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Lactoferrin receptor mediates apo- but not holo-lactoferrin internalization via clathrin-mediated endocytosis in trophoblasts. LfR [Lf (lactoferrin) receptor] is expressed in most mammalian tissues, including placental trophoblasts, and is presumed to mediate the internalization of public competition, Lf. In Essay? However, the international law essay physiological significance of player, trophoblast LfR is not understood. Public International Law Essay? Using the CT (cytotrophoblast) cell model BeWo, we demonstrated that transfection with LfR siRNA (small interfering RNA) significantly decreased apo- but not holo-Lf uptake compared with mock-transfected controls and that apo- but not holo-Lf significantly increased MMP (matrix metalloproteinase)-2 activity. As Lf functionality is related to the presence (holo-Lf) or absence (apo-Lf) of iron within the Lf molecule, our results suggest that apo-Lf may play a role in cellular invasion. Essays? Moreover, we detected LfR (?105 kDa) in association with the plasma membrane, and public law essay competition ligand blotting confirmed that Lf binds to a LfR of ?105 kDa. Apo-Lf treatment significantly increased LfR abundance at the plasma membrane and leadership essay internalization probably occurs via clathrin-mediated endocytosis through early and recycling endosomes, as LfR was co-localized with EEA1 (early endosome antigen 1) and international competition TfR (transferrin receptor) using confocal microscopy, and hypertonic medium (0.4 M sucrose) significantly inhibited apo-Lf internalization. On Emotional? In summary, our data demonstrate that apo- but not holo-Lf is internalized by LfR and suggest that, following internalization via LfR, apo-Lf plays a role in CT invasiveness by inducing MMP-2 activity.
Moreover, LfR facilitates apo-Lf uptake specifically through clathrin-mediated endocytosis into early endosomes and international law essay potentially into underline author, a recycling pathway. Competition? Taken together, our data provide a new dimension in understanding ligand-dependant function that may be directly related to the ability of LfR to selectively internalize apo- but not holo-Lf. BeWo cell endocytosis lactoferrin lactoferrin receptor matrix metalloproteinase-2 (MMP-2) The presence of a LfR [Lf (lactoferrin) receptor] was first identified in human small intestine . It presumably facilitates the uptake of its protein ligand Lf, which is present in milk and mucosal secretions .
Therefore, kinetic studies on glass LfR have primarily been conducted in small intestine [3–6] and cultured enterocytes . We have previously cloned the LfR gene (GenBank® accession number NM_017625)  and determined that the intestinal LfR is an ?110 kDa trimeric (?37 kDa subunits), glycosylated protein . However, LfR is also expressed in non-intestinal tissues, such as heart, liver, spleen, thymus and skeletal muscle [3,8,9]. International Law Essay Competition? Recently, we detected LfR expression in mouse placenta associated with trophoblast cells , suggesting that LfR plays a role in this highly specialized cell type. Trophoblasts are divided into two subtypes; CTs (cytotrophoblasts) and STs (syncytiotrophoblasts), which are both maintained throughout pregnancy. Mckinsey? The primary function of international, CTs is mckinsey invasion into the uterine epithelium to international competition, secure successful implantation , whereas STs further implantation progression by establishing the nutrient and gas exchange barrier (maternal–fetal interface) and secreting hormones such as progesterone which are essential for successful pregnancy maintenance . The invasive capacity of CTs relies on thesis player the secretion of MMPs (matrix metalloproteinases), which function to degrade the surrounding ECM (extracellular matrix). Several disorders of pregnancy, including pre-eclampsia and spontaneous abortion, are believed to relate to ineffective CT proliferation, migration and invasion into the uterine endothelium . Lf has been attributed with multifunctional properties, including increased cell proliferation [12–16] and changes in cell morphology through activation of law essay, MMP transcription .
The pleuripotency of essay mckinsey, Lf may reflect the dramatic conformational change associated with iron binding and release [18,19] as holo-Lf (iron saturated Lf) and apo-Lf (iron free Lf) are known to adopt “closed” and “open” forms respectively [18,20]. For example, holo-Lf increases but apo-Lf decreases cell proliferation in human enterocytes (Caco-2 cells)  and international while apo-Lf stimulates microvessel proliferation in rats, holo-Lf is ineffective . Essay Mckinsey? Currently, the international competition mechanisms through which apo- and holo-Lf exert their differential functions are unknown. Herein, we hypothesized that the opposing functionality of glass essay, apo- and international law essay holo-Lf reflects differences in cellular internalization mechanisms mediated via LfR. In order to address this, we utilized the underline human placental CT cell line BeWo and public determined: (1) the effects of apo- and holo-Lf on essays on emotional MMP-2 activity, the law essay competition major MMP in trophoblasts; (2) the ability of the LfR to author in essay, facilitate apo- and holo-Lf uptake and internalization; and international law essay competition (3) the subsequent intracellular compartmentalization of LfR following Lf uptake. Anger? We also determined that apo- but not holo-Lf stimulates MMP-2 activity, that apo-Lf is internalized by LfR in a clathrin-mediated process and that internalized LfR associates with the early endosome/recycling compartment. This suggests that functional differences between apo- and holo-Lf are probably related to divergent uptake and intracellular trafficking mechanisms and international law essay provides a functional role for LfR in trophoblasts. BeWo cells were purchased from the author in essay American Type Culture Collection (CCL-98, Rockville, MD).
Cells were maintained in complete medium containing F12 Nutrient Mixture (Ham) (Life Technologies), supplemented with 10% fetal bovine serum (Gemini Bio-Products), 1.5 g/l sodium bicarbonate, 10 ?g/ml streptomycin and 10 units/ml penicillin (Sigma) at 37 °C in international competition, a humidified atmosphere with 5% CO 2 until 90% confluent. For experiments, cells were sub-cultured at a ratio of mckinsey, 1:3 and public international cultured until confluent. Lf was isolated from essay online vocabulary human milk as previously described with slight modifications . Casein was precipitated from pooled human milk with 0.06 M CaCl 2 , pH 4.6, for international 1 h at room temperature (25 °C) with constant stirring and centrifuged at 8600 g for 20 min at 4 °C. The whey fraction was collected, centrifuged at 8600 g for 20 min at 4 °C and then mixed with heparin–Sepharose Fast Flow resin (GE Health-Amersham Biosciences) overnight at glass menagerie essay 4 °C. The whey/resin mixture was centrifuged at public international 800 g for 5 min at 4 °C, the supernatant discarded and the remaining resin was packed on to a column and washed several times with wash buffer 1 (50 mM Tris/HCl, pH 8.0). To remove non-specific proteins, the column was rinsed with 5 bed volumes of elution buffer 1 (0.05 M Tris/HCl, 0.3 M NaCl, pH 8.0).
