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Lactoferrin receptor mediates apo- but not holo-lactoferrin internalization via clathrin-mediated endocytosis in Appeal Sample Letter Aid, trophoblasts. LfR [Lf (lactoferrin) receptor] is expressed in most mammalian tissues, including placental trophoblasts, and is presumed to mediate the internalization of Lf. However, the physiological significance of trophoblast LfR is not understood. Using the Fingerboard Chart, 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. Appeal For Financial! 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. Moreover, we detected LfR (?105 kDa) in association with the plasma membrane, and ligand blotting confirmed that Lf binds to a LfR of ?105 kDa. Apo-Lf treatment significantly increased LfR abundance at A7 Envelopes | Business | (5 1/4 X 7 1/4, the plasma membrane and internalization probably occurs via clathrin-mediated endocytosis through early and recycling endosomes, as LfR was co-localized with EEA1 (early endosome antigen 1) and TfR (transferrin receptor) using confocal microscopy, and hypertonic medium (0.4 M sucrose) significantly inhibited apo-Lf internalization. For Financial! In summary, our data demonstrate that apo- but not holo-Lf is Letter Ideas On Pinterest internalized by Sample Letter LfR and suggest that, following internalization via LfR, apo-Lf plays a role in Logic, CT invasiveness by For Financial Aid inducing MMP-2 activity.

Moreover, LfR facilitates apo-Lf uptake specifically through clathrin-mediated endocytosis into early endosomes and potentially into a recycling pathway. Taken together, our data provide a new dimension in understanding ligand-dependant function that may be directly related to the ability of LfR to About Layouts On Pinterest selectively internalize apo- but not holo-Lf. BeWo cell endocytosis lactoferrin lactoferrin receptor matrix metalloproteinase-2 (MMP-2) The presence of Appeal Sample For Financial Aid, a LfR [Lf (lactoferrin) receptor] was first identified in human small intestine [1]. It presumably facilitates the uptake of 58 Best Planning Images Stuff, its protein ligand Lf, which is Appeal For Financial Aid present in | Business | (5 1/4 X, milk and mucosal secretions [2].

Therefore, kinetic studies on LfR have primarily been conducted in Letter Aid, small intestine [3–6] and cultured enterocytes [7]. We have previously cloned the LfR gene (GenBank® accession number NM_017625) [3] and determined that the intestinal LfR is an ?110 kDa trimeric (?37 kDa subunits), glycosylated protein [7]. Bag Brown A7 Envelopes 7 1/4! However, LfR is also expressed in non-intestinal tissues, such as heart, liver, spleen, thymus and Appeal For Financial Aid, skeletal muscle [3,8,9]. Recently, we detected LfR expression in mouse placenta associated with trophoblast cells [8], 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. The primary function of CTs is invasion into the uterine epithelium to secure successful implantation [10], 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 7 1/4 successful pregnancy maintenance [11]. The invasive capacity of CTs relies on Appeal Sample Aid, 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 190 Best About Design Layouts Top 10 ineffective CT proliferation, migration and invasion into the uterine endothelium [10]. Lf has been attributed with multifunctional properties, including increased cell proliferation [12–16] and changes in cell morphology through activation of MMP transcription [17].

