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In the periphery, hypoxia is a powerful regulator of angiogenesis. However, vascular endothelial cells are remarkably heterogeneous and little is known about how brain endothelial cells respond to hypoxic challenge. The objective of this study is to characterize the effect of hypoxic challenge on the angiogenic response of cultured brainderived microvascular endothelial cells. Brain endothelial cell cultures were initiated from isolated rat brain microvessels and subjected to hypoxia (1% O 2) for various time periods. The results showed that hypoxia induced rapid (≤ 0.5 h) expression of hypoxia-inducible factor 1α (HIF-1α) and that cell viability, assessed by MTT assay, was unaffected within the first 8 h. Examination of brain endothelial cell cultures for pro-and anti-angiogenic proteins by western blot, RT-PCR and ELISA revealed that within 0.5 to 2 h of hypoxia levels of vascular endothelial growth factor and endothelin-1 mRNA and protein were elevated. The expression of heme oxygenase-1 also increased but only after 8 h of hypoxia. In contrast, similar hypoxia exposure evoked a decrease in endothelial nitric oxide synthase and thrombospondin-2 levels. Exposure of brain endothelial cell cultures to hypoxia resulted in a significant (p\u003c0.001) decrease (94%) in tube length, an in vitro index of angiogenesis, compared to control cultures. The data indicate, despite a shift toward a pro-angiogenic phenotype, hypoxia inhibited vessel formation in brain endothelial cells. These results suggest that in brain endothelial cells expression of angiogenic factors is not sufficient for the development of new vessels. Further work is needed to determine what factors/conditions prevent hypoxia-induced angiogenic changes from culminating in the formation of new brain blood vessels and what role this may play in the pathologic changes observed in AD and other diseases characterized by cerebral hypoxia.","publication_date":"2012,,","publication_name":"Microvascular Research","grobid_abstract_attachment_id":"101609578"},"document_type":"paper","pre_hit_view_count_baseline":null,"quality":"high","language":"en","title":"Hypoxia induces angiogenic factors in brain microvascular endothelial cells","broadcastable":false,"draft":null,"has_indexable_attachment":true,"indexable":true}}["work"]; window.loswp.workCoauthors = [267578687]; window.loswp.locale = "en"; window.loswp.countryCode = "SG"; window.loswp.cwvAbTestBucket = ""; window.loswp.designVariant = "ds_vanilla"; window.loswp.fullPageMobileSutdModalVariant = "full_page_mobile_sutd_modal"; window.loswp.useOptimizedScribd4genScript = false; 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data-login_uri="https://www.academia.edu/registrations/google_one_tap" data-moment_callback="onGoogleOneTapEvent" id="g_id_onload"></div><div class="ds-top-related-works--grid-container"><div class="ds-related-content--container ds-top-related-works--container"><h2 class="ds-related-content--heading">Related papers</h2><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="0" data-entity-id="80857526" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/80857526/Brain_microvasculature_and_hypoxia_related_proteins_in_Alzheimers_disease">Brain microvasculature and hypoxia-related proteins in Alzheimer's disease</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="47845662" href="https://independent.academia.edu/AlmaSanchezB">Alma Sanchez B</a></div><p class="ds-related-work--metadata ds2-5-body-xs">International journal of clinical and experimental pathology, 2011</p><p class="ds-related-work--abstract ds2-5-body-sm">Alzheimer&#39;s disease (AD) is a progressive, neurodegenerative disease of increasing incidence. 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The objectives of this study were to determine expression of HIF-1α, angiogenic proteins, angiopoietin-2 (Ang-2), and matrix metalloproteinase 2 (MMP2), and survival/apoptotic proteins (Bcl-xL, cas...</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Brain microvasculature and hypoxia-related proteins in Alzheimer's disease","attachmentId":87099253,"attachmentType":"pdf","work_url":"https://www.academia.edu/80857526/Brain_microvasculature_and_hypoxia_related_proteins_in_Alzheimers_disease","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/80857526/Brain_microvasculature_and_hypoxia_related_proteins_in_Alzheimers_disease"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="1" data-entity-id="100641919" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/100641919/Effect_of_Hypoxia_on_the_Proliferation_of_Microvascular_Cells_in_Vitro">Effect of 'Hypoxia' on the Proliferation of Microvascular Cells in Vitro</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="173435165" href="https://independent.academia.edu/PatrickMoriarty13">Patrick Moriarty</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Experimental Eye Research, 1994</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Effect of 'Hypoxia' on the Proliferation of Microvascular Cells in Vitro","attachmentId":101406682,"attachmentType":"pdf","work_url":"https://www.academia.edu/100641919/Effect_of_Hypoxia_on_the_Proliferation_of_Microvascular_Cells_in_Vitro","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/100641919/Effect_of_Hypoxia_on_the_Proliferation_of_Microvascular_Cells_in_Vitro"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="2" data-entity-id="5450221" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/5450221/Angiogenesis_as_a_predictive_marker_of_neurological_outcome_following_hypoxia_ischemia">Angiogenesis as a predictive marker of neurological outcome following hypoxia–ischemia</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="7640545" href="https://otago.academia.edu/IvanSammut">Ivan