced as the melting point of waxes increased. The composites and blends showed 2.5 to 17 fold lower lysozyme release rates than controls. The films were effectively plasticized by using catechin. The catechin also provided antioxidant activity of films (up to 69 mu mol Trolox/cm(2)) and contributed to their controlled release properties by reducing film porosity. The films showed antimicrobial activity against Listeria innocua. This work showed the possibility of obtaining advanced edible films having flexibility, antimicrobial and antioxidant activity and controlled release properties.Scientific and Technical Research Council of Turkey (108M353

Trebouxia lynnae sp. nov. (Former Trebouxia sp. TR9): Biology and Biogeography of an Epitome Lichen Symbiotic Microalga

[EN] Simple Summary In this work, we present the formal description of a new species of lichen photobiont (i.e., Trebouxia lynnae) isolated from the lichen Ramalina farinacea. The findings reported here provide an exhaustive characterization of the cellular ultrastructure, physiological traits and genetic and genomic diversity of the new species. Our results contribute to the knowledge of lichen-forming symbiotic green microalgae with their diversity and distribution. Two microalgal species, Trebouxia jamesii and Trebouxia sp. TR9, were detected as the main photobionts coexisting in the thalli of the lichen Ramalina farinacea. Trebouxia sp. TR9 emerged as a new taxon in lichen symbioses and was successfully isolated and propagated in in vitro culture and thoroughly investigated. Several years of research have confirmed the taxon Trebouxia sp. TR9 to be a model/reference organism for studying mycobiont-photobiont association patterns in lichen symbioses. Trebouxia sp. TR9 is the first symbiotic, lichen-forming microalga for which an exhaustive characterization of cellular ultrastructure, physiological traits, genetic and genomic diversity is available. The cellular ultrastructure was studied by light, electron and confocal microscopy; physiological traits were studied as responses to different abiotic stresses. The genetic diversity was previously analyzed at both the nuclear and organelle levels by using chloroplast, mitochondrial, and nuclear genome data, and a multiplicity of phylogenetic analyses were carried out to study its intraspecific diversity at a biogeographical level and its specificity association patterns with the mycobiont. Here, Trebouxia sp. TR9 is formally described by applying an integrative taxonomic approach and is presented to science as Trebouxia lynnae, in honor of Lynn Margulis, who was the primary modern proponent for the significance of symbiosis in evolution. The complete set of analyses that were carried out for its characterization is provided.PROMETEO 2021/005 (Excellence in research, Generalitat Valenciana) and the Grants New Generation EU (Ministry of Universities) to Salvador Chiva (MS21-058) and Cesar Bordenave (ZA21-046)Barreno, E.; Muggia, L.; Chiva, S.; Molins, A.; Bordenave, C.; García-Breijo, F.; Moya, P. (2022). Trebouxia lynnae sp. nov. (Former Trebouxia sp. TR9): Biology and Biogeography of an Epitome Lichen Symbiotic Microalga. Biology. 11(8):1-19. https://doi.org/10.3390/biology1108119611911

Responses of evergreen and deciduous Quercus species to enhanced ozone levels

Plants of one evergreen oak (Quercus ilex) and three deciduous oaks (Q. faginea, with small leaves; Q. pyrenaica and Q. robur, with large leaves) were exposed both to filtered air and to enhanced ozone levels in Open-Top Chambers. Q. faginea and Q. pyrenaica were studied for the first time. Based on visible injury, gas exchange, chlorophyll content and biomass responses, Q. pyrenaica was the most sensitive species, and Q. ilex was the most tolerant, followed by Q. faginea. Functional leaf traits of the species were related to differences in sensitivity, while accumulated ozone flux via stomata (POD1.6) partly contributed to the observed differences. For risk assessment of Mediterranean vegetation, the diversity of responses detected in this study should be taken into account, applying appropriate critical levels. © 2010 Elsevier Ltd. All rights reserved.We thank both the Ministerio de Medio Ambiente y Medio Rural y Maritimo (in collaboration with ICP-Forests), and the Conselleria de Medi Ambient, Aigua i Habitatge and Interreg III (ForMedOzone and VegetPollOzone projects) for supporting the OTC activity. Institut Universitario CEAM-UMH is also supported by Generalitat Valenciana and Fundacion Bancaja, benefiting from CONSOLIDER-INGENIO 2010 (GRACCIE) and Prometeo (Generalitat Valenciana) Programs. Filippo Bussotti and two anonymous referees are thanked for their useful comments. Carmen Martin is also thanked for taking care of the plants.Calatayud, V.; Cervero, J.; Calvo, E.; García Breijo, FJ.; Reig Armiñana, J.; Sanz, M. (2011). Responses of evergreen and deciduous Quercus species to enhanced ozone levels. Environmental Pollution. 159(1):55-63. doi:10.1016/j.envpol.2010.09.024S5563159