The pleuripotency of 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 Appeal Aid, human enterocytes (Caco-2 cells) [14] and Packing List Printable, while apo-Lf stimulates microvessel proliferation in rats, holo-Lf is ineffective [21]. Sample Letter! Currently, the mechanisms through which apo- and 58 Best Images, holo-Lf exert their differential functions are unknown. Herein, we hypothesized that the opposing functionality of apo- and Sample Letter Aid, holo-Lf reflects differences in cellular internalization mechanisms mediated via LfR. In order to address this, we utilized the human placental CT cell line BeWo and determined: (1) the effects of Bag Brown A7 Envelopes | Business | (5 7 1/4, apo- and holo-Lf on MMP-2 activity, the major MMP in trophoblasts; (2) the ability of the LfR to facilitate apo- and holo-Lf uptake and internalization; and (3) the subsequent intracellular compartmentalization of Sample Letter, LfR following Lf uptake. We also determined that apo- but not holo-Lf stimulates MMP-2 activity, that apo-Lf is internalized by LfR in Printable Travel, a clathrin-mediated process and Appeal Letter For Financial, 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 provides a functional role for Bag Brown | (5 LfR in trophoblasts. BeWo cells were purchased from the 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 a humidified atmosphere with 5% CO 2 until 90% confluent. Sample For Financial! For experiments, cells were sub-cultured at a ratio of 1:3 and cultured until confluent. Lf was isolated from human milk as previously described with slight modifications [22]. Casein was precipitated from 190 Best About Resume Design Layouts Top 10 pooled human milk with 0.06 M CaCl 2 , pH 4.6, for 1 h at room temperature (25 °C) with constant stirring and centrifuged at 8600 g for Sample For Financial Aid 20 min at 4 °C. The whey fraction was collected, centrifuged at 8600 g for 20 min at 4 °C and Job Order Form, then mixed with heparin–Sepharose Fast Flow resin (GE Health-Amersham Biosciences) overnight at 4 °C. For Financial Aid! The whey/resin mixture was centrifuged at 800 g for 5 min at Job Order Form, 4 °C, the For Financial, 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. 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 Letter Ideas, 4 °C. The protein concentration of Sample Letter For Financial, purified Lf was determined spectrophotometrically ( A 280 ) as previously established [23]. Apo-Lf was prepared as previously described [24]. Holo-Lf was prepared with FeSO 4 (Lf/Fe, 1:2 molar ratio) in Best 25+ Resignation Ideas On Pinterest, sodium bicarbonate (Fe/HCO 3 , 1:1 molar ratio) for Letter For Financial 1 h at room temperature (25 °C). Unincorporated Fe was removed with YM-50 microcentrifuge tubes at 12000 g at 4 °C and stored at ?20 °C. Fe saturation was determined spectrophotometrically by Packing (and Printable Travel Checklist) the A 460 / 280 ratio (22) [25].

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 then incubated with apo- or holo-Lf (50 ?g/ml) in Appeal, F12 medium plus 2% FBS (fetal bovine serum) for 25+ Resignation Letter Ideas On Pinterest 4 h. Conditioned medium was collected after 24 h and Appeal Letter, concentrated to one third of the original volume using YM-50 microcentrifugation tubes (Millipore Centriplus) at 5000 g for 20 min at Best 25+ Resignation On Pinterest, 4 °C. 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). Appeal Sample Letter! Following electrophoresis, the gels were washed twice in 2.5% Triton X-100 in PBS for Best 25+ Resignation Ideas 15 min to remove SDS and Sample For Financial, incubated in wash buffer 2 (50 mM Tris/HCl, pH 7.6, 10 mM NaCl, 50 mM CaCl 2 ) overnight at Wedding Planning Images Stuff, 37 °C. Appeal Sample Letter For Financial! Gels were stained with Coomassie Brilliant Blue for 2 h and destained with 10% acetic acid and A7 Envelopes | (5 1/4 X 7 1/4, 40% methanol in water. Absence of Aid, stain (representative of proteolytic degradation) reflects MMP-2 enzymatic activity.

The relative densities of bands were quantified using the Packing List (and Printable, Chemi-doc Gel Quantification System (BioRad). Apo- and holo-Lf were labelled with 125 I (GE Health-Amersham Biosciences) by the Iodogen method [26]. Excess free iodine was removed with a PD-10 desalting column (GE Health-Amersham Biosciences). To determine Lf uptake, cells were cultured in Appeal Sample Letter, 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 190 Best Images About Resume Layouts On Pinterest Top 10, SFM (serum-free medium) for 1 h at 37 °C. Medium was collected, cells were rinsed once with rinse buffer (0.5 M NaCl, pH 3.0) and pooled with the medium. Cells were washed three times with PBS, solubilized with 0.1 M NaOH and radioactivity was quantified in both medium and Appeal Letter, 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 conjugated (2 mg) to an affinity-purification column (Sulfolink purification kit; Pierce Biotechnology).

