On the Formation of Lipid Droplets in Human Adipocytes: The Organization of the Perilipin–Vimentin Cortex
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{"title"=>"On the formation of lipid droplets in human adipocytes: The organization of the perilipin-vimentin cortex", "type"=>"journal", "authors"=>[{"first_name"=>"Hans", "last_name"=>"Heid", "scopus_author_id"=>"7003796695"}, {"first_name"=>"Steffen", "last_name"=>"Rickelt", "scopus_author_id"=>"18635041800"}, {"first_name"=>"Ralf", "last_name"=>"Zimbelmann", "scopus_author_id"=>"6701756475"}, {"first_name"=>"Stefanie", "last_name"=>"Winter", "scopus_author_id"=>"7202247340"}, {"first_name"=>"Heiderose", "last_name"=>"Schumacher", "scopus_author_id"=>"15840486700"}, {"first_name"=>"Yvette", "last_name"=>"Dörflinger", "scopus_author_id"=>"55165382400"}, {"first_name"=>"Caecilia", "last_name"=>"Kuhn", "scopus_author_id"=>"7202056585"}, {"first_name"=>"Werner W.", "last_name"=>"Franke", "scopus_author_id"=>"35473669800"}], "year"=>2014, "source"=>"PLoS ONE", "identifiers"=>{"scopus"=>"2-s2.0-84896538646", "doi"=>"10.1371/journal.pone.0090386", "sgr"=>"84896538646", "isbn"=>"1932-6203 (Electronic)\\r1932-6203 (Linking)", "pmid"=>"24587346", "issn"=>"19326203", "pui"=>"372643332"}, "id"=>"34185270-56a7-3ecf-a7c8-ca4999d80952", "abstract"=>"We report on the heterogeneity and diversity of lipid droplets (LDs) in early stages of adipogenesis by elucidating the cell and molecular biology of amphiphilic and cytoskeletal proteins regulating and stabilizing the generation of LDs in human adipose cells. A plethora of distinct and differently sized LDs was detected by a brief application of adipocyte differentiation medium and additional short treatment with oleic acid. Using these cells and highly specific antibodies for LD-binding proteins of the perilipin (PLIN) family, we could distinguish between endogenously derived LDs (endogenous LDs) positive for perilipin from exogenously induced LDs (exogenous LDs) positive for adipophilin, TIP47 and S3-12. Having optimized these stimulation conditions, we used early adipogenic differentiation stages to investigate small-sized LDs and concentrated on LD-protein associations with the intermediate-sized filament (IF) vimentin. This IF protein was described earlier to surround lipid globules, showing spherical, cage-like structures. Consequently - by biochemical methods, by immunofluorescence microscopy and by electron- and immunoelectron microscopy - various stages of emerging lipid globules were revealed with perilipin as linking protein between LDs and vimentin. For this LD-PLIN-Vimentin connection, a model is now proposed, suggesting an interaction of proteins via opposed charged amino acid domains respectively. In addition, multiple sheaths of smooth endoplasmic reticulum cisternae surrounding concentrically nascent LDs are shown. Based on our comprehensive localization studies we present and discuss a novel pathway for the LD formation.", "link"=>"http://www.mendeley.com/research/formation-lipid-droplets-human-adipocytes-organization-perilipinvimentin-cortex", "reader_count"=>48, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>2, "Researcher"=>5, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>21, "Student > Postgraduate"=>2, "Student > Master"=>6, "Other"=>1, "Student > Bachelor"=>4, "Professor"=>4}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>2, "Researcher"=>5, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>21, "Student > Postgraduate"=>2, "Student > Master"=>6, "Other"=>1, "Student > Bachelor"=>4, "Professor"=>4}, "reader_count_by_subject_area"=>{"Engineering"=>1, "Biochemistry, Genetics and Molecular Biology"=>12, "Agricultural and Biological Sciences"=>22, "Medicine and Dentistry"=>9, "Neuroscience"=>1, "Sports and Recreations"=>1, "Chemistry"=>2}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>9}, "Neuroscience"=>{"Neuroscience"=>1}, "Chemistry"=>{"Chemistry"=>2}, "Sports and Recreations"=>{"Sports and Recreations"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>22}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>12}}, "reader_count_by_country"=>{"Argentina"=>1}, "group_count"=>2}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/1404296"], "description"=>"<p>(a, a, a) AIM treatment for 1–3 days reveals many newly generated small and medium-sized LDs positively staining with perilipin antibodies (red). Additional OA-treatment for 2 hours leads to the appearance of LDs positive for adipophilin (green) in higher numbers and bigger sizes when compared to non-stimulated or AIM-stimulated cells solely (cp. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone-0090386-g001\" target=\"_blank\">Fig. 1</a>). (a) Enlarged part taken from double staining seen in (a). (a) Enlarged part from (a) showing single color and mosaic double-stained droplets, obviously in the midst of a fusion process with involvement of different types of LD-binding PLIN proteins and with yellow, partly mixed colored LDs for co-localization. (b) A different perilipin/adipophilin antibody combination reveals a similar heterogeneous staining of LDs as seen in a-a. (c) Perilipin (red) vs. S3-12 (green) double staining. Several smaller S3-12-positive droplets are seen attaching and combining with larger perilipin-positive droplets. Many newly generated, very small and middle-sized rings of S3-12-positive droplet staining with punctate patterns can be seen by single green color staining only (examples marked by arrowheads). (d) Perilipin (red) vs. TIP47 (green) double staining. Some of the many newly appearing TIP47-positive small droplets adhere occasionally to the much bigger perilipin positive droplets. (e) Adipophilin (red) vs. TIP47 (green) double staining. Most of the droplets are visible with sizes estimated smaller than 1 µm in diameter. Some heterogeneously colored droplets are marked by arrowheads. Note, OA-treatment leads to the re-appearance of a tremendous number of small droplets positive for adipophilin, TIP47 and S3-12. Note in addition, the complexity of the PLIN staining of LDs. Different LD-binding proteins are expressed and different types of LDs appear in single cells by specific “endogenous” and “exogenous” stimulations. DAPI (blue). Bar in a: 5 µm; all other Bars: 20 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "scanning", "microscopy", "ld", "labeling", "perilipin", "aim-stimulated", "preadipocytes", "additionally", "treated", "oleic"], "article_id"=>949280, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g002", "stats"=>{"downloads"=>2, "page_views"=>36, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Laser_scanning_microscopy_showing_LD_labeling_of_perilipin_proteins_in_briefly_AIM_stimulated_human_preadipocytes_which_were_additionally_treated_shortly_with_oleic_acid_OA_/949280", "title"=>"Laser scanning microscopy showing LD labeling of perilipin proteins in briefly AIM-stimulated human preadipocytes which were additionally treated shortly with oleic acid (OA).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404309"], "description"=>"<p>(a) Short AIM stimulation and simultaneous fixation method, often lead to the appearance of small LDs (approximately 1 µm in diameter) which are completely surrounded by multiple cisternae and layers of smooth ER (white double arrows). Smaller LDs of 0.2– 0.5 µm in diameter without such ER layers can also be seen (right side). Embedded in a meshwork of filaments (black arrows) are mitochondria (M) in the neighborhood of these ER layers with no direct contact to LDs. (b) Addition of OA to the cell media for one day leads to the decay of the droplet surrounding ER layers (here shown with sequential fixation method). Arrowheads mark periodically arranged vimentin cage-like structures seen directly in contact with the LD surface. These regular arrays of anchored IFs have been described already by Franke et al. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Franke1\" target=\"_blank\">[15]</a>. Several IF bundles within the cytoplasma and with connections to LDs are marked by arrows. Note, with both treatments and fixation methods, mitochondria (M) are not seen in direct contact to the LDs. Bars: 0.50 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "micrographs", "comparing", "aim-stimulated", "oa-treated"], "article_id"=>949293, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g010", "stats"=>{"downloads"=>6, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Electron_micrographs_comparing_briefly_AIM_stimulated_versus_briefly_AIM_stimulated_plus_OA_treated_human_preadipocytes_/949293", "title"=>"Electron micrographs comparing briefly AIM-stimulated versus briefly AIM-stimulated plus OA-treated human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404294"], "description"=>"<p>(a-c) Non-stimulated cells. (a) Adipophilin monoclonal antibody (mab) staining reveals many small LDs (green). (b) Positive TIP47 polyclonal antibody (pab; green) vs. negative perilipin (mab; red) staining. (c) Pab S3-12 shows many very small LDs – some can be seen like arranged and placed along rows of filaments. (d-g) Adipocyte differentiation of cells with AIM medium. 