Next, Lf was eluted with 5 bed volumes of elution buffer 2 (0.05 M Tris/HCl, 1.0 M NaCl, pH 8.0) and fractions were collected. Underline Author? Lf-containing fractions were identified following Coomassie Brilliant Blue staining of SDS/PAGE (12%) gels. Samples were pooled and concentrated in YM-50 centrifugation tubes (Millipore Centriplus, Fisher Scientific) at 3000 g for 90 min at public 4 °C. The protein concentration of purified Lf was determined spectrophotometrically ( A 280 ) as previously established . Apo-Lf was prepared as previously described . Holo-Lf was prepared with FeSO 4 (Lf/Fe, 1:2 molar ratio) in sodium bicarbonate (Fe/HCO 3 , 1:1 molar ratio) for underline in essay 1 h at room temperature (25 °C). Unincorporated Fe was removed with YM-50 microcentrifuge tubes at 12000 g at international 4 °C and stored at ?20 °C. Essays Wellness? Fe saturation was determined spectrophotometrically by the A 460 / 280 ratio (22) .
To determine effects of apo- and holo-Lf on MMP-2 activity, cells were seeded in 24-well plates in complete medium until cells were 90–95% confluent. Cells were rinsed twice with 1?PBS and law essay then incubated with apo- or holo-Lf (50 ?g/ml) in F12 medium plus 2% FBS (fetal bovine serum) for 4 h. Conditioned medium was collected after 24 h and author concentrated to one third of the original volume using YM-50 microcentrifugation tubes (Millipore Centriplus) at 5000 g for 20 min at 4 °C. International? Protein concentration of conditioned media was determined using the Bradford assay (Bio-Rad) and proteins (20 ?g) were electrophoresed by SDS/PAGE (7.5% gel containing 1 mg/ml gelatin). Following electrophoresis, the gels were washed twice in 2.5% Triton X-100 in PBS for 15 min to remove SDS and menagerie essay incubated in wash buffer 2 (50 mM Tris/HCl, pH 7.6, 10 mM NaCl, 50 mM CaCl 2 ) overnight at 37 °C. Gels were stained with Coomassie Brilliant Blue for international law essay 2 h and destained with 10% acetic acid and 40% methanol in college admission online vocabulary, water. Absence of public international competition, stain (representative of proteolytic degradation) reflects MMP-2 enzymatic activity.
The relative densities of bands were quantified using the Chemi-doc Gel Quantification System (BioRad). Apo- and holo-Lf were labelled with 125 I (GE Health-Amersham Biosciences) by the Iodogen method . Excess free iodine was removed with a PD-10 desalting column (GE Health-Amersham Biosciences). To determine Lf uptake, cells were cultured in 24-well dishes in growth medium until confluent. Cells were incubated with 125 I-labelled apo- or holo-Lf (10000–100000 c.p.m.) in SFM (serum-free medium) for wellness 1 h at 37 °C. Law Essay Competition? Medium was collected, cells were rinsed once with rinse buffer (0.5 M NaCl, pH 3.0) and pooled with the leadership medium. Cells were washed three times with PBS, solubilized with 0.1 M NaOH and radioactivity was quantified in both medium and solubilized cells in a gamma counter. Affinity purification and validation of LfR antibody. LfR peptide (SSSPSLPRSCKEIKDE-Cys) was produced (Genemed Synthesis, South San Francisco, CA, U.S.A.) and international law essay competition conjugated (2 mg) to college admission essay vocabulary, an affinity-purification column (Sulfolink purification kit; Pierce Biotechnology).
LfR antibody was purified from law essay rabbit antiserum according to the manufacturer's instructions. To demonstrate antibody specificity, affinity-purified LfR antibody (1 ?g/ml) was pre-incubated with LfR peptide (1 mg/ml) in 5% (w/v) non-fat milk in on emotional wellness, PBS-T (PBS, 0.1% Tween) for 2 h at room temperature (25 °C) prior to immunoblotting of cell lysate as described below. Ligand binding and immunoblotting. Confluent cells were lysed in homogenization buffer [20 mM Tris/HCl, pH 7.5, 150 mM NaCl, 1% Triton X-100, 1 mM EGTA, 1?complete protease inhibitor (Roche Diagnostics)], sonicated three times for 10 s, centrifuged at 3300 g for 15 min at law essay competition 4 °C and glass menagerie the post-nuclear supernatant was collected. Protein concentration of the supernatant was determined by the Bradford method and public international cell lysates (50–100 ?g protein) were resolved by electrophoresis. To determine Lf binding, cell proteins were resolved by non-reducing SDS/PAGE at 80 V for example 4 h and transferred on to nitrocellulose membrane. Membranes were incubated with 5% BSA in PBS-T for 1 h at room temperature (25 °C) to block non-specific binding. Membranes were incubated overnight with 125 I-labelled apo-Lf, blots were rinsed twice with PBS-T and public international law essay bound ligand was visualized by autoradiography. For immunoblotting, cell extracts were elecrophoresed by SDS/PAGE (10% gel, 200 V for 1 h) under reducing conditions and membranes were incubated with LfR antibody for 1 h. Membranes were washed in PBS-T and primary antibody was detected with donkey anti-rabbit IgG-HRP (IgG-horseradish peroxidase; GE Health-Amersham Biosciences) in 5% (w/v) nonfat milk in PBS-T.
Membranes were washed in PBS-T for wellness 30 min, visualized with Super Femto Detection Chemiluminescence Reagent (Pierce Biotechnology) and public competition exposed to autoradiography film. Relative band density was quantified using the Chemi-doc Gel Quantification System (Bio-Rad). Transient transfection with LfR siRNA (small interfering RNA) Cells were seeded on to 24-well or 6-well plates and wellness cultured until 50% confluent, then transfected with 80 pmol of LfR siRNA (Ambion) in Lipofectamine™ 2000 (Invitrogen) for 24 h according to the manufacturer's instructions. Cells treated with Lipofectamine™ 2000 alone were used as mock-transfected controls. After 24 h, cells were incubated with 125 I-labelled apo- or holo-Lf in SFM at 37 °C. Lf uptake was measured after 30 min as described above. Public? To verify LfR suppression, transfected cells were scraped into 25 ?l of homogenization buffer (20 mM Tris/HCl, pH 7.5, 150 mM NaCl, 1% Triton X-100, 1 mM EGTA, 1?complete protease inhibitor) and immunoblotted as described above. Biotin-labelling of cell surface proteins.
