The identified patterns were analysed for their functional properties and evaluated for their use as clinical biomarkers. Using bioinformatics correlation analysis, the underlying regulatory networks were analysed. Furthermore the molecular mechanisms of endolysosomal lipid storage phospholipidoses and lipid-droplet formation in macrophages were studied. For this purpose monocyte-derived macrophages were challenged with the modified lipoproteins enzymatically degraded LDL Eldl and oxidised LDL oxLDL and the cellular lipid storage pattern visualised by fluorescence microscopy as well as performing lipidomic and transcriptomic analysis.
Cells from Niemann-Pick type C NPC disease patients were used to further characterise endolysomal cholesterol export, phospholipidosis and intracellular lipid trafficking. Special emphasis was put on BMP and cardiolipin species as endolysosomal and mitochondrial stress reporter molecules. The effects of lipid unloading using HDL and apoAI were studied in vitro in monocyte-derived-macrophages. The oxLDL induced LD response was also investigated under normoxic and hypoxic conditions and the role of PAT family proteins in macrophages during development of insulin resistance, in atherosclerosis and during chronic myocardial ischemia was studied.
Chronic and acute exposure to modified lipoproteins also influences leukocyte PMN lipid homeostasis and induces paracrine cell signalling. To address this subject, novel methods such as nanoparticle tracking analysis NTA were established and evaluated. Here direct effects of lipoproteins on granulocytes PMN were compared with effects on monocyte-derived-macrophages. EVs from granulocytes were charactersed in their lipidomic, miRNA and proteomic properties and functional aspects in atherosclerotic plaque maintenance studied.
EVs were also isolated from platelet preparations compared to the lipid and protein compositions of PMN-EV and evaluated as surrogate markers for platelet function. WP6 - Cellular lipid entry pathways into LDs and lamellar bodies WP6 was related to cellular lipid entry pathways into endolysosomal lamellar bodies and LDs. The major tasks were to establish and to apply methods for analysing lipid entry into different types of cells.
A major focus was on the role of lipid binding proteins in this process; especially the role of different ORPs oxysterol binding protein-related proteins , NPC proteins, and the function of saposins has been clarified with the aid of cells deficient in relevant proteins, the production of recombinant proteins, and the development of new tools for their analysis.
The characterisation of lipotoxic drugs and their role in LD formation has been addressed with the aid of lipid probes that have been modified by fluorescence, isotopes, and other structural modifications. MS has been applied for the characterisation of lipid uptake pathways, and its application to lipidomics has been extended. WP7 - Bioinformatics WP7 aimed to maximise the value of the data generated throughout the project, and also at providing new software tools for the processing, analysis and storage in the specific case of lipidomics data.
Task forces consisting of domain experts and bioinformaticians defined SPPs and defined the corresponding standard operating procedures SOPs for the different data types used in the project. The task forces developed a new nomenclature system and a data format standard to allow correct reporting and storage of MS MS -based lipidomics data. The wiki concept was employed to allow and encourage annotations and comments from the community at large.
These were all essential objectives to have the basic building blocks to provide a basic bioinformatics infrastructure for MS-based lipidomics data. In addition, a lipid identifier reference system was also created and is available via a new resource called 'LipidHome'. Furthermore, the curation of lipid-related pathways was one of the main objectives of this WP. An innovative tool for biochemical pathway tracking of non-labelled and stable isotope labelled lipid species analysed by MS was developed, including features such as the visualisation of the direction and significance of concentration changes in lipid species.
WP8 - Dissemination and training WP8 enhanced communication tools being continuously optimised in parallel to newest developments was planned to be used in the project. In close cooperation with WP1, WP8 members together with the project manager and integrated investigators disseminated project related information to consortium members and when was applicable to general public, organising meetings, conferences, workshops and exchange visits.