1-3 days of AIM treatment newly generates many small and medium-sized LDs, which stain with antibodies for perilipin (red). In contrast after AIM treatment, LDs positive for other perilipin (PLIN) family members are reduced in numbers and size (green). Cells still appear fibroblast-like elongated, not roundish. (d) Perilipin vs. adipophilin staining. (e) Perilipin and TIP47 double staining. (f) Perilipin localization at surface of LDs (g) Perilipin comparison with S3-12. Note, whereas antibodies specific for adipophilin, S3-12 and TIP47 show plenty of small LDs in non-stimulated cells and staining for perilipin is negative, the situation changes completely with the start of AIM stimulation. (For additional examples of small LD staining conspicuously arranged along rows of filaments, see e.g. TIP47 staining patterns obtained with PLC epithelial cells given in Fig. S5 by Heid et al., 2013 <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Heid1\" target=\"_blank\">[13]</a>). Nuclear staining was with DAPI (blue). Bars: 20 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "scanning", "single-", "double-label", "microscopy", "lipid", "droplet", "labeling", "perilipin", "non-stimulated", "aim-stimulated"], "article_id"=>949278, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g001", "stats"=>{"downloads"=>1, "page_views"=>39, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Laser_scanning_single_and_double_label_immunofluorescence_microscopy_showing_lipid_droplet_LD_labeling_of_perilipin_proteins_in_untreated_non_stimulated_and_in_briefly_AIM_stimulated_human_preadipocytes_/949278", "title"=>"Laser scanning single- and double-label immunofluorescence microscopy showing lipid droplet (LD) labeling of perilipin proteins in untreated, non-stimulated and in briefly AIM-stimulated human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404304"], "description"=>"<p>(a) Pab Peri-hCT and mab Vim 3B4 reveal similar staining as seen with alike antibodies (cp. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone-0090386-g006\" target=\"_blank\">Fig. 6a</a>). (b) At closer inspection, pab Peri-hCT often shows additional localization. Fine punctated rows of very small droplets can be seen on perinuclear distinct winded structures (perilipin staining pattern P1). (c) Enlarged perinuclear area of a cell starting adipocyte conversion. Many, tiny lipid droplets in the cytoplasm, can be seen especially at perinuclear structures (pattern P1) and in close association with the vimentin IF network. (d,d,d) Different stages of adipocyte conversion are present within the two neighboring cells shown. In the cell in the lower part of the pictures, conversion appears to begin, showing 2 different perilipin patterns - many tiny droplets binding in rows on winded perinuclear structures (pattern P1; cp. b,c) and plenty of small droplets with sizes estimated below 1 µm in diameter and distributed mainly above the perinuclear region in the cytoplasm (pattern P2). In the advanced conversion stage of the cell shown in the upper part of the picture, fairly well accomplished, bigger LDs are seen distributed all over the cytoplasm and surrounded by vimentin cages (pattern P3, examples of vimentin cages shown by arrowheads). Note, by increasing the time for AIM stimulation, we noticed an increased P3 pattern paralleled by decreasing P1 and P2 patterns. By conventionally longer AIM treatment (1–3 weeks), the P3 pattern could be seen almost exclusively in all differentiated cells. DAPI (blue). Bars in a-c: 20 µm. Bar in d: 10 µm;</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "scanning", "microscopy", "vimentin", "c-terminal", "perilipin", "antibodies", "aim-stimulated", "preadipocytes", "detection"], "article_id"=>949288, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g007", "stats"=>{"downloads"=>0, "page_views"=>33, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Laser_scanning_microscopy_showing_comparison_of_vimentin_and_C_terminal_specific_perilipin_antibodies_using_briefly_AIM_stimulated_human_preadipocytes_and_detection_of_different_perilipin_staining_patterns_/949288", "title"=>"Laser scanning microscopy showing comparison of vimentin and C-terminal specific perilipin antibodies using briefly AIM-stimulated human preadipocytes and detection of different perilipin staining-patterns.