To determine LfR abundance at example the plasma membrane, cells were seeded on to 100-mm-diameter dishes and cultured until confluent. Cells were rinsed with PBS and the cell surface was biotinylated with sulfo- N -hydroxysuccinimido-biotin (0.5 mg/ml; EZ-Link Sulfo-NHSS-Biotin; Pierce) at 4 °C for 1 h. Cells were rinsed with ice-cold 0.2 M glycine buffer, pH 5.0, and public international then with cold PBS to remove unbound biotin. Leadership Essay Mckinsey? Cells were lysed with lysis buffer [50 mM Tris/HCl, pH 7.4, 2 mM EDTA, 2 mM EGTA, 1?protease inhibitor (Roche Diagnostics)], sonicated (three 10 s bursts) and centrifuged at 100000 g for 20 min at 4 °C. The supernatant was discarded and the pellet was resuspended in lysis buffer with 0.2% SDS and incubated at public law essay 60 °C for 5 min. As the samples cooled, 1% Triton X-100 was added, the samples were re-sonicated and centrifuged at 100000 g for 20 min at 4 °C to essays, pellet any insoluble material. The supernatant was collected and incubated with 300 ?l of international law essay, slurry (1:1) of immobilized avidin beads (UltraLink Immunobilized Monomeric Avidin; Pierce) for 1 h at room temperature (25 °C) with constant rotation. To collect proteins bound to the resin, the samples were centrifuged at 2300 g for 1 min. Underline Author? The supernatant was discarded and the beads were rinsed with PBS and centrifuged for competition 1 min at 2300 g . To elute biotinylated proteins, an equal volume of Laemmli sample buffer (Bio-Rad) containing 2-mercaptoethanol (5%) was added and incubated at thesis team 90 °C for competition 5 min.
The samples were centrifuged at leadership 2300 g for 1 min and biotinylated proteins were resolved by SDS/PAGE and immunoblotted for LfR as described above. Determination of GPI (glycosylphosphatidylinositol)-anchoring by public international law essay, Pi-PLC (phosphatidylinositol-specific phospholipase C) cleavage. To determine if LfR is a GPI-anchored protein, cells were seeded on to 24-well plates and pretreated with Pi-PLC (1 unit/ml; Invitrogen) in SFM for wellness 1 h at 37 °C to public international law essay, cleave GPI-anchored proteins. To verify the glass effects of Pi-PLC on LfR abundance at public international competition the plasma membrane, cell surface proteins were biotinylated and college online vocabulary immunoblotted for LfR as described above. To determine the effects of public competition, Pi-PLC on Lf uptake, cells were rinsed with PBS, fresh SFM was added and cells were incubated with 125 I-labelled apo-Lf at 37 °C and uptake was measured after 1 h as described above. To illustrate the efficacy of the enzyme, spent medium was collected and concentrated to one third of the original volume using YM-50 microcentrifugation tubes (Millipore Centriplus) at glass menagerie 5000 g for 20 min at 4 °C and immunoblotted for ALP (alkaline phosphatase), a known GPI-anchored protein. To inhibit endocytosis, cells were seeded on to 24-well plates and public competition treated with hypertonic medium (0.4 M sucrose). Cells were pre-treated in hypertonic medium for essays wellness 30 min and incubated with 125 I-labelled apo-Lf in SFM at 37 °C in the presence of 0.4 M sucrose and uptake was measured after 60 min. International Competition? Lf uptake was measured as described above.
To determine if de novo LfR protein synthesis is required for Lf uptake, cells were seeded on to 12- or 24-well plates until confluent and pretreated with cycloheximide (Cyhex, 10 ?g/ml, Sigma) in SFM for 6 h at 37 °C. Cells were either collected for SDS/PAGE analysis or incubated with 125 I-labelled apo-Lf in the presence of in essay, Cyhex at 37 °C and Lf uptake was measured over 7 h. To determine the public subcellular localization of LfR, cells were seeded on thesis to glass coverslips, cultured for 3–4 days in public international law essay, growth medium and incubated with apo-Lf (where indicated, 200 ?g/ml) in SFM for leadership essay up to competition, 1 h at 37 °C. After treatment, cells were rinsed with PBS and fixed with 4% phosphate-buffered paraformaldehyde for college essay 10 min and permeabilized with 0.2% Triton X-100 in public law essay, PBS for 5 min. To eliminate non-specific binding, cells were incubated in blocking buffer (5% goat serum in PBS) for 30 min followed by incubation with affinity-purified LfR antibody (0.8 ?g/ml) in PBS containing 5% goat serum, 1% BSA and 0.2% Triton X-100 for on emotional 1 h. Cells were rinsed with PBS and LfR was detected with Alexa 488-conjugated-anti-rabbit IgG (1 ?g/ml; Molecular Probes) in PBS containing 0.2% Triton X-100 and 1% BSA for 20 min. Co-localization of LfR with specific subcellular markers was determined following incubation with antibodies for clathrin (1:500, Abcam); EEA1 (early endosome antigen 1; 1:500, Abcam), TfR (transferrin receptor, localizes to recycling endosomes, 1 ?g/ml; Zymed Laboratories); LAMP1 (lysosome-associated membrane protein 1; 1:200, Abcam); M6PR (mannose 6-phosphate receptor, localizes to international competition, late endosomes; 1:500, Abcam) and detected with Alexa 568-conjugated-anti-mouse IgG (1 ?g/ml; Molecular Probes). College Vocabulary? After several rinses, coverslips were mounted with ProLong Gold (Molecular Probes) and sealed with nail polish. Immunofluorescence imaging was performed using an Olympus BX50WI microscope with UPlanApo ?100 magnification under an oil-immersion lens [NA (numerical aperture), 1.00], and digital images were captured using the Bio-Rad Radiance 2100 confocal system with LaserSharp2000 software, version 4.1 (Bio-Rad). Results are presented as means±S.D. of triplicate samples from 2–3 separate experiments. Statistical comparisons were performed using Student's t test or one-way ANOVA (Prism Graph Pad) and significance was set at P 0.05.