Efforts have been made to prepare and publish special issues on focused themes of the project. WP8 assisted the involved partners in promotion, dissemination and public release of the achievements of the consortium. WP8 ensured extensive dissemination activities taking place within the network of involved institutions as well as beyond. Amongst others, integration of partners, active involvement and training of young scientists and collection of publications were also objectives and subsequently, achievements of WP8. Project results: Lipids are central to the regulation and control of cellular processes by acting as basic building units for biomembranes, the platforms for the vast majority of cellular functions.
The recent developments in lipid MS have set the scene for a completely new way to understand the composition of membranes, cells and tissues in space and time by allowing the precise identification and quantification of alterations of the total lipid profile after specific perturbations. Lipid storage and release in multiple cells and tissues leads to transdifferentiation of multiple organs creating, fatty liver, obesity, white muscle and macrophage foam cells which are the hallmark of all energy overload diseases.
The project exploited recent advances in lipidomics technology to establish HT methods to define drugable targets and novel biomarkers related to LD lipid and protein species, their interaction and regulation during assembly, disassembly, storage and release. The research groups of various backgrounds and interested have been united in six WPs WPs 2 to 7 to study lipid protein interactions and investigate the dynamics of fat deposition and release in relevant cells as a hallmark of energy overload diseases with major health care impact in Europe.
Work progress and achievements during the period for WP2 Imaging technology: the focus in this project section was development of methods for LD imaging beyond the standard fluorescence imaging using commercially available dyes. LD is a lipophilic dye, based on the Bodipy fluorophore, for microscopic imaging of LDs. In contrast to previous LD dyes, it can be resolved spectrally from both green and red fluorophores allowing multi-colour imaging in both fixed and living cells.
Due to its slightly higher lipophilicity the dye can be used at about fold lower concentrations than the popular Bodipy, resulting in a significantly lower unspecific membrane background staining. Its improved specificity, brightness and photostability supports live cell imaging, which was used to demonstrate by two-colour imaging LD motility along microtubules.
For biological systems that either do not fluoresce or cannot tolerate the toxicity associated with staining and the photo-bleaching of fluorophores, their intrinsic chemical properties can be used as contrast mechanisms through coherent anti-Stokes Raman scattering CARS , To illustrate the method's potential the consortium gave a subcontract to Dr A.
Without a priori knowledge, the full Raman spectrum is recorded for each image voxel with high spatial resolution sub-femtoliter volume and with fast acquisition times in the order of milliseconds per voxel. As an example, quantitative chemical structure analysis of intra-organelle lipids has been demonstrated by mapping their degree of acyl chain unsaturation.
Major efforts were devoted to advanced electron microscopy and have led to significant improvements in EM immunocytochemistry of lipid structures like lipoproteins and LDs, in EM-tomographic analysis of LDs and in EM lipid tracing using click chemistry. These methods are particularly useful to study the complex organisation and the rapid dynamics of hepatic lipid metabolism and have direct impact on the work in other WPs, in particular WP5 and WP6. Using freeze fracture electron microscopy, possible pathways of LD growth were studied. Homotypic fusion of small LDs to create larger ones, is one proposed mechanism though the evidence for this process continues to be debated.
Possible fusion has been suggested from live cell fluorescence imaging, but the low resolution and other technical factors, such as vertical movement of structures out of the viewing plane, have prevented definitive conclusions to be reached. While thin-section electron microcopy has adequate resolution, extraction of lipids together with the surface phospholipid monolayers of closely apposed droplets during processing creates false fusion-like images. Freeze-fracture electron microcopy offers some advantages over other approaches.
Freeze-fracture typically reveals the LDs as closely associated but discrete spheroid or ovoid bodies with a characteristic layered structure. These features help define each droplet as an individual entity. Despite being so tightly pact that individual droplets appear to touch and even become flattened by mutual pressure at such contact regions, the boundaries of the individual droplets are normally quite distinct, suggesting a strong resistance to fusion.