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404302"], "description"=>"<p>(a) Mab Vim3B4 reveals a dense filamentous network with additional prominent ring- and cage-like structures (red staining; some examples of cages are marked by arrowheads). (a) Double-staining of mab Vim3B4 (red) with pab Peri-hNT (green) unambiguously shows that these cages of intermediate-sized filaments (IFs) are marked by perilipin staining. DAPI (blue). Bar: 20 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "scanning", "microscopy", "vimentin", "n-terminal", "perilipin", "antibodies", "aim-stimulated"], "article_id"=>949286, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g006", "stats"=>{"downloads"=>3, "page_views"=>18, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Laser_scanning_microscopy_showing_comparison_of_vimentin_and_N_terminal_specific_perilipin_antibodies_using_briefly_AIM_stimulated_human_preadipocytes_/949286", "title"=>"Laser scanning microscopy showing comparison of vimentin and N-terminal specific perilipin antibodies using briefly AIM-stimulated human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404299"], "description"=>"<p>(a) Fractions obtained by cell disintegration using nitrogen cavitation, iodixanol gradient centrifugation followed by SDS-PAGE separations and Coomassie blue staining are shown (cp. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Heid1\" target=\"_blank\">[13]</a>). Separated fractions were tested with Western blotting (WB) using pabs TIP47-hCT(b), S3-12-hNT (c,c), Adipo-hCT (d,d) and mabs Peri112.17 (e,é) and Vim3B4 (f,f´). Positions of molecular weight markers are given on the left margin of (a) and fraction numbers on top of (a,b). (c,d,é,f´) represent prolonged exposures of reactions shown in (c,d,e,f) respectively. The higher molecular band reactions of adipophilin and perilipin seen in LD1 fractions are unknown modifications of PLIN proteins (asterisks in d,e). Note, the separation shown here is given with AIM-stimulated preadipocytes and not with AIM-stimulated <u>plus</u> OA-treated preadipocytes. Therefore only the few small (i.e. freshly endocytosed) LDs are detected. These are mostly coalesced with the bigger endogenously generated LDs and therefore preferentially found here within the LD1 fraction. Importantly PLIN proteins can be detected also in these non OA-treated preadipocytes together with vimentin within <u>all</u> three major LD gradient areas LD1, LD2, LD3 - including the top layer LD1.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "vimentin", "gradient", "centrifugation", "fractions", "aim-stimulated"], "article_id"=>949283, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g004", "stats"=>{"downloads"=>1, "page_views"=>19, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_PLIN_proteins_and_vimentin_detected_in_different_density_gradient_centrifugation_fractions_of_briefly_AIM_stimulated_preadipocytes_/949283", "title"=>"PLIN proteins and vimentin detected in different density gradient centrifugation fractions of briefly AIM-stimulated preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404314"], "description"=>"<p>(<b>a</b>) Shown are groups of LDs positive for perilipin by nanogold-label and silver enhancement. Localization was with mab Peri112.17. The LDs are seen closely associated and anchored with IF bundles. (<b>b</b>) Enlarged perilipin labeling with two small LDs (approximately 300–400 nm in diameter) approaching a big LD (approximately 2.0–2.5 µm in diameter) for combining and coalescence. Bundles of intermediate-sized filaments are associated to grains of the immunolabeled perilipin (arrows). Bars: 0.50 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "microscopic", "localization", "perilipin", "aim-stimulated"], "article_id"=>949297, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g012", "stats"=>{"downloads"=>0, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Immunoelectron_microscopic_localization_of_perilipin_in_briefly_AIM_stimulated_human_preadipocytes_/949297", "title"=>"<i>Immunoelectron microscopic localization of perilipin in briefly AIM-stimulated human preadipocytes.