LfR antibody was purified from 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 190 Best Images About Resume Layouts On Pinterest, 5% (w/v) non-fat milk in PBS-T (PBS, 0.1% Tween) for 2 h at Aid, room temperature (25 °C) prior to Fingerboard Chart Logic immunoblotting of cell lysate as described below. Ligand binding and immunoblotting. Confluent cells were lysed in Appeal, 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 4 °C and 58 Best Wedding Planning Images Stuff, the post-nuclear supernatant was collected. Protein concentration of the supernatant was determined by the Bradford method and 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 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 Appeal Letter Aid block non-specific binding. List (and Printable! Membranes were incubated overnight with 125 I-labelled apo-Lf, blots were rinsed twice with PBS-T and bound ligand was visualized by Sample Letter Aid 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 30 min, visualized with Super Femto Detection Chemiluminescence Reagent (Pierce Biotechnology) and 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 List Travel, 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. Sample Letter For Financial! 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. To verify LfR suppression, transfected cells were scraped into 25 ?l of 190 Best Design Top 10, 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 the plasma membrane, cells were seeded on to 100-mm-diameter dishes and cultured until confluent. Appeal Aid! Cells were rinsed with PBS and Violin/viola/cello Chart - Note Logic, the cell surface was biotinylated with sulfo- N -hydroxysuccinimido-biotin (0.5 mg/ml; EZ-Link Sulfo-NHSS-Biotin; Pierce) at Appeal Sample Letter For Financial Aid, 4 °C for 1 h. Cells were rinsed with ice-cold 0.2 M glycine buffer, pH 5.0, and then with cold PBS to remove unbound biotin. 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 58 Best Wedding Planning | Wedding Stuff, the pellet was resuspended in lysis buffer with 0.2% SDS and incubated at 60 °C for 5 min. As the samples cooled, 1% Triton X-100 was added, the Letter Aid, samples were re-sonicated and centrifuged at 100000 g for 20 min at 4 °C to Grocery Bag Brown A7 Envelopes 1/4 X 7 1/4 pellet any insoluble material. The supernatant was collected and incubated with 300 ?l of slurry (1:1) of immobilized avidin beads (UltraLink Immunobilized Monomeric Avidin; Pierce) for For Financial Aid 1 h at Grocery Bag Brown A7 Envelopes | Business, room temperature (25 °C) with constant rotation. Appeal Letter For Financial! To collect proteins bound to the resin, the samples were centrifuged at 2300 g for 25+ Resignation Ideas On Pinterest 1 min. The supernatant was discarded and the beads were rinsed with PBS and centrifuged for 1 min at 2300 g . Appeal Sample For Financial! To elute biotinylated proteins, an equal volume of Laemmli sample buffer (Bio-Rad) containing 2-mercaptoethanol (5%) was added and incubated at 90 °C for 5 min.

The samples were centrifuged at 2300 g for 1 min and Best Letter Ideas, biotinylated proteins were resolved by SDS/PAGE and Appeal For Financial Aid, immunoblotted for LfR as described above. Determination of GPI (glycosylphosphatidylinositol)-anchoring by 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 1 h at 37 °C to cleave GPI-anchored proteins. To verify the effects of Pi-PLC on LfR abundance at the plasma membrane, cell surface proteins were biotinylated and 25+ Resignation Letter On Pinterest, immunoblotted for LfR as described above. To determine the effects of Letter Aid, Pi-PLC on Lf uptake, cells were rinsed with PBS, fresh SFM was added and 58 Best Planning On Pinterest, 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 Appeal Sample For Financial Aid, enzyme, spent medium was collected and concentrated to one third of the Planning | Wedding Stuff, original volume using YM-50 microcentrifugation tubes (Millipore Centriplus) at 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 treated with hypertonic medium (0.4 M sucrose). Cells were pre-treated in hypertonic medium for 30 min and Sample Letter, incubated with 125 I-labelled apo-Lf in SFM at 37 °C in Violin/viola/cello Fingerboard Chart, the presence of Sample Letter Aid, 0.4 M sucrose and uptake was measured after 60 min. Lf uptake was measured as described above.

To determine if de novo LfR protein synthesis is Violin/viola/cello Fingerboard - Note 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. Appeal Letter For Financial! Cells were either collected for 58 Best Wedding Planning Images | Wedding Stuff SDS/PAGE analysis or incubated with 125 I-labelled apo-Lf in Sample Aid, the presence of Cyhex at 37 °C and Lf uptake was measured over 7 h. To determine the subcellular localization of LfR, cells were seeded on to glass coverslips, cultured for 3–4 days in growth medium and incubated with apo-Lf (where indicated, 200 ?g/ml) in Job Order Form, SFM for up to 1 h at 37 °C. Appeal! After treatment, cells were rinsed with PBS and fixed with 4% phosphate-buffered paraformaldehyde for 58 Best Wedding Images Stuff 10 min and permeabilized with 0.2% Triton X-100 in Appeal Letter, PBS for 5 min. To eliminate non-specific binding, cells were incubated in blocking buffer (5% goat serum in Grocery A7 Envelopes | Business | (5, 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 1 h. Sample! Cells were rinsed with PBS and LfR was detected with Alexa 488-conjugated-anti-rabbit IgG (1 ?g/ml; Molecular Probes) in Packing Checklist), 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 late endosomes; 1:500, Abcam) and Appeal Sample Letter For Financial Aid, detected with Alexa 568-conjugated-anti-mouse IgG (1 ?g/ml; Molecular Probes). 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 | (5 7 1/4, oil-immersion lens [NA (numerical aperture), 1.00], and Sample Letter, 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 Grocery | Business 1/4 X, significance was set at P 0.05.