Gibberellic acid in Citrus spp. flowering and fruiting: A systematic review

[EN] Background In Citrus spp., gibberellic acid (GA) has been proposed to improve different processes related to crop cycle and yield. Accordingly, many studies have been published about how GA affects flowering and fruiting. Nevertheless, some such evidence is contradictory and the use of GA applications by farmers are still confusing and lack the expected results. Purpose This review aims to collate, present, analyze and synthesize the most relevant empirical evidence to answer the following questions: (i) how does gibberellic acid act on flowering and fruiting of citrus trees?; (ii) why is all this knowledge sometimes not correctly used by farmers to solve yield problems relating to flowering and fruit set? Methods An extensive literature search to obtain a large number of records about the topic was done. Searches were done in five databases: WoS, Scopus, Google Academics, PubMed and Scielo. The search string used was "Gibberellic acid" AND "Citrus". Records were classified into 11 groups according to the development process they referred to and initial data extraction was done. Records related with flowering and fruit set were drawn, and full texts were screened. Fifty-eight full text records were selected for the final data extraction. Results Selected studies were published from 1959 to 2017 and were published mainly in Spain, USA, Brazil and Japan. Twelve species were studied, and Citrus sinensis, C. reticulata and C. unshiu were the principal ones. Most publications with pre-flowering treatments agreed that GA decreases flowering, while only 3 out of 18 did not observe any effect. In most of these studies, the effect on fruit set and yield was not evaluated. Studies with treatments at full bloom or some weeks later mostly reported increased fruit set. However, these increases did not imply higher yields. The results on yield were highly erratic as we found increases, decreases, no effects or variable effects. Conclusions Despite some limitations, the action of GA related to cell division and growth, stimulating the sink ability of the organ and discouraging its abscission, has been clearly established through reviewed studies. GA applications before flowering counteract the floral induction caused by stress reducing flowering. However, on adult trees under field conditions, reducing flowering by applying GA would be difficult because it would be necessary to previously estimate the natural floral induction of trees. During flowering and fruit set, many problems may arise that limit production. Only when the problem is lack of fruit set stimulus can GA applications improve yields. However, much evidence suggests that the main factor-limiting yield would be carbohydrate availability rather than GA levels. GA applications increased fruit set (often transiently), but this increase did not mean improved yields.This research was supported by the Asociacion Club de Variedades Vegetales Protegidas as part of a project undertaken with the Universitat Politecnica de Valencia (Spain, UPV 20170673), of which Merle H, was the principal researcher. There was no additional external funding received for this study.Garmendia, A.; Beltrán, R.; Zornoza, C.; García-Breijo, F.; Reig, J.; Merle Farinós, HB. (2019). Gibberellic acid in Citrus spp. flowering and fruiting: A systematic review. PLoS ONE. 14(9):1-24. https://doi.org/10.1371/journal.pone.0223147S124149Kende, H., & Zeevaart, J. (1997). The Five «Classical» Plant Hormones. The Plant Cell, 1197-1210. doi:10.1105/tpc.9.7.1197Roux, S. le, & Barry, G. H. (2010). Vegetative Growth Responses of Citrus Nursery Trees to Various Growth Retardants. HortTechnology, 20(1), 197-201. doi:10.21273/horttech.20.1.197Tan, M., Song, J., & Deng, X. (2007). Production of two mandarin × trifoliate orange hybrid populations via embryo rescue with verification by SSR analysis. Euphytica, 157(1-2), 155-160. doi:10.1007/s10681-007-9407-5Greenberg, J., Holtzman, S., Fainzack, M., Egozi, Y., Giladi, B., Oren, Y., & Kaplan, I. (2010). EFFECTS OF NAA AND GA3 SPRAYS ON FRUIT SIZE AND THE INCIDENCE OF CREASING OF «WASHINGTON» NAVEL ORANGE. Acta Horticulturae, (884), 273-279. doi:10.17660/actahortic.2010.884.32Bermejo, A., Primo-Millo, E., Agustí, M., Mesejo, C., Reig, C., & Iglesias, D. J. (2015). Hormonal Profile in Ovaries of Mandarin Varieties with Differing Reproductive Behaviour. Journal of Plant Growth Regulation, 34(3), 584-594. doi:10.1007/s00344-015-9492-yDavies, F. S., & Zalman, G. (2006). Gibberellic Acid, Fruit Freezing, and Post-freeze Quality of `Hamlin’ Oranges. HortTechnology, 16(2), 301-305. doi:10.21273/horttech.16.2.0301Guardiola JL. Overview of flower bud induction, flowering and fruit set. Proceedings of Citrus Flowering and Fruit short course IFAS Citrus Research and Education Center, University of Florida. 1997. pp. 5–21.Stover, E. (2000). Relationship of Flowering Intensity and Cropping in Fruit Species. HortTechnology, 10(4), 729-732. doi:10.21273/horttech.10.4.729Stephenson, A. G. (1981). Flower and Fruit Abortion: Proximate Causes and Ultimate Functions. Annual Review of Ecology and Systematics, 12(1), 253-279. doi:10.1146/annurev.es.12.110181.001345TAKAGI, T., TOMIYASU, A., MATSUSHIMA, M., & SUZUKI, T. (1989). Seasonal Changes of GA-like Substances in Fruit and Current Shoots of Satsuma Mandarin Trees. Journal of the Japanese Society for Horticultural Science, 58(3), 569-573. doi:10.2503/jjshs.58.569Goldschmidt, E. E., Aschkenazi, N., Herzano, Y., Schaffer, A. A., & Monselise, S. P. (1985). A role for carbohydrate levels in the control of flowering in citrus. Scientia Horticulturae, 26(2), 159-166. doi:10.1016/0304-4238(85)90008-1Mahouachi, J., Iglesias, D. J., Agustí, M., & Talon, M. (2009). Delay of early fruitlet abscission by branch girdling in citrus coincides with previous increases in carbohydrate and gibberellin concentrations. Plant Growth Regulation, 58(1), 15-23. doi:10.1007/s10725-008-9348-6Gallasch, P. (1978). Attempts to control alternate cropping of Valencia orange by inhibiting flower formation with gibberellic acid. Australian Journal of Experimental Agriculture, 18(91), 309. doi:10.1071/ea9780309Martínez-Fuentes, A., Mesejo, C., Muñoz-Fambuena, N., Reig, C., González-Mas, M. C., Iglesias, D. J., … Agustí, M. (2013). Fruit load restricts the flowering promotion effect of paclobutrazol in alternate bearing Citrus spp. Scientia Horticulturae, 151, 122-127. doi:10.1016/j.scienta.2012.12.014Moss, G., & Bevington, K. (1977). The use of gibberellic acid to control alternate cropping of Late Valencia sweet orange. 