</i>", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404311"], "description"=>"<p>(<b>a</b>) Small and characteristic LDs can be found by short-time adipose conversion showing droplets with several flat layers and cisternae of smooth endoplasmic reticulum (LD<sub>ER</sub>, white double arrows; simultaneous fixation method). Occasionally, very regularly spaced dots of vimentin filaments can be seen at the surface of such droplets sandwiched between droplets and ER (black arrowheads). In addition, smaller LDs of 0.2– 0.5 µm in diameter without such surrounding ER layers - but with interaction of vimentin IFs (marked by black arrows; left side) – and larger, fully accomplished LDs of several µm in diameter with mitochondria (M) in the neighborhood can be seen. (<b>b</b>) The regular pattern of vimentin arrays at the surface of LDs can be recognized best where fewer and less densely packed ER sheaths surround the droplets (black arrowheads), which might constitute a specific stage of adipogenesis. In this situation LDs seem to be matured, i.e. grown enough in size and the multiple layers of ER cisternae are just about to be released. Note, such EM pictures suggest that different stages and types of LDs in differentiating adipocytes can be pictured and characterized. Bars: 0.50 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "micrographs", "associations", "lds", "er", "aim-stimulated"], "article_id"=>949295, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g011", "stats"=>{"downloads"=>1, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Electron_micrographs_showing_special_associations_of_LDs_and_ER_within_briefly_AIM_stimulated_human_preadipocytes_/949295", "title"=>"Electron micrographs showing special associations of LDs and ER within briefly AIM-stimulated human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404340", "https://ndownloader.figshare.com/files/1404341", "https://ndownloader.figshare.com/files/1404343", "https://ndownloader.figshare.com/files/1404344", "https://ndownloader.figshare.com/files/1404345", "https://ndownloader.figshare.com/files/1404346"], "description"=>"<div><p>We report on the heterogeneity and diversity of lipid droplets (LDs) in early stages of adipogenesis by elucidating the cell and molecular biology of amphiphilic and cytoskeletal proteins regulating and stabilizing the generation of LDs in human adipose cells. A plethora of distinct and differently sized LDs was detected by a brief application of adipocyte differentiation medium and additional short treatment with oleic acid. Using these cells and highly specific antibodies for LD-binding proteins of the perilipin (PLIN) family, we could distinguish between endogenously derived LDs (endogenous LDs) positive for perilipin from exogenously induced LDs (exogenous LDs) positive for adipophilin, TIP47 and S3-12. Having optimized these stimulation conditions, we used early adipogenic differentiation stages to investigate small-sized LDs and concentrated on LD-protein associations with the intermediate-sized filament (IF) vimentin. This IF protein was described earlier to surround lipid globules, showing spherical, cage-like structures. Consequently - by biochemical methods, by immunofluorescence microscopy and by electron- and immunoelectron microscopy - various stages of emerging lipid globules were revealed with perilipin as linking protein between LDs and vimentin. For this LD-PLIN-Vimentin connection, a model is now proposed, suggesting an interaction of proteins via opposed charged amino acid domains respectively. In addition, multiple sheaths of smooth endoplasmic reticulum cisternae surrounding concentrically nascent LDs are shown. Based on our comprehensive localization studies we present and discuss a novel pathway for the LD formation.</p></div>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "lipid", "droplets"], "article_id"=>949320, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0090386.s001", "https://dx.doi.org/10.1371/journal.pone.0090386.s002", "https://dx.doi.org/10.1371/journal.pone.0090386.s003", "https://dx.doi.org/10.1371/journal.pone.0090386.s004", "https://dx.doi.org/10.1371/journal.pone.0090386.s005", "https://dx.doi.org/10.1371/journal.pone.0090386.s006"], "stats"=>{"downloads"=>11, "page_views"=>24, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_On_the_Formation_of_Lipid_Droplets_in_Human_Adipocytes_The_Organization_of_the_Perilipin_8211_Vimentin_Cortex_/949320", "title"=>"On the Formation of Lipid Droplets in Human Adipocytes: The Organization of the Perilipin–Vimentin Cortex", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404307"], "description"=>"<p>(a) Grazing section of two very small LDs (approx. 