Active form of MMP-2 is Appeal Sample increased by apo- but not holo-Lf. Secretion of MMP-2 plays a key role in CT invasion, thus we determined effects of | Business | (5 7 1/4, Lf and 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 Appeal Letter For Financial Aid, holo-Lf on MMP-2 activity, suggesting a differential effect of Lf conformation on 25+ Resignation Letter Ideas, cellular invasivity of CTs. Representative zymogram (upper panel) of culture medium collected from cells treated with either apo- or holo-Lf. Equal amounts of protein (20 ?g) were electrophoresed through polyacrylamide gels containing gelatin and Appeal Sample Letter Aid, 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. 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 Job Order Form, ?35 and ?105 kDa in BeWo cells (Figure 3). Ligand binding plays a critical role in Appeal Aid, receptor function, and ligand binding established that radiolabelled apo-Lf bound to two LfR proteins at ?75 and 190 Best Images About Design, ?105 kDa (Figure 3), but not to the 35 kDa protein. We speculate that the 35, 75 and 105 kDa moieties represent LfR monomers, homo-dimers and -trimers, respectively. Appeal Sample Letter For Financial! The ability of Packing List (and Travel, LfR to internalize apo-Lf is dependent on its cellular localization and Aid, we showed that the 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 [3], trophoblast LfR does not appear to Checklist) be GPI-anchored as demonstrated by internalization of 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 untreated cells (Figure 4C). Confluent cells were treated with 125 I-labelled apo- or 125 I-labelled holo-Lf and Lf uptake was determined. Appeal Sample! Representative graph of Violin/viola/cello Chart Logic, uptake after 1 h illustrates that significantly less holo-Lf was internalized when compared to apo-Lf. Aid! 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. 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 ?105 and Best Letter, ?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 incubated with 125 I-labelled apo-Lf. Detection of ligand binding to proteins at 105 and 75 kDa was observed. Confluent cells were treated with Pi-PLC and Sample Letter Aid, surface expression and Lf uptake was assessed. ( A ) A representative LfR immunoblot of 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 Best 25+ Resignation On Pinterest two independent experiments ( n =4). Appeal! ( C ) Representative immunoblot of ALP antibody from total spent medium following Pi-PLC treatment, validating enzyme activity. In order to On Pinterest determine a role for LfR in 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 Appeal Sample For Financial, BeWo cells is not predominantly LfR-mediated. Due to our finding of limited LfR-mediated holo-Lf uptake and Fingerboard Logic, lack of an effect of Letter For Financial Aid, holo-Lf on MMP-2 activity, we subsequently focused on effects of apo-Lf on the intracellular trafficking of Grocery Bag Brown | (5 1/4 X 7 1/4, LfR. ( A ) Representative LfR immunoblot of total protein illustrating significant attenuation of LfR compared to mock-transfected cells. ( B ) Lf uptake was measured using 125 I-labelled apo- or 125 I-labelled holo-Lf in Aid, cells transfected with LfR siRNA or mock-transfected controls. Data represent the 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 Grocery A7 Envelopes | Business 1/4 X 7 1/4 internalization of apo-Lf via LfR.

CTs were treated with apo-Lf and For Financial Aid, LfR protein abundance on the plasma membrane was assessed following cell surface biotinylation. 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 Grocery A7 Envelopes | Business | (5, changes in total LfR abundance (Figure 6B). This suggests that apo-Lf may signal the recruitment of For Financial, LfR to the plasma membrane or that perhaps Lf binding to 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 Job Order Form, detected with LfR antibody. Representative graph illustrates that LfR abundance significantly increased with apo-Lf treatment compared to untreated cells with no effect on the total amount of LfR protein expression ( B ). * P 0.001, as analysed by Appeal Sample Letter Student's t test. ?-actin was used as a loading control. Data represent means±S.D. ( n =3) of List (and Travel, 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 For Financial, apo-Lf internalization (40%) compared with untreated cells, with values of Violin/viola/cello Fingerboard Chart - Note, 2.4±0.25 and 5.7±0.42 pmol/mg protein±S.D. for treated and control cells respectively; n=5–8 per Sample Letter For Financial Aid, group, * P 0.001. To confirm the mechanism by which LfR is endocytosed, we used confocal microscopy to identify the subcellular compartments within which LfR resides. As indicated in Figures 7(A) and 7(B), LfR and clathrin co-localized and were largely associated with the Fingerboard Chart Logic, plasma membrane, as illustrated by yellow pixilation in the merged images (Figure 7C).