Active form of MMP-2 is increased by apo- but not holo-Lf. Secretion of MMP-2 plays a key role in CT invasion, thus we determined effects of Lf and public international competition Lf conformation on MMP-2 activity in BeWo cells. Using gelatin zymography, our data demonstrated that apo-Lf significantly increased (20%) MMP-2 activity compared to untreated cells (Figure 1). In contrast, we observed no effect of holo-Lf on MMP-2 activity, suggesting a differential effect of Lf conformation on cellular invasivity of CTs. Representative zymogram (upper panel) of culture medium collected from cells treated with either apo- or holo-Lf. Thesis Player? Equal amounts of protein (20 ?g) were electrophoresed through polyacrylamide gels containing gelatin and stained with Coomassie blue. MMP-2 activity was detected as clear bands (digested gelatin) on a blue background.
MMP-2 activity was significantly increased in cells treated with apo-Lf as compared with untreated (control) and holo-Lf treated cells (lower panel). Data shown represent MMP-2 activity as a percentage of untreated control cells (mean±S.D.) for two independent experiments ( n =4). * P 0.05. LfR facilitates uptake of apo- but not holo-Lf. To determine if LfR mediates apo- and/or holo-Lf uptake, uptake studies using 125 I-labelled Lf were performed. Public International Law Essay? Our data demonstrated that BeWo cells internalized 9-fold more apo-Lf compared with holo-Lf (Figure 2). Using affinity-purified LfR antibody, we confirmed that BeWo cells expressed LfR, as specifically immunoreactive proteins were detected at ?35 and ?105 kDa in BeWo cells (Figure 3). Ligand binding plays a critical role in receptor function, and ligand binding established that radiolabelled apo-Lf bound to two LfR proteins at ?75 and essays ?105 kDa (Figure 3), but not to the 35 kDa protein. We speculate that the 35, 75 and international competition 105 kDa moieties represent LfR monomers, homo-dimers and -trimers, respectively. The ability of LfR to internalize apo-Lf is dependent on its cellular localization and we showed that the college admission online 105 kDa LfR was localized to the plasma membrane using cell surface biotinylation followed by immunoblotting (Figure 4A). In contrast to what was reported for the intestinal LfR , trophoblast LfR does not appear to international competition, be GPI-anchored as demonstrated by internalization of thesis player, Lf following GPI-anchored protein cleavage with Pi-PLC (Figure 4B).
Confirmation of Pi-PLC cleavage was verified as Pi-PLC increased ALP abundance in spent medium compared to public international, untreated cells (Figure 4C). Confluent cells were treated with 125 I-labelled apo- or 125 I-labelled holo-Lf and Lf uptake was determined. Essay Online Vocabulary? Representative graph of uptake after 1 h illustrates that significantly less holo-Lf was internalized when compared to apo-Lf. Public Competition? Data shown represent mean Lf uptake (Lf uptake/mg protein)±S.D. from two independent experiments ( n =4). * P 0.001. Representative immunoblot ( A and B ) and ligand blot ( C ) of cell lysate proteins. Underline In Essay? Proteins (50 ?g) were resolved by SDS/PAGE and immunoblotted with LfR antibody ( A ; LfR Ab, 1 ?g/ml) or pre-absorbed with excess Lf antigen ( B ; LfR Ab+peptide) for 1 h. The LfR antibody detected specific proteins at law essay competition ?105 and glass menagerie ?35 kDa ( A ) which were both completely eliminated following pre-absorption with LfR peptide ( B ). ( C ) Cell lysate (100 ?g) was resolved under non-reducing and non-denaturing conditions, and law essay incubated with 125 I-labelled apo-Lf. Detection of ligand binding to proteins at glass essay 105 and 75 kDa was observed. Confluent cells were treated with Pi-PLC and surface expression and Lf uptake was assessed. International? ( A ) A representative LfR immunoblot of admission vocabulary, biotinylated cell surface proteins documents the presence of LfR on the plasma membrane after Pi-PLC treatment. ( B ) Cleavage of GPI-anchored proteins with Pi-PLC did not significantly reduce 125 I-labelled apo-Lf uptake in BeWo cells.
Data represent mean Lf uptake/mg protein±S.D. for two independent experiments ( n =4). International Law Essay? ( C ) Representative immunoblot of ALP antibody from underline total spent medium following Pi-PLC treatment, validating enzyme activity. In order to determine a role for LfR in law essay, Lf uptake, LfR expression was attenuated in cells transfected with LfR siRNA (Figure 5A). We demonstrated that reduced LfR abundance resulted in significantly lower apo-Lf uptake (30%) compared with mock-transfected controls (Figure 5B). In contrast, holo-Lf uptake was not significantly reduced (10%), suggesting that internalization of holo-Lf in BeWo cells is not predominantly LfR-mediated. Due to our finding of limited LfR-mediated holo-Lf uptake and lack of an effect of holo-Lf on MMP-2 activity, we subsequently focused on effects of leadership, apo-Lf on the intracellular trafficking of LfR. ( A ) Representative LfR immunoblot of total protein illustrating significant attenuation of LfR compared to mock-transfected cells. Law Essay? ( B ) Lf uptake was measured using 125 I-labelled apo- or 125 I-labelled holo-Lf in glass menagerie, cells transfected with LfR siRNA or mock-transfected controls. Data represent the competition means±S.D. ( n =2–3) of triplicate experiments. * P 0.05, ** P 0.01, as analysed by Student's t test. LfR internalization is a clathrin-dependent process and potentially follows a recycling pattern. As our data clearly demonstrated that apo-Lf is internalized via LfR, we then determined the mechanisms responsible for anger essay internalization of apo-Lf via LfR.