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Growth and Metabolic Activity of Lemon Juice Vesicle Explants in Vitro. Plant Physiology, 69(1), 1-6. doi:10.1104/pp.69.1.1Amo-Marco, J. B. (1997). EFFECT OF PLANT GROWTH REGULATORS AND ORANGE JUICE ON GROWTH OF CALLUS FROM FRUIT TISSUES OF WASHINGTON NAVEL ORANGE. Israel Journal of Plant Sciences, 45(4), 293-296. doi:10.1080/07929978.1997.10676692Gmitter, F. G., Ling, X. B., & Deng, X. X. (1990). Induction of triploid Citrus plants from endosperm calli in vitro. Theoretical and Applied Genetics, 80(6), 785-790. doi:10.1007/bf00224192Spiegel-Roy, P., & Saad, S. (1986). Effect of carbohydrates and inhibitors of GA3 biosynthesis on embryogenenic potential of salt tolerant and non-tolerant callus lines of orange (Citrus sinensis osbeck). Plant Science, 47(3), 215-220. doi:10.1016/0168-9452(86)90181-0Shen, X., Orbović, V., Dutt, M., Castle, W. S., & Gmitter, F. G. (2013). Direct Shoot Organogenesis in Murraya paniculata (L.) Jack: A Prerequisite for Genetic Transformation. 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Regulation of Color Break in Citrus Fruits. Changes in Pigment Profiling and Gene Expression Induced by Gibberellins and Nitrate, Two Ripening Retardants. Journal of Agricultural and Food Chemistry, 54(13), 4888-4895. doi:10.1021/jf0606712Coggins C, Henning GL. Grapefruit rind blemish caused by interaction of gibberellic acid and wetting agents. International Citrus. Congress (6th: 1988: Tel Aviv, Israel): Margraf. 1988. pp. 333–338.Coggins, C. W., Scora, R. W., Lewis, L. N., & Knapp, J. C. F. (1969). Gibberellin-delayed senescence and essential oil changes in the Navel orange rind. Journal of Agricultural and Food Chemistry, 17(4), 807-809. doi:10.1021/jf60164a034El-Otmani, M., & Coggins, C. W. (1991). Growth regulator effects on retention of quality of stored citrus fruits. Scientia Horticulturae, 45(3-4), 261-272. doi:10.1016/0304-4238(91)90072-7El-Otmani, M., M’Barek, A. A., & Coggins, C. W. (1990). GA3 and 2,4-D prolong on-tree storage of citrus in Morocco. Scientia Horticulturae, 44(3-4), 241-249. doi:10.1016/0304-4238(90)90124-wFidelibus, M. W., Davies, F. S., & Campbell, C. A. (2002). Gibberellic Acid Application Timing Affects Fruit Quality of Processing Oranges. HortScience, 37(2), 353-357. doi:10.21273/hortsci.37.2.353Fidelibus, M. W., Koch, K. E., & Davies, F. S. (2008). Gibberellic Acid Alters Sucrose, Hexoses, and Their Gradients in Peel Tissues During Color Break Delay in ‘Hamlin’ Orange. Journal of the American Society for Horticultural Science, 133(6), 760-767. doi:10.21273/jashs.133.6.760Mcdonald, R. E., Greany, P. D., Shaw, P. E., & Mccollum, T. G. (1997). Preharvest applications of gibberellic acid delay senescence of Florida grapefruit. Journal of Horticultural Science, 72(3), 461-468. doi:10.1080/14620316.1997.11515534Zea-Hernández, L. O., Saucedo-Veloz, C., Cruz-Huerta, N., Ramírez-Guzmán, M. E., … Robles-González, M. M. (2016). Evaluation of post-harvest applications of gibberellic acid on the quality and shelf life of three varieties of Mexican lime. Revista Chapingo Serie Horticultura, XXII(1), 17-26. doi:10.5154/r.rchsh.2015.01.005Bower JP. The physiological control of citrus creasing. In: Albrigo LG, Sauco VG, editors. Citrus and Other Subtropical and Tropical Fruit Crops: Issues, Advances and Opportunities. 2004. pp. 111–115.Treeby, M. T., & Storey, R. (2002). Calcium-spray treatments for ameliorating albedo breakdown in navel oranges. Australian Journal of Experimental Agriculture, 42(4), 495. doi:10.1071/ea00149Li, J., Liang, C., Liu, X., Huai, B., Chen, J., Yao, Q., … Luo, X. (2016). Effect of Zn and NAA co-treatment on the occurrence of creasing fruit and the peel development of ‘Shatangju’ mandarin. Scientia Horticulturae, 201, 230-237. doi:10.1016/j.scienta.2016.01.039García-Hurtado, N., Carrera, E., Ruiz-Rivero, O., López-Gresa, M. P., Hedden, P., Gong, F., & García-Martínez, J. L. (2012). The characterization of transgenic tomato overexpressing gibberellin 20-oxidase reveals induction of parthenocarpic fruit growth, higher yield, and alteration of the gibberellin biosynthetic pathway. Journal of Experimental Botany, 63(16), 5803-5813. doi:10.1093/jxb/ers229Kojima, K., Shiozaki, K., Koshita, Y., & Ishida, M. (1999). Changes of Endogenous Levels of ABA, IAA and GA-like Substances in Fruitlets of Parthenocarpic Persimmon. Engei Gakkai zasshi, 68(2), 242-247. doi:10.2503/jjshs.68.242Salazar-Garcia, S., & Lovatt, C. J. (1999). Winter trunk injections of gibberellic acid altered the fate of `Hass’ avocado buds: Effects on inflorescence type, number and rate of development. The Journal of Horticultural Science and Biotechnology, 74(1), 69-73. doi:10.1080/14620316.1999.11511074El-Otmani, M., Lovatt, C. J., Coggins, C. W., & Agustí, M. (1995). Plant Growth Regulators in Citriculture: Factors Regulating Endogenous Levels in Citrus Tissues. 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Trebouxia lynnae sp. nov. (former Trebouxia sp. TR9): biology and biogeography of an epitome lichen symbiotic microalga