Only occasionally is a possible site of continuity between the contents of apposed droplets seen. Whether such sites represent droplets caught in the act of fusion or the fracture path has simply failed to be deviated sufficiently to show the boundaries clearly at these sites cannot be determined with certainty. It is nevertheless striking that, from the appearance observed in freeze-fracture, the larger LDs frequently appear to be aggregates of smaller droplets.
The overwhelming impression created by these images is that numerous smaller LDs have amalgamated to create a larger one. One of the theoretical objections to fusion as a mechanism of LD growth is that, owing to the reduction in overall surface area, an excess of surface monolayer would result that could not be accommodated via the mechanisms operating in membrane-bound vesicles. MS Large amounts of different methods for mass spectrometrical lipid analysis were either newly developed or were improved and standardised or automated high throughput analysis.
This method is currently under DAkkS accreditation. In contrast to most published methods based on reversed phase chromatography, hydrophilic interaction liquid chromatography HILIC achieved good peak shapes, a short analysis time of 4. Also this method was validated according to FDA guidelines and showed excellent precision, accuracy, detection limits and robustness. The main advantage of the method is the rapid 8 min and simultaneous determination of a set of cholesterol precursors, plant sterols, oxysterols, and osyphytosterols with a short analysis time and excellent peak resolution.
The hyphenation of GC and tandem MS has successfully enhanced the performance of approved GC methods, both in analytical sensitivity and in time savings. Because the main applications of sterol profiling in human plasma can be seen in neurodegenerative diseases, atherosclerosis, and cholesterol metabolism disorders, we excluded the implementation of cholesterol epoxides, and other autoxidative species. The method offers a valuable tool for direct therapeutic stratification by discriminating with a single analysis hyperabsorber from hypersynthesizer patients.
Furthermore, a sterol profile identifies metabolic overload dependent uncoupling of cholesterol biosynthesis. The method contributes to the diagnosis of some rare genetic diseases like cerebrotendinous xanthomatosis, NPC disease or Smith-Lemli-Opitz-Syndrome in a single run. The system has been charactersed and is now generating analytical data from adipose and liver tissue samples, as well as various cell lines. This analysis system is complemented by an integrated platform for lipidomic analysis at the level of molecular species covering the workflow from sample preparation to data analysis.
The platform enables analysis of a broad range of lipids within one chromatographic run combined with identification of minor compounds present in amounts as low as 0. Furthermore it allows for fast profiling of lipid molecular species with regard to up and down regulation in search for lipid biomarkers. Flexibility is even further broadened with suitable internal standards added to samples.
In this case the platform affords quantitative determination of the lipidome. A special highlight is the analysis of phosphoinositides by MS. There has been a long standing difficulty with determining the phosphoinositides using MS. There are a number of reasons for this: the phosphoinositides are signalling molecules, consequently there generation and removal is tightly regulated and the more highly phosphorylated forms particularly PtdIns3,4,5P3 are short lived entities; additionally the phosphate groups on the inositol ring are labile and lost or they migrate around the inositol ring during extraction; the phosphoinositides are strongly charged and thus bind to cellular proteins making extraction difficult; the highly charged lipids also bind to glass and steel making recoveries low.
A number of methods have been reported, though none have been reproducible and easily adaptable. We have addressed the problem by adopting a derivatisation procedure which has the advantage that it stabilises the lipids and by shielding the charges reduces non-specific losses. Lipids are isolated from cell or tissue extracts and then the free phosphate groups are methylated using trimethylsilyldiazomethane. Following partial purification on a C4 reverse phase column, the diacyl groups of the lipids are identified following neutral loss of amu.
Inclusion of appropriate standards allows semi-quantification. Raman and CARS microspectroscopies The underlying hypothesis for this project has been the unique diagnostic capability of Raman and CARS microspectroscopies in offering non-invasive, label-free, and quantitative in-situ chemical analysis of lipid species with high spatial resolution sub-femtolitre volume and chemical structure information.