CTs were treated with apo-Lf and LfR protein abundance on the plasma membrane was assessed following cell surface biotinylation. International Law Essay? We noted that treatment with apo-Lf resulted in a rapid (10 min) and significant increase in cell surface-associated LfR (Figure 6A), independent of changes in total LfR abundance (Figure 6B). This suggests that apo-Lf may signal the essays on emotional wellness recruitment of LfR to the plasma membrane or that perhaps Lf binding to public, LfR transiently interferes with its internalization. Confluent cells were treated with or without apo-Lf and LfR surface expression was assessed. ( A ) Representative immunoblot of crude membrane proteins from cells treated with apo-Lf or untreated and detected with LfR antibody. Representative graph illustrates that LfR abundance significantly increased with apo-Lf treatment compared to college admission essay, untreated cells with no effect on the total amount of LfR protein expression ( B ). Public? * P 0.001, as analysed by Student's t test. ?-actin was used as a loading control. Data represent means±S.D. ( n =3) of team player, 3 independent experiments. Multiple pathways for internalization of plasma membrane proteins have been characterized. To identify the mechanism responsible for LfR internalization, CTs were treated with hypertonic medium (0.4 M sucrose) to inhibit clathrin-dependent endocytosis, which resulted in a robust inhibition of public international, apo-Lf internalization (40%) compared with untreated cells, with values of 2.4±0.25 and team player 5.7±0.42 pmol/mg protein±S.D. for treated and control cells respectively; n=5–8 per public international law essay group, * P 0.001. Leadership Essay? To confirm the mechanism by international, which LfR is glass endocytosed, we used confocal microscopy to public international law essay competition, identify the subcellular compartments within which LfR resides. As indicated in Figures 7(A) and glass menagerie essay 7(B), LfR and clathrin co-localized and were largely associated with the international law essay plasma membrane, as illustrated by team, yellow pixilation in the merged images (Figure 7C).
As internalization is ligand-stimulated, cells were treated with apo-Lf, and indirect immunofluorescence was used to public, visualize specific sub-cellular compartments within which LfR was associated. Following ligand stimulation, we demonstrated partial co-localization of LfR and the early endosome marker EEA1 in underline author, vesicular/endosomal-like structures (Figures 7D–7F). As co-localization between EEA1 and public law essay competition LfR was not exclusive, we sought to examine other sub-cellular compartments containing LfR. Clathrin-mediated endocytosis is one mechanism cells utilize to control cell surface receptor abundance and turnover via recycling. To investigate whether LfR trafficking from the plasma membrane potentially includes endocytic recycling, we used the underline author in essay indirect immunofluorescence of law essay competition, TfR as a marker for recycling endosomes. Thesis Team Player? We showed partial co-localization of international law essay competition, LfR and TfR within distinct internalized vesicles in cells following ligand stimulation (Figures 7G–7I).
No co-localization between LfR and M6PR (a late endosome marker) and minimal co-localization with LAMP1 (a lysosome marker) was detected (results not shown). These results indicate that, in response to apo-Lf stimulation, LfR is internalized via clathrin-mediated endocytosis and leadership mckinsey consequently enters a distinct intracellular endocytotic pathway. Confocal microscopy identifying specific sub-cellular compartments associated with LfR. Cells were fixed with paraformaldehyde and public international law essay competition imaged for LfR (green) and sub-cellular markers (red); clathrin ( A and C ), EEA1 ( D and F ) and leadership essay mckinsey TfR ( G and I ). Clathrin ( A ) and LfR ( B ) were co-localized (yellow, C ) and international law essay competition stained in a plasma membrane-like pattern in cells. Following apo-Lf treatment, partial co-localization with the early endosomal marker EEA1 ( D ) and LfR ( E ) was detected (yellow, F ) and glass menagerie treatment of cells with apo-Lf clearly resulted in vesicularization of LfR.
Furthermore, co-localization with the public international recycling endosome marker TfR ( G ) and player LfR ( H ) was detected (yellow, I ) following treatment of cells with apo-Lf. Finally, to determine if increased abundance of LfR at the plasma membrane in response to apo-Lf stimulation reflects changes in law essay, intracellular LfR recycling, we blocked de novo protein synthesis with Cyhex and measured apo-Lf uptake. Inhibition of new receptor synthesis did not diminish apo-Lf uptake, indicating that LfR is recycled. In fact, apo-Lf uptake was significantly higher in Cyhex treated cells (Figure 8A), suggesting that apo-Lf enhances LfR recycling. On Emotional? Moreover, our data indicated that CT-derived LfR is a relatively long-lived protein (Figure 8B), as abundant LfR remained detectable after 7 h of Cyhex treatment and that de novo protein synthesis may play a minor role in sustaining total and surface receptor abundance. Confluent cells were pre-treated with Cyhex and international competition uptake of 125 I-labelled apo-Lf and LfR abundance was assessed compared with untreated cells. ( A ) Apo-Lf uptake from cells significantly increased in the presence of cycloheximide (Lf+Cyhex) after 3 h compared with untreated cells (Lf). Data represent means±S.D. for two independent experiments ( n =3–4), and wellness * P 0.05 compared with untreated cells, as analysed by Student's t test. Competition? ( B ) Representative immunoblot of total protein extract isolated from example cells pre-treated with Cyhex then incubated with apo-Lf for 3 and 7 h compared with untreated cells. The immunoblot demonstrates that Cyhex did not affect LfR abundance. ?-Actin was used as a loading control. CTs are specialized cells in the placenta that are initially responsible for invasion into the uterine endothelium, assuring optimal fetal development . Public Competition? Our previous detection of LfR along the maternal and fetal interface in the placenta implicates LfR in CT function [8,9,27].
The aim of the present study was to explore the function of Lf in essay, the placenta as mediated via LfRs using a CT cell model. International Law Essay? While primarily regarded as a component of anger, milk , Lf is also found in decidua (9–95 ?g/g), amniotic fluid (2–37 ?g/g) and chorion (2–26 ?g/g) , and is secreted directly from international competition uterine epithelia [29,30]. This suggests that Lf secreted from the uterine endothelium may regulate CT function in on emotional, a paracrine manner. Law Essay Competition? Results from this study documented an enhancing effect of apo- but not holo-Lf on underline author in essay MMP-2 activity in international law essay competition, CTs, suggesting a regulatory role for essay apo-Lf in CT invasivity as this process is dependent upon the secretion and activation of public international law essay, MMPs. Leadership? There is precedence for law essay competition effects of Lf on MMP regulation as over-expression of Lf activates MMP-1 expression through binding to essay, AP-1 promoter elements and through stimulation of the p38 MAPK (mitogen activated protein kinase)-dependant pathway in mouse fibroblasts and human COS-1 cells . International Law Essay? We speculate that apo-Lf may similarly regulate the transcription of MMP-2 via AP-1 promoter elements which are present in the MMP-2 gene . In support of the essays concept of Lf acting as a transcription factor, a nuclear localization signal within the N-terminal region of the Lf molecule (amino acids 1–5) has been previously identified , and Lf binds to DNA [33,34]. Law Essay Competition? Alternatively, the induction of MMP-2 via apo-Lf could be through the essays p38 MAPK pathway as p38 activation is required for the up-regulation of MMP-2 protein expression . Public International Competition? The ability of Lf to activate p38 MAPK has been previously shown in other cell types [16,35], providing support for a direct role of apo-Lf on CT invasivity.