Two microalgal species, Trebouxia jamesii and Trebouxia sp. TR9, were detected as the main photobionts coexisting in the thalli of the lichen Ramalina farinacea. Trebouxia sp. TR9 emerged as anew taxon in lichen symbioses and was successfully isolated and propagated in in vitro culture andthoroughly investigated. Several years of research have confirmed the taxon Trebouxia sp. TR9 tobe a model/reference organism for studying mycobiont–photobiont association patterns in lichensymbioses. Trebouxia sp. TR9 is the first symbiotic, lichen-forming microalga for which an exhaustivecharacterization of cellular ultrastructure, physiological traits, genetic and genomic diversity is available.The cellular ultrastructure was studied by light, electron and confocal microscopy; physiologicaltraits were studied as responses to different abiotic stresses. The genetic diversity was previouslyanalyzed at both the nuclear and organelle levels by using chloroplast, mitochondrial, and nucleargenome data, and a multiplicity of phylogenetic analyses were carried out to study its intraspecificdiversity at a biogeographical level and its specificity association patterns with the mycobiont.Here, Trebouxia sp. TR9 is formally described by applying an integrative taxonomic approach and ispresented to science as Trebouxia lynnae, in honor of Lynn Margulis, who was the primary modernproponent for the significance of symbiosis in evolution. The complete set of analyses that werecarried out for its characterization is provided