The specific aim has been the demonstration of the full potential of these enabling technologies in lipidomic research. Schmitz, P01 : A. The non-invasive chemical mapping of LD composition in living human macrophages in collaboration with University Hospital Regensburg, G. Schmitz, P The Raman-spectroscopic characterisation of polyene lipids as versatile labels in multi-photon imaging of cells and tissues in collaboration with University Bonn, Ch.
Thiele, P Our research achievements demonstrate the benefits and high potential of label-free Raman and CARS microscopies as complementary and non-invasive tools for further compelling applications in lipidomic research that circumvent current limitations in fluorescent-based microscopy approaches, and provide information that cannot be obtained by conventional biomedical microscopies and analytical techniques in a non-invasive manner.
The advances made within this project are currently prepared for publications in peer-reviewed journals.
Work progress and achievements during the period for WP3 The most significant results from WP3 are the in depth analyses of several new pathways that are key players in altered LD formation and contribute massively to the etiology of obesity and diabetes diabesity. Due to a collaborative effort we were able to identify the major components of some of these pathways also uncovering new druggable targets that might possibly be used for diabetes therapy.
Furthermore, we could show by combing analyses from human and mouse data that considerable proportion of these pathways are conserved allowing us to use the mouse model to better understand the progression of human disease. The detailed analyses are listed in the report below. All expression clones exhibiting a lipid binding protein insert can be used for further lipid binding studies.
To combine the protein array technology with MS, initially proteins have been coated on nitrocellulose stripes, followed by incubation and washing steps. The bound protein was then extracted from the nitrocellulose matrix and identified by MS. Additionally, when increasing the number of immobilised proteins, technical limitations such as spot finding were experienced. As an alternative, a bead-based protein array based on the Luminex technology was established for interaction studies.
To set up such an assay, two different coupling procedures were developed with regard to proteins purified in two different buffer systems. One buffer system is compatible to physiological conditions, the other system is based on denatured conditions. Refolding of structural epitopes may occur directly on the bead during sample incubation. This technology can be used in future studies not only for studying the interaction of lipids and proteins but also to study other molecules that might interact with different protein classes. In a targeted approach we used a gene array screen to identify genes which are deregulated in obese and type 2 diabetic patients and mice WP3 and WP4.
The list was filtered for possible lipid interactors which led to the identification of the orphan transcription factor ROR-gamma as a key driver of adipocyte formation. Through an MS coupled approach we were able to identify a tetra-hydroxylated bile acid as one endogenous ligand of ROR-gamma. We could demonstrate that dietary supplementation of THBA was able to protect from obesity associated type-2 diabetes development. However, mice treated with THBA had significantly smaller adipocytes due to a massive increase in adipogenesis.
Taken together we identified a functional lipid-protein interaction which might be important for modulating adipose tissue response to energy overload. As THBA is a natural occurring lipid it can be used in dietary intervention to treat obesity associated insulin resistance. Metabolic disorders and aberrant LD formation in the adipocyte The objective of this part of WP3 was to establish pathways that are essential for aberrant LD formation in adipose tissue using animal models of metabolic disorders.
Several new pathways were identified which might contribute to the etiology of type 2 diabetes development. To identify key signalling nodes that regulate LD formation and degradation in the mature adipocyte transcriptomics data from seven different animal models of metabolic disorders was analysed in collaboration between several partners using the bioinformatics platform Explain. The workflow to identify keynodes that can be used to model the transcriptional changes can be has been for the first time applied to the analysis of metabolic disorders. From these differentially regulated genes promoter information was extracted and used to calculate a Transcription factor matrix.
The matrix was used as a basis to identify key signalling nodes that regulate adipocyte function and that might be responsible for altered function in states of metabolic disorders. Caveolins form plasmalemmal invaginated caveolae, and the total absence of caveolin-1 by global gene invalidation in mice results in defective adipose tissue lipid storage capacity leading to progressive lipoatrophy, and resistance to high fat feeding induced obesity.