The effect of apo- but not holo-Lf on MMP-2 activity probably reflects that distinct functions are associated with either apo- or holo-Lf. In contrast to reports in wellness, rat hepatocytes , CTs bound and public law essay internalized both apo- and holo-Lf. However, in CTs, apo-Lf was internalized more avidly than holo-Lf, suggesting preferential uptake mechanisms specific to apo-Lf which may reflect the requirement of this molecule for CT-specific functions such as invasivity, while precise Lf-dependant functions in hepatocytes remain unknown. Gene attenuation of LfR conclusively determined that LfR primarily facilitates apo-Lf uptake in essay, CTs. Recently, distinct Arg-Gly-Asp (RGD) binding motifs were found within the Lf molecule , which have been postulated to determine ligand binding and receptor interactions.
It is possible that iron-saturation, which alters Lf conformation, may result in LfR inaccessibility to these binding motifs. Competition? While this is not to imply that holo-Lf binding and essays on emotional wellness uptake does not elicit cellular events, it does not appear that these effects are mediated via LfR internalization in CTs. Possibly, holo-Lf internalization is not a prerequisite to stimulate cellular events at public international competition the plasma membrane as suggested by other investigators [15,35]. As apo-Lf was preferentially taken up by on emotional wellness, LfR, we sought to public competition, characterize the placental LfR more extensively. It has previously been reported that intestinal LfR is a GPI-anchored protein . Contrary to this observation, we determined that, in CTs, the LfR receptor is not a GPI-anchored protein and is in fact internalized via clathrin-mediated endocytosis following ligand binding. Our observations in trophoblasts are consistent with previous reports in other cell types such as hepatocytes, where inhibition of clathrin-mediated endocytosis by hypertonic medium  significantly reduced Lf internalization.
We speculate that differences in Lf uptake between intestinal and CT LfR may reflect differences in mckinsey, cellular function, as CTs are a semi-pleuripotent and invasive cell type, whereas enterocytes are terminally differentiated polarized cells involved in public, vectorial nutrient transport. Leadership Essay? Our determination that LfR is internalized via a clathrin-dependent mechanism may provide insight into international competition, LfR structure, as receptors utilizing clathrin-mediated endocytosis require a cytoplasmic, tyrosine-based sorting signal for anger essay example interaction with adaptor proteins, such as AP2 . Public Law Essay? To identify potential AP2-interacting motifs within the LfR sequence we used a proteomics server (www.elm.eu.org) and indeed two tyrosine-sorting motifs were identified at amino acids 120–123 and 129–142. This further suggests that a portion of LfR protein does reside intracellularly and that one or more of glass menagerie essay, these motifs may functionally interact with AP2 to facilitate LfR internalization. LfRs have been identified in various cell-types and reports have determined a molecular mass of ?110 kDa . Recently, Tsuji et al.  examined the structure of human LfR in greater detail and determined that it is an 105.5 kDa oligomeric protein made up of three 35.5 kDa subunits, similar to what we observed in public international law essay competition, trophoblasts. Interestingly, while we detected both the monomeric (35 kDa) and author in essay trimeric (105 kDa) forms of international law essay competition, LfR under reducing conditions, the essays abundance of the trimer was always greater that the monomer.
For some reason, we were unable to extensively reduce the trimeric form to monomers even when using dithiothreitol and iodoacetic acid (V. Lopez, unpublished observations). Law Essay Competition? Interestingly, we did not detect the dimeric form of LfR under reducing conditions, which is similar to in essay, the findings of Tsuji et al. . We further demonstrated that apo-Lf binds to both dimeric and public law essay trimeric LfR multimers, which agrees with previously published reports demonstrating that human LfR forms a homo-trimer [7,40] as a result of subunit oligomerization . We speculate that the trimeric form is an unusually stabilized form, mediated by several intracellular disulfide bonds and possibly also hydrophobic interactions, whereas the dimeric form is less stable and on emotional can be reduced to public international competition, the monomeric form. In contrast, a 45 kDa calcium-dependent LfR has been reported in anger essay example, rat hepatocytes ; however, it is unknown how this LfR relates specifically to Lf function as rats do not express Lf. It is well established that cell signalling and cellular homoeostasis are altered by increasing or decreasing cell surface receptor abundance . As LfR abundance at the plasma membrane did not require de novo synthesis, our data suggest that recycling of public law essay, LfR to and from the plasma membrane occurs. Endocytic recycling requires internalization of cell surface receptors and their ligands within clathrin coated pits , with the interaction between transferrin and TfR being the classic example of this process . On Emotional? Receptor recycling is public law essay competition one strategy by which cells maintain a mobilizable receptor pool to quickly traffic to the cell surface in underline author in essay, response to physiological stimuli, to allow cells to redistribute cell-surface molecules to public international law essay, specific cell locations or compartments undergoing a physiological activity .
We speculate that Lf may either stimulate the recruitment of LfR to underline author, the plasma membrane, or that perhaps Lf binding to the receptor results in transient accumulation of LfR prior to internalization via clathrin-mediated mechanisms. We propose that CTs may require Lf/LfR internalization to public international law essay, rapidly facilitate implantation, allowing for optimal embryo-fetal development. Further studies are needed to address this issue. Collectively, the results from this study demonstrated a specific role for apo-Lf but not holo-Lf in regulating MMP-2 activity and underline thus potentially endothelial invasion in CTs. Importantly, we have determined that apo- but not holo-Lf is internalized via LfR, which occurs through a clathrin-dependent pathway in CTs suggesting a mechanistic and functional difference between apo- and holo-Lf. These results may help to explain the pleuripotent and cell-specific functions which have been associated with Lf.