Influencia de las lesiones secundarias en el ictus

Introducción: el ictus es la enfermedad neurológica más prevalente que ingresa en la unidad de cuidados intensivos, evitar las lesiones secundarias que influyen en su evolución garantiza una mayor supervivencia. Objetivo:valorar la influencia de las lesiones secundarias en la evolución de  los pacientes con ictus mayores de 18 años ingresados en la unidad de cuidados intensivos 3 del Hospital General Docente Abel Santamaría Cuadrado de Pinar del Río entre septiembre 2014 a agosto 2015. Método: se realizó un estudio descriptivo, retrospectivo,  longitudinal de corte transversal, utilizándose las variables: etiología, mortalidad, sexo, color de la piel, factores de riesgo, lesiones secundarias y estadía hospitalaria. Para el procesamiento estadístico se usó Chi cuadrado, Odds Ratio, frecuencia absoluta y la media. El universo estuvo constituido por 70 pacientes y la muestra fueron 38 que cumplieron crietrios pre establecido de inclusión en el estudio. Resultados: la incidencia fue baja  con 14,86%. La forma isquémica predominó y la mortalidad fue mayor para la  hemorrágicacon  76,47%,  el sexo femenino yel  color de la piel constituyó un factor para padecer enfermedad cerebro vascular siendoesta última estadísticamente significativa. La hipertensión arterial, tabaquismo, cardiopatía isquémica y diabetes mellitus como  factores de riesgo y la hipertensión arterial, hipertermia, hipoxemia, edema cerebral y convulsiones como lesiones secundarias, fueron los más reportados. El 50% tenían entre 2-5  lesiones secundarias, con mayor mortalidad los que más presentaban, siendo la primera semana donde más se registraba. Conclusiones: para seguir una  conducta adecuada con los pacientes ingresados con enfermedad cerebro vasculary poder reducir la estadía y complicaciones,  se debe evitar estrictamente las lesiones secundarias, que empeoran la evolución y enlentecen la pronta recuperación