To answer this question, we took advantage of a mice model with tissue-specific caveolin deficiency to dissect the respective roles of adipocyte and endothelial caveolin. Adipose tissue development was have analysed in Cav1 null mice with global invalidation for Cav1 Cav1-KO and in Cav1 null mice in which caveolin-1 expression was rescued exclusively in endothelial cells but not in adipocytes Cav1-RC.
We could show that a lack of caveolin in adipocytes but not in endothelial cells accounts for adipose tissue lipoatrophy, decreased adipocyte size and adipocyte LD diameter reduction. In addition, we also observed that endothelial caveolin regulates macrophage infiltration and extravasation of immune cells into adipose tissue, thus unravelling distinct roles of endothelial and adipocyte caveolins. In addition we could show identify new pathways linking inflammation and especially immunoglobulins Ig released by B-lymphocytes as key drivers in the pathogenesis of diet-induced insulin resistance and type 2 diabetes.
Recent studies have suggested that during the development of obesity Igs play a key role in this pathology, although their mechanism of action is still unknown. Ig activate cellular responses by cross linking membrane receptors specific for the Fc portion of the Ig molecule FcR. We have investigated whether functional FcR play a role in the development of diet-induced adipose tissue inflammation and diet-induced insulin resistance. Taken together, these results demonstrate that the FcR gamma-chain acts as a sensor for HFD induced obesity leading to insulin resistance and adipose tissue inflammation.
Both proteins locate to the surface of LD and are believed to be lipases. Interestingly, in humans a non-synonymous genetic polymorphism in ADPN IM has been correlated to fatty liver diseases, even up to cirrhoses, but underlying mechanism were not known. These data indicate that ADPN, though having a slow TAG hydrolase activity, is not involved in degrading lipid, rather in lipid synthesis. Of note, activities of the human proteins were always higher than those of the murine proteins.
These date provide evidence that this mouse model is excellently suited for elucidation of the human situation. One pertinent regulator of this process is the protein TIMP1. TIMP1 mRNA and protein levels were measured in lean and obese animals also we analysed human samples for circulating Timp1 levels as part of the translational analyses. TIMP1 expression is increased in the serum and adipose tissue of obese mouse models. Recombinant murine TIMP1 is an inhibitor of adipocyte differentiation in subcutaneous primary preadipocytes.
This is of special interest since it is possible that altered TIMP1 levels in visceral adipose tissue might explain the increased contribution of this fat depot to the development of metabolic disorders. TIMP1-treated mice develop an impaired metabolic profile with increased circulating free fatty acid FFA levels, hepatic triacylglycerol accumulation and accelerated insulin resistance. Altered glucose clearance in TIMP1-injected mice is due to changes in adipose tissue glucose uptake, whereas muscle glucose clearance remains unaffected.
In conclusion it can be stated that the keynode TIMP1 identified by HTP screening as reported above is important for mouse and human physiology is a negative regulator of adipogenesis.
In vivo, TIMP1 leads to enlarged adipocytes in the state of over-nutrition. This might contribute to the detrimental metabolic consequences, such as systemic fatty acid overload, hepatic lipid accumulation and insulin resistance. Metabolic disorders and aberrant LD formation in the hepatocyte The objective of this part of the WP was to establish pathways that are essential for aberrant LD formation in hepatocytes and hepatic stellate cells HSCs using animal models of metabolic disorders.
It is interesting to note that nutritional stress exerted by starvation or HFD and genetic stress exerted by malfunctioning adipose triglyceride lipase ATGL always result in various forms of non-alcoholic fatty liver diseases NAFLD , the latter known in humans as 'neutral lipid storage disease with cardiomyopathy'. We postulated that the different etiologies for the steatoses observed in mouse models should have an impact on the lipidome of respective hepatocyte LD. From excised livers hepatocytes and there from LD were isolated, lipids extracted and subjected to lipidomic analysis.