The authors would like to international, express their gratitude to Dr. Lisa L. Gough for critical reading of the admission essay manuscript. This work was supported by NIH grant number HD-043240. Abbreviations: ALP, alkaline phosphatase; apo-Lf, iron free lactoferrin; CT, cytotrophoblast; Cyhex, cycloheximide; EEA, early endosomal antigen; GPI, glycosylphosphatidylinositol; holo-Lf, iron saturated lactoferrin; LAMP, lysosome-associated membrane protein; Lf, lactoferrin; LfR, Lf receptor; M6PR, mannose 6-phosphate receptor; MAPK, mitogen activated protein kinase; MMP, matrix metalloproteinase; Pi-PLC, phosphatidylinositol-specific phospholipase C; siRNA, small interfering RNA; SFM, serum-free medium; ST, syncytiotrophoblast; TfR, transferrin receptor.
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MLA Essay Format: Help with Writing Your Essay. MLA (Modern Language Association) style is international law essay, used in writing custom essays, research and term papers in many fields. MLA essay format is most widely used in the field of humanities and essay liberal arts. The Modern Language Association recommendations to formatting essays were updated in 2009. Among the new rules of formatting, the Works Cited or References list is the requirement of indicating the medium of publication for every cited source.
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10 essays that will get you hooked on international Joan Didion. I first read Joan Didion when I was an editorial assistant at a big publishing house in Manhattan. Essays Wellness! Blue Valentine was the movie everyone in public international competition, the bullpen was discussing, and underline in essay, I YouTubed Ryan Gosling and Michelle Williams to see how their chemistry translated off-screen. Fortuitously, I came across a Nightline segment where Michelle discussed Heath Ledger’s death, and how she read Didion’s The Year of Magical Thinking to help cope with his passing. “It didn#8217;t seem unlikely to me that he could walk through a door or could appear from behind a bush,” she said in the interview. “It was a year of very magical thinking.” I hadn’t yet known The Year of Magical Thinking was a memoir about the grief Didion endured after losing her husband, novelist John Gregory Dunne, nor did I know it had won her the law essay, 2005 National Book Award for Nonfiction. All I knew was the term “magical thinking” intrigued me; that in the midst of college admission essay, pain we believe the impossible is possible. I went to the Borders (RIP) by Madison Square Garden that night, and devoured the book in law essay, one sitting. Thus began my new life as a Didion fanboy. I’m often asked why I love Joan Didion so much. My response is simple: I don’t read her essays, they read me. When I picture her in my mind, I see a bespectacled mercenary driving through an apocalyptic landscape in a white corvette, not in underline in essay, search of international law essay, bounty, but of underline in essay, answers. In celebration of her upcoming documentary, We Tell Ourselves Stories In Order to Live , and the Didion biography The Last Love Song out later this month, we’ve compiled a list of the best nonfiction works to get into law essay, the work of essay, Didion.
Below are the top ten essays that have made us cry, think, and, above all else, believe. “In The Realm of the Fisher King” ( After Henry ) After Henry , originally published in 1992, is perhaps Didion’s most unsung collection. Public International Competition! In it she shatters illusions constructed by the American media in the 80s, while simultaneously chronicling the political atmosphere across the author, nation. “The Realm of the public international law essay, Fisher King” gives a raw and personal look at the Regan years through the lens of Peggy Noonan, Regan’s speechwriter, and example, First Lady Nancy Regan. What makes this essay interesting isn’t just that she compares the Regans to actors on set, but rather the inside look at their everyday life. For example, President Regan kept photos of families who wrote to him in his desk drawer and Mrs. Regan was shocked they had to pay for their own meals. International Law Essay! This is a must read for historical buffs. Without a doubt, this is one of Didion’s most celebrated works and with good reason. In it she defines the glass menagerie, parameters of self-respect, and preaches the importance of character. “People with self-respect have the courage of their mistakes. They know the price of things.” I first read this essay during a dark period in international law essay, my life, when I thought none of my actions had consequences. Man, was I ever wrong. Essay! The price I paid was heavy, and when I decided to shape up, I kept a copy of the above quote in my wallet to remind myself never again to public compromise my integrity.
Want a kick-ass Easter egg? Did you know the essay first appeared in thesis team, the pages of Vogue back in 1961? Yup, Didion worked there after she won a competition in college. This essay is so legendary, it can be viewed on Vogue.com in its original format. Check it out! Okay, I’m cheating a little here. Technically this next selection on our list is a book, not an essay. But hear me out. Miami began to change when many Cubans sought asylum in the Sunshine State back in the early 60s. Spanish-speaking households, quinceaneras, and salsa may all be common aspects of Miami’s culture today, but in the 1980s they weren’t. Miami is Didion’s analysis of the significant transformation Miami underwent.
As a Cuban-American myself, I struggled with some of Didion’s opinions in this book. Public Law Essay! In particular, she has a unique perspective about the Cuban exodus and its ramifications on Miami#8217;s African-American community. It’s a tough read, but worth it. #8220;John Wayne: A Love Song” ( Slouching Towards Bethlehem ) This is anger essay example, my favorite Joan Didion essay because she describes her loss of innocence. Law Essay Competition! In this essay she describes the on emotional wellness, first time she saw John Wayne in the summer of 1943, when she was eight years old and watching a film called War of the Wildcats . This is an essay about coming to public law essay competition terms with your idol’s mortality. She, like many others, did not believe John Wayne could fall ill and die.
And when it happened, it called into question everything she believed. Forty plus years later, this essay still inspires much debate. Joan Didion writing about the college essay online vocabulary, women’s movement may seem like pure fan service today, but in public international law essay competition, 1972 this was actually a contentious essay. Written in anger essay, the early years of Second Wave feminism, “The Women’s Movement” quotes legendary feminist Shulamith Firestone (author of The Dialect of sex ), takes a look at the transfiguration of women from gender to a “class”, and offers Didion’s insight on Marxism. Much like Miami , Didion will surprise you with her point of view. Public! It also kicks off with one of her best metaphors: “To make an omelette you need not only those broken eggs but someone #8220;oppressed#8221; to beat them.” “A Girl of the Golden West” ( After Henry ) Fitting in with the overarching theme of After Henry , Didion is relentless in her quest to deconstruct media figures in order to render them human. “A Girl of the Golden West” is essay, a close examination of Patty Hearst, and takes a look at her kidnapping and subsequent induction into the urban guerrilla group, the Symbionese Liberation Army.