Sammut</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Brain Research, 2007</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Angiogenesis as a predictive marker of neurological outcome following hypoxia–ischemia","attachmentId":49288157,"attachmentType":"pdf","work_url":"https://www.academia.edu/5450221/Angiogenesis_as_a_predictive_marker_of_neurological_outcome_following_hypoxia_ischemia","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/5450221/Angiogenesis_as_a_predictive_marker_of_neurological_outcome_following_hypoxia_ischemia"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="3" data-entity-id="28879735" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/28879735/Cerebral_angiogenic_factors_angiogenesis_and_physiological_response_to_chronic_hypoxia_differ_among_four_commonly_used_mouse_strains">Cerebral angiogenic factors, angiogenesis, and physiological response to chronic hypoxia differ among four commonly used mouse strains</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="54384473" href="https://independent.academia.edu/NicoleWard10">Nicole Ward</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Applied Physiology, 2007</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Cerebral angiogenic factors, angiogenesis, and physiological response to chronic hypoxia differ among four commonly used mouse strains","attachmentId":49310394,"attachmentType":"pdf","work_url":"https://www.academia.edu/28879735/Cerebral_angiogenic_factors_angiogenesis_and_physiological_response_to_chronic_hypoxia_differ_among_four_commonly_used_mouse_strains","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/28879735/Cerebral_angiogenic_factors_angiogenesis_and_physiological_response_to_chronic_hypoxia_differ_among_four_commonly_used_mouse_strains"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="4" data-entity-id="98562700" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/98562700/Effects_of_Hypoxia_Reoxygenation_on_Microvascular_Endothelial_Function_in_the_Rat_Hippocampal_Slice_">Effects of Hypoxia–Reoxygenation on Microvascular Endothelial Function in the Rat Hippocampal Slice </a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32291038" href="https://independent.academia.edu/CathyDrexler">Cathy Drexler</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Anesthesiology, 1999</p><p class="ds-related-work--abstract ds2-5-body-sm">Background Cerebral ischemia and hypoxia may cause injury to both neuronal and vascular tissue. The direct effects of hypoxia on endothelial function in intraparenchymal cerebral arterioles are unknown. Using a modification of the rat brain slice preparation, allowing continuous imaging of these previously inaccessible vessels, microvessel dilation was evaluated before and after a brief hypoxic episode. Methods Rat brain slices were superfused with oxygenated artificial cerebrospinal fluid. Hippocampal arterioles were visualized using computerized videomicroscopy, and their diameters (range, 12-27 microm) were measured using image analysis. After preconstriction with prostaglandin F2alpha and controlled pH and carbon dioxide tension, graded concentrations of either acetylcholine (endothelium-dependent vasodilation) or sodium nitroprusside (endothelium-independent vasodilation) were given before and after a 10-min period of hypoxia. Results Sodium nitroprusside (100 microM) caused si...</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Effects of Hypoxia–Reoxygenation on Microvascular Endothelial Function in the Rat Hippocampal Slice ","attachmentId":99879141,"attachmentType":"pdf","work_url":"https://www.academia.edu/98562700/Effects_of_Hypoxia_Reoxygenation_on_Microvascular_Endothelial_Function_in_the_Rat_Hippocampal_Slice_","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/98562700/Effects_of_Hypoxia_Reoxygenation_on_Microvascular_Endothelial_Function_in_the_Rat_Hippocampal_Slice_"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="5" data-entity-id="102145155" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/102145155/Hypoxia_induced_adaptation_of_cerebral_microvasculature">Hypoxia-induced adaptation of cerebral microvasculature</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="34797888" href="https://independent.academia.edu/IwaoKanno">Iwao Kanno</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Microvascular Reviews and Communications, 2013</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" 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js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/17870707/Hypoxia_inducible_factor_1_HIF_1_independent_microvascular_angiogenesis_in_the_aged_rat_brain">Hypoxia-inducible factor-1 (HIF-1)-independent microvascular angiogenesis in the aged rat brain</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32868221" href="https://case.academia.edu/JosephLamanna">Joseph C LaManna</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Brain Research, 2010</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Hypoxia-inducible factor-1 (HIF-1)-independent microvascular angiogenesis in the aged rat brain","attachmentId":39754680,"attachmentType":"pdf","work_url":"https://www.academia.edu/17870707/Hypoxia_inducible_factor_1_HIF_1_independent_microvascular_angiogenesis_in_the_aged_rat_brain","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/17870707/Hypoxia_inducible_factor_1_HIF_1_independent_microvascular_angiogenesis_in_the_aged_rat_brain"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="7" data-entity-id="76834103" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" 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href="https://independent.academia.edu/kathiravangovindarajan">kathiravan