Lipidclass-specific analyses were carried out at levels of lipid species profiling and lipid molecular species. First, we could show that LD numbers in freshly isolated hepatocytes were strongly induced upon HFD or starvation. Lipidomic profiling revealed that the TAG class responded most sensibly to stresses applied to the animals. Taken together, TG species and molecular species in hepatocyte LD provide the lipidome best suited to phenotyping and diagnosing steatoses of various etiologies.
Moreover, quantitative determination and structural analysis of acylglycerol and phospholipid species furnish metabolic insights. Under physiological conditions, HSC have been defined as the most important reservoir for vitamin A in mammals, stored in a large number of cytoplasmic LDs containing cholesteryl esters CE , triacylglycerols TAG and retinyl esters RE , the storage form of vitamin A.
Activation of HSCs has been recognised as one of the first steps in liver injury and repair. During activation, HSCs transform into myofibroblasts with concomitant loss of their LD and production of excessive extracellular matrix. Changes in the metabolism of retinoids and release of the hydrophobic lipid content associated with transformation of LDs are considered to be one of the main factors connected with HSC activation.
We have analysed different animal models based on our findings from transcriptomic analyses established in year of the project. The molecular identity of the enzymes involved in LD degradation in HSCs during activation is largely unexplored. In fact, SNPs rs and rs in the ATP10D gene were significantly associated with circulating C and C glucosylceramide species levels, obesity and insulin resistance. Furthermore, significantly elevated glucose and insulin levels and partially reduced insulin sensitivity were visible in Def mice. Def mice show also elevated hexosylceramide species levels.
Recently this family was found to regulate energy homeostasis. Cidec-KO mice are protected from diet-induced obesity, show elevated glucose uptake rates and better insulin sensitivity, and increased lipolysis rate. These findings demonstrate that the observed elevated expression of SCD1 has functional relevance and leads to enhanced desaturation of its substrates. As shown also in this study, these effects become ameliorated by the repaired gene in the transgenic ATP10D mice. Moreover, obesity is the entry gate for T2D.
Thus, it is not surprising that pandemics of obesity will be soon followed by an epidemic of diabetes. By it is estimated that million people will be afflicted by T2D diabetes with an estimated cost of USD 95 billion for health care.
Uniting academic and industrial members, 25 partners 20 academic research groups and 5 small and medium-sized enterprises SMEs jointed their forces investigating lipid storage and release. The technical advances, HT technologies, provided the optimal methodological possibilities to map the entire spectrum of lipids in cells, tissues and whole organisms.
Great condition for a used book! Lucas, C. Sperry, W. Matter from the University of Zurich, Switzerland. In essence, the endogenous water in the tissue was considered as a ternary component of the extraction system and sufficient chloroform and methanol were added to give a single-phase system for homogenization. Phase I: evaluation of secondary reference materials.
Translational research from mouse to man applied to LD pathology was another cornerstone of this large-scale project interfacing research and development. Net klinik. Deliverables Deliverables not available. Publications Publications via OpenAire. Viral genotype correlates with distinct liver gene transcription signatures in chronic hepatitis C virus infection Author s : Robinson, Mark W. Liver International Permanent ID: doi OSBP-related protein 2 is a sterol receptor on lipid droplets that regulates the metabolism of neutral lipids Author s : Hynynen, R.
Inflammasome is a central player in the induction of obesity and insulin resistance Author s : Stienstra, R. Liquid chromatographic-mass spectrometric analysis of phospholipids. Chromatography ionization and quantification. Caspase-1 deficiency in mice reduces intestinal triglyceride absorption and hepatic triglyceride secretion Author s : Diepen, J. Purification and characterization of phospholipase A2 from the pyloric caeca of red sea bream, Pagrus major. Purification and characterization of bile salt-activated lipase from the hepatopancreas of red sea bream, Pagrus major.
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