200 nm in diameter) surrounded by numerous vimentin IF bundles of the “wide-spacing type” with approximately 30–50 nm spacing distance (cp.<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Franke1\" target=\"_blank\">[15]</a>) and with interconnection of LDs by filaments (arrows). (b) Similar small LDs associated with filaments as seen in (a). (c) Survey electron micrograph showing ordered arrays of vimentin bundles at the surface of LDs. Some LDs can be seen interconnected by IFs (arrows). First evidences of specific assembly of smooth ER tubules forming cisternae sheaths and layers can be noticed (brown-colored system of tubules). Note, at this stage of adipocyte differentiation mitochondria (M) are not directly associated to these forming small LDs. Bars: 0.50 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "micrographs", "vimentin", "filaments", "lds", "stimulation"], "article_id"=>949291, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g009", "stats"=>{"downloads"=>4, "page_views"=>21, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Electron_micrographs_showing_the_association_of_vimentin_filaments_with_the_forming_of_LDs_after_brief_AIM_stimulation_of_human_preadipocytes_/949291", "title"=>"Electron micrographs showing the association of vimentin filaments with the forming of LDs after brief AIM stimulation of human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404322"], "description"=>"<p>Delineated stages of the endogenous LD-formation are schematically presented with additional brief descriptions of individual steps and involved proteins and organelles, respectively. Sequential events in recruiting and combining proteins and organelles are drawn. The LDs exhibit phospholipid monolayer membranes with perilipin densely covering their surfaces. Perilipin attracts vimentin which further in turn seems to attract a specific ER tubular system (see results for more details). Outlined are double (LD-PLIN), triple (LD-PLIN-IF) and quadruple (LD-PLIN-IF-ER) complexes together with increasing sizes of LDs. Finally, the filament and ER parts of these complexes were relieved by so far unknown influences. In the last stages of differentiation, after the disappearance of the ER sheaths and vimentin, some mitochondria are seen to approach the perilipin-covered large LDs. EM, electron microscopy; for EM stages II-IV, see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone-0090386-g009\" target=\"_blank\"><b>Figs. 9</b></a><b>–</b><a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone-0090386-g011\" target=\"_blank\"><b>11</b></a> (EM stage I with the direct budding of tiny LDs from ER is rather difficult to see and therefore is drawn here as a suggested mechanism); for correlated different patterns of perilipin in immunofluorescence microscopy (IFM), see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone-0090386-g007\" target=\"_blank\"><b>Fig. 7</b></a>. ER, endoplasmic reticulum; M, mitochondria; PLIN, perilipin; LD, lipid droplet; IF, intermediated-sized filaments, i.e. vimentin in adipocytes; O =  and O∼, membrane phospholipids and cholesterol in ER bilayer membrane.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "pathway", "ld", "adipose", "cells", "featured", "perilipin", "vimentin"], "article_id"=>949305, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g015", "stats"=>{"downloads"=>2, "page_views"=>16, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_pathway_of_LD_formation_in_adipose_cells_featured_by_perilipin_and_vimentin_localization_/949305", "title"=>"The pathway of LD formation in adipose cells featured by perilipin and vimentin localization.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404306"], "description"=>"<p>(a) With short AIM stimulation for one day many tiny LDs positively stained for adipophilin can still be seen within the dense vimentin meshwork. (a) Additional short OA treatment for 3 hours immediately leads to exogenously derived, larger adipophilin positive droplets. Arrowheads mark two huge, vimentin surrounded cage-like structures. These are probably perilipin containing droplets (cp. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone-0090386-g007\" target=\"_blank\">Fig. 7d,d</a>) which have not yet been amalgamated by the new exogenously generated adipophilin droplets. DAPI (blue). Bars: 20 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "scanning", "microscopy", "vimentin", "adipophilin", "antibodies", "aim-stimulated", "compared", "oa-treated"], "article_id"=>949290, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g008", "stats"=>{"downloads"=>2, "page_views"=>20, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Laser_scanning_microscopy_showing_comparison_of_vimentin_and_adipophilin_antibodies_in_AIM_stimulated_compared_to_AIM_stimulated_plus_shortly_OA_treated_human_preadipocytes_/949290", "title"=>"Laser scanning microscopy showing comparison of vimentin and adipophilin antibodies in AIM-stimulated compared to AIM-stimulated plus shortly OA-treated human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404319"], "description"=>"<p>(<b>a</b>) Specific hydrophobic interactions are thought to be responsible for the general binding of PLIN proteins to surfaces of LDs (cp. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Heid1\" target=\"_blank\">[13]</a>). (<b>b</b>) Perilipin exhibits these LD targeting sites too. In contrast and uniquely compared to the other PLIN proteins, perilipin owns no helical domains at the C-terminus, but an additional acidic <b>E-rich</b> domain (see alignments given in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Heid1\" target=\"_blank\">[13]</a>, <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386-Hickenbottom1\" target=\"_blank\">[43]</a>). The possible interaction based on differently charged domains of perilipin and vimentin is presented. For details and for the implications of the proposed binding with the basic <b>R-rich</b> head region of vimentin see results and discussion. (<b>c</b>) A schematic summary of immunolocalization and of spatial arrangement in the proximity of emerging nascent LDs is depicted (not true-to-scale). Perilipin binds to the surface of LDs due to hydrophobic – hydrophilic short sequence domains. In a next layer, vimentin is attracted by perilipin and wraps the LDs tightly in “cage-like” spherical structures, followed by multiple concentric layers of smooth ER cisternae. This arrangement is delineated in a simplified bar-illustrated transversal section (bottom). (<b>c</b>) The schematic scenario shown in (<b>c</b>) is illustrated with a corresponding EM picture showing sheaths of non-fenestrated ER cisternae, regularly spaced dots of transversal sectioned vimentin IFs and a small rim of PLIN proteins directly bound to the LD surface. Abbreviations: E  =  glutamic acid; R  =  arginine; NT  =  N-terminal; CT  =  C-terminal; P  =  perilipin; Vim  =  vimentin; ER  =  endoplasmic reticulum. Bar in (<b>c</b>): 0.20 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "ld-plin-vimentin"], "article_id"=>949302, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g014", "stats"=>{"downloads"=>0, "page_views"=>14, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_LD_PLIN_vimentin_model_/949302", "title"=>"The LD-PLIN-vimentin model.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404316"], "description"=>"<p>(a) Cells reveal besides perilipin-positive also adipophilin-positive LDs. Localization was performed with mab AP125. A few positive, mostly smaller LDs are seen as well as plenty of non-labeled, larger LDs. Almost all LDs are closely associated and anchored with IF bundles. (b) A small, strongly immunolabeled LD is seen approaching a big, scarcely labeled LD. These LDs are obviously at the rim of combining. (c,d) Enlargements with details of filament attachment and immunolabeling sites which were highlighted by arrows (vimentin IFs) and arrowheads (adipophilin immunoreaction). Bars: a,b: 1 µm; c,d: 0.50 µm.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "microscopic", "localization", "adipophilin", "aim-stimulated", "oa-treated"], "article_id"=>949299, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g013", "stats"=>{"downloads"=>1, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Immunoelectron_microscopic_localization_of_adipophilin_in_briefly_AIM_stimulated_and_OA_treated_human_preadipocytes_/949299", "title"=>"Immunoelectron microscopic localization of adipophilin in briefly AIM-stimulated and OA-treated human preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404300"], "description"=>"<p>Mabs Peri112.17, Vim3B4 and control VE-cadherin were used. Immunoprecipitations (IPs) were analyzed by SDS-PAGE and CB staining (<b>a</b>) and corresponding WB using pab Peri-hNT (<b>b</b>). Lane 1: control unspecific binding of cell lysate to beads, i.e. magnetic beads after incubation