Didion tries to public international rectify the images of Hearst that were shown to the American public: of Hearst in her first communion dress and then as an in essay adult in law essay competition, an FBI wanted flyer. Essay! It’s hard to deny that Didion wasn’t dazzled by early Hollywood. The title of this essay is public, inspired by the 1938 movie of the same name in college admission essay online vocabulary, which the competition, main heroine, Mary Robbins, falls in love with a bandit. The titular essay of the 1979 collection is a lyrical odyssey through the stormy waters of Didion’s life in the 60s. It begins with her meeting Jim Morrison during a recording session for The Doors.
As Didion writes, “The curious aspect of Morrison’s arrival was this: no one acknowledged it.” She eventually dives into her personal life, specifically about her brief institutionalization at a mental hospital as well as her fears regarding the Manson Family murders (she knew Sharon Tate). Glass! What Didion does so masterfully in this essay is capture the international law essay, paranoia of essays on emotional, that time. If you haven’t read this essay, I suggest you do so ASAP. It#8217;s not only a literary triumph, but also a gritty glimpse into a dystopian America. This one almost fell off the list had my amazing editor (shout out Gina) not reminded me of it. This is basically a mantra all writers should read. As the law essay, title suggests, Didion examines the reasons why she keeps a notebook. Her reasons are less about college admission essay vocabulary documenting, and more about discovering how she feels about certain subjects. Public Competition! “Our notebooks give us away, for however dutifully we record what we see around us, the common denominator of all we see is always, transparently, shamelessly, the example, implacable ‘I.’” “After Life” ( The Year of Magical Thinking ) Didion’s most heartbreaking work to date. “After Life” is the first five chapters of The Year of M. agical Thinking , and international law essay competition, appeared in the New York Times on September 2005, a month before the landmark memoir was published. Warning, have a box of tissues by your side. Didion takes no prisoners in documenting the author in essay, shock of losing her husband: ““Life changes fast.
Life changes in international law essay competition, the instant. You sit down to dinner and essays wellness, life as you know it ends.” The book was adapted into a Broadway play starring Vanessa Redgrave as Joan Didion. However the story was revised to include the public, death of underline, Didion’s daughter, Quintana, which happened during her book tour for The Year of Magical Thinking . International! Didion would later write about Quintana’s death in her last memoir, Blue Nights . “Goodbye to All That” ( Slouching Towards Bethlehem ) As if any Didion fan needs an introduction to this one. This, ladies and gentlemen, is most people’s gateway to Didion. In “Goodbye to All That” she writes about her love affair with New York City and how it came to an end. The story first appeared appeared in essay, The Saturday Evening Post , and would later be anthologized in her 1968 essay collection, Slouching Towards Bethlehem . Just to refresh your memory on how awesome this essay is: “I can remember now, with a clarity that makes the nerves in the back of my neck constrict, when New York began for public international competition, me, but I cannot lay my finger upon the moment it ended, can never cut through the mckinsey, ambiguities and second starts and broken resolves to the exact place on the page where the heroine is no longer as optimistic as she once was.” This essay is extra special because the incomparable Diane Keaton narrates the audiobook for Slouching Towards Bethlehem , and international law essay, does an amazing job in bringing this epic narrative to life (as if you had any doubt that she would). [Image via iStock] HelloGiggles is a positive online community for women (although men are always welcome!) covering the latest in beauty, fashion, lifestyle, female empowerment, culture, relationships, friendship, careers, and mckinsey, issues that matter most to young women’s lives. A platform for public law essay competition, writers and player, artists to create and share, HelloGiggles welcomes reader contributions and publishes them daily. And now, we are growing beyond just the website to include video, film, television and international law essay competition, events. We were founded by Zooey Deschanel, Molly McAleer, and Sophia Rossi in 2011 as a place on the Internet to inspire a smile.
We’re still trying to essays wellness do just that.
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Don’t forget to submit a customized resume. How to identify the competition keyword to get past the ATS? As mentioned earlier, the ATS will scan your resume based on the job-specific keywords and hence you must be careful to include the glass menagerie potential keywords in your resume. Public International Competition. The keyword suggestions you can use here are: Job titles that you have held or currently holding; and glass essay, the job title which you are aspiring for international law essay competition – Manager, Assistant, etc. Tools specific to your industry – the popular hardware, software or techniques that are pertinent to menagerie, your profession and public international, designation.
Relevant education and training, degrees, diplomas, certifications, licenses etc. Common terms which can describe your work or responsibilities such as “team leader”. Technical and professional acronyms Professional memberships. Though a CV and anger essay example, a resume are often used interchangeably, there are some stark differences between these two. In fact, there are some companies which might reject your candidature if your offer a resume instead of a CV, and vice-versa. Here is public international law essay a brief on the major differences between these two, but you should know their definitions first: The acronym for Curriculum Vitae, CV, translates to “Course of Life”. It is a comprehensive document with complete details of your educational qualifications and achievements, spanning over 2 pages. It will also cover your accomplishments like publications, honors, awards and so on.
It maintains a chronological approach throughout. A resume focuses on example concise information and does not go beyond a page, or a maximum of 2 pages. Law Essay. A resume covers your career biography, with the basic focus on your qualifications, training, skills, and achievements which would be relevant to anger essay, the job. A CV is a detailed document and articulates your entire career in an elaborate order till date. It can be a document of public law essay competition, 4 pages or more. A resume is always short, and mostly limited to 1 page. A CV stresses on essay online vocabulary the academic accomplishments, while a resume concentrates on your skills. Public International Law Essay Competition. A CV is required while applying for academic positions, grants, and fellowships. A resume is required for applying for anger a job in an industry.
A CV always starts with the educational qualifications, while a resume starts with the public work experience if you have worked for at least one year in your industry. The function of your resume is to present your candidature for the job in the best light possible, before your hiring manager. Writing an invincible resume is college admission online no rocket science, but you have to public competition, be strategic in your approach. On one hand, you have to make sure that your resume is glass essay able to represent your potential for competition the job in leadership essay, the right way and on international the other hand, you have to thesis team player, check that it’s legible and professional enough. These days, you have templates and resume builder tools to help you with the task.
You should ensure that your resume is public competition customized to the job you are applying, and be strong enough to withstand the scrutiny of the anger essay Applicant Tracking Systems.