govindarajan</a></div><p class="ds-related-work--metadata ds2-5-body-xs">… journal of oncology, 2007</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Glioma cells suppress hypoxia-induced endothelial cell apoptosis and promote angiogenic process","attachmentId":72840385,"attachmentType":"pdf","work_url":"https://www.academia.edu/58417889/Glioma_cells_suppress_hypoxia_induced_endothelial_cell_apoptosis_and_promote_angiogenic_process","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" 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Castillo-melendez</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Pediatric Research, 2012</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"VEGF expression and microvascular responses to severe transient hypoxia in the fetal sheep brain","attachmentId":42061105,"attachmentType":"pdf","work_url":"https://www.academia.edu/18127677/VEGF_expression_and_microvascular_responses_to_severe_transient_hypoxia_in_the_fetal_sheep_brain","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" 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ds2-5-body-link" data-author-id="187206366" href="https://independent.academia.edu/JoAnnGensert">JoAnn Gensert</a></div><p class="ds-related-work--metadata ds2-5-body-xs">2010</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Hypoxia Inducible Factor1 as Novel Targets for Stroke Therapy Translation of Ischemic Preconditioning to the Patient: Prolyl Hydroxylase Inhibition and","attachmentId":89848706,"attachmentType":"pdf","work_url":"https://www.academia.edu/85018256/Hypoxia_Inducible_Factor1_as_Novel_Targets_for_Stroke_Therapy_Translation_of_Ischemic_Preconditioning_to_the_Patient_Prolyl_Hydroxylase_Inhibition_and","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link 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activation, proliferation, and increased expression of the α6β4 integrin and dystroglycan</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32868221" href="https://case.academia.edu/JosephLamanna">Joseph C LaManna</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Glia, 2010</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"In the hypoxic central nervous system, endothelial cell proliferation is followed by astrocyte activation, proliferation, and increased expression of the α6β4 integrin and 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class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="13" data-entity-id="14522623" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/14522623/Effects_of_hypoxia_on_endothelial_pericytic_co_culture_model_of_the_blood_brain_barrier">Effects of hypoxia on endothelial/pericytic co-culture model of the blood–brain barrier</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="33477131" href="https://manoa-hawaii.academia.edu/KentaroHayashi">Kentaro Hayashi</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Regulatory Peptides, 2004</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Effects of hypoxia on endothelial/pericytic 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class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Cerebral adaptations to chronic anemia in a model of erythropoietin-deficient mice exposed to hypoxia","attachmentId":47217655,"attachmentType":"pdf","work_url":"https://www.academia.edu/10671470/Cerebral_adaptations_to_chronic_anemia_in_a_model_of_erythropoietin_deficient_mice_exposed_to_hypoxia","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/10671470/Cerebral_adaptations_to_chronic_anemia_in_a_model_of_erythropoietin_deficient_mice_exposed_to_hypoxia"><span class="ds2-5-text-link__content">View PDF</span><span 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href="https://www.academia.edu/55319197/Differential_activation_of_vascular_genes_by_hypoxia_in_primary_endothelial_cells">Differential activation of vascular genes by hypoxia in primary endothelial cells</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="47887892" href="https://independent.academia.edu/StefanWennstrom">Stefan Wennström</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Experimental Cell Research, 2004</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Differential activation of vascular genes by hypoxia in primary endothelial 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ds2-5-body-link" href="https://www.academia.edu/76248219/Thrombin_a_mediator_of_cerebrovascular_inflammation_in_AD_and_hypoxia">Thrombin, a mediator of cerebrovascular inflammation in AD and hypoxia</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="215567674" href="https://independent.academia.edu/JosephMartinez143">Joseph Martinez</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Frontiers in aging neuroscience, 2013</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Thrombin, a mediator of cerebrovascular inflammation in AD and hypoxia","attachmentId":83977804,"attachmentType":"pdf","work_url":"https://www.academia.edu/76248219/Thrombin_a_mediator_of_cerebrovascular_inflammation_in_AD_and_hypoxia","alternativeTracking":true}"><span 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microvascular endothelium during long-term hypoxia: Alterations in cell growth, monolayer permeability, and cell surface coagulant properties</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32024011" href="https://independent.academia.edu/FedericoCozzolino">Federico Cozzolino</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Journal of Cellular Physiology, 1991</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Macrovascular and microvascular endothelium during long-term hypoxia: Alterations in cell growth, monolayer permeability, and cell surface coagulant 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