with lysate and PBS washing; lane 2: bound lysate material of perilipin antibody beads; lane 3: control unbound supernatant of perilipin antibody beads; lane 4: bound lysate material of vimentin antibody beads; lane 5: control unbound supernatant of vimentin antibody beads; lane 6: bound lysate material of VE antibody beads; lane 7: control unbound supernatant of VE antibody beads; lane 8: control magnetic beads after incubation with crosslinked lysate; lane 9: bound crosslinked lysate material of perilipin antibody beads; lane 10: bound crosslinked lysate material of vimentin antibody beads; lane 11: bound crosslinked lysate material of VE antibody beads. Positions of molecular weight marker proteins are given on the left margin. Of importance, perilipin and notably vimentin antibodies precipitate and co-precipitate perilipin (<b>lanes 2,4;</b> perilipin position at 65 kD is marked at left margin by an arrow) in contrast to the negative control antibody (<b>lane 6</b>). In addition, similar positive reactions are obtained with crosslinked lysate by detecting crosslinked perilipin (<b>lanes 9,10</b>) whereas the control antibody shows no reaction with crosslinked material at all (<b>lane 11</b>).</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "perilipin", "vimentin", "antibodies", "aim-stimulated"], "article_id"=>949284, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g005", "stats"=>{"downloads"=>1, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Immunoprecipitations_using_perilipin_and_vimentin_antibodies_and_briefly_AIM_stimulated_preadipocytes_/949284", "title"=>"Immunoprecipitations using perilipin and vimentin antibodies and briefly AIM-stimulated preadipocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}
  • {"files"=>["https://ndownloader.figshare.com/files/1404297"], "description"=>"<p>Major treatments and involved PLIN proteins are shown. Preadipocytes containing many small LDs are <u>not</u> differentiated for several weeks with AIM containing media as conventionally described (top row; cp. <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0090386#pone.0090386.s001\" target=\"_blank\">Fig. S1</a>), but <u>only</u> very briefly (1–3 days), giving rise to “Adipocytes” (boxed area). Additional, short OA-treatment leads to “OA-Adipocytes” (boxed area, right side). Treatment with OA only - without AIM stimulation - leads to “OA-Preadipocytes” (bottom). Note the huge heterogeneity of sizes and colors of LDs seen within “OA-Adipocytes”. LDs are endogenously generated at the endoplasmic reticulum and stained positively for perilipin (“Endogenous-LDs”, red). Other LDs are obtained from the exogenous uptake of OA and stained positively for adipophilin, TIP47 and S3-12 (“Exogenous-LDs”, green). Merged LDs by fusion and mixed-type expression are seen by yellow color. The backway arrows indicate possible routes of LDs during lipolysis.</p>", "links"=>[], "tags"=>["Biochemistry", "lipids", "Fatty acids", "Lipid aggregates", "Lipid metabolism", "Lipid structure", "metabolism", "Metabolic pathways", "proteins", "Cytoskeletal proteins", "lipoproteins", "Protein interactions", "Structural proteins", "immunochemistry", "immunology", "Immunologic techniques", "immunofluorescence", "stimulation", "methods", "adipose", "types", "lipid"], "article_id"=>949281, "categories"=>["Biological Sciences"], "users"=>["Hans Heid", "Steffen Rickelt", "Ralf Zimbelmann", "Stefanie Winter", "Heiderose Schumacher", "Yvette Dörflinger", "Caecilia Kuhn", "Werner W. Franke"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0090386.g003", "stats"=>{"downloads"=>1, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Summary_scheme_and_brief_description_of_stimulation_methods_used_for_adipose_conversion_and_for_the_generation_of_different_distinct_types_of_lipid_droplets_/949281", "title"=>"Summary scheme and brief description of stimulation methods used for adipose conversion and for the generation of different, distinct types of lipid droplets.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-02-28 04:20:13"}

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Relative Metric

{"start_date"=>"2014-01-01T00:00:00Z", "end_date"=>"2014-12-31T00:00:00Z", "subject_areas"=>[{"subject_area"=>"/Biology and life sciences/Cell biology", "average_usage"=>[286]}, {"subject_area"=>"/Biology and life sciences/Developmental biology", "average_usage"=>[285]}]}
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