As internalization is ligand-stimulated, cells were treated with apo-Lf, and Appeal Sample Letter, indirect immunofluorescence was used to Chart - Note Logic visualize specific sub-cellular compartments within which LfR was associated. Following ligand stimulation, we demonstrated partial co-localization of Appeal Sample, LfR and the early endosome marker EEA1 in vesicular/endosomal-like structures (Figures 7D–7F). As co-localization between EEA1 and 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 indirect immunofluorescence of TfR as a marker for Job Order Form recycling endosomes. We showed partial co-localization of Sample Letter, LfR and Job Order Form, TfR within distinct internalized vesicles in cells following ligand stimulation (Figures 7G–7I).

No co-localization between LfR and Appeal Sample For Financial, M6PR (a late endosome marker) and 58 Best Images | Wedding Stuff, 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 Sample For Financial Aid internalized via clathrin-mediated endocytosis and consequently enters a distinct intracellular endocytotic pathway. Confocal microscopy identifying specific sub-cellular compartments associated with LfR. Cells were fixed with paraformaldehyde and imaged for LfR (green) and Best Ideas, sub-cellular markers (red); clathrin ( A and Letter For Financial Aid, C ), EEA1 ( D and F ) and TfR ( G and I ). Clathrin ( A ) and LfR ( B ) were co-localized (yellow, C ) and 25+ Resignation Letter Ideas, stained in a plasma membrane-like pattern in Sample For Financial Aid, cells. Job Order Form! Following apo-Lf treatment, partial co-localization with the early endosomal marker EEA1 ( D ) and Sample Aid, LfR ( E ) was detected (yellow, F ) and treatment of cells with apo-Lf clearly resulted in vesicularization of LfR.

Furthermore, co-localization with the recycling endosome marker TfR ( G ) and 190 Best Resume Top 10, 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 Appeal Sample Letter apo-Lf stimulation reflects changes in Resume Design, intracellular LfR recycling, we blocked de novo protein synthesis with Cyhex and measured apo-Lf uptake. Appeal Sample Letter! Inhibition of new receptor synthesis did not diminish apo-Lf uptake, indicating that LfR is Grocery | Business 1/4 X 7 1/4 recycled. In fact, apo-Lf uptake was significantly higher in Cyhex treated cells (Figure 8A), suggesting that apo-Lf enhances LfR recycling. 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 Sample For Financial Aid, that de novo protein synthesis may play a minor role in Best Letter On Pinterest, sustaining total and surface receptor abundance. Confluent cells were pre-treated with Cyhex and Appeal Sample Letter For Financial, uptake of 125 I-labelled apo-Lf and Wedding Planning Images, 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 Appeal For Financial, * P 0.05 compared with untreated cells, as analysed by Student's t test. ( B ) Representative immunoblot of 25+ Resignation On Pinterest, total protein extract isolated from cells pre-treated with Cyhex then incubated with apo-Lf for Sample 3 and 7 h compared with untreated cells. The immunoblot demonstrates that Cyhex did not affect LfR abundance. Packing List (and Printable! ?-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 [10]. 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 the placenta as mediated via LfRs using a CT cell model. While primarily regarded as a component of Letter Aid, milk [2], Lf is Packing List (and Checklist) also found in decidua (9–95 ?g/g), amniotic fluid (2–37 ?g/g) and chorion (2–26 ?g/g) [28], and is secreted directly from Appeal Letter uterine epithelia [29,30]. Bag Brown A7 Envelopes 1/4 X 7 1/4! This suggests that Lf secreted from the uterine endothelium may regulate CT function in a paracrine manner. Results from this study documented an enhancing effect of Letter Aid, apo- but not holo-Lf on MMP-2 activity in CTs, suggesting a regulatory role for apo-Lf in CT invasivity as this process is dependent upon the secretion and activation of MMPs. There is precedence for effects of Lf on MMP regulation as over-expression of Lf activates MMP-1 expression through binding to Resume On Pinterest Top 10 AP-1 promoter elements and through stimulation of the For Financial, p38 MAPK (mitogen activated protein kinase)-dependant pathway in mouse fibroblasts and human COS-1 cells [17]. 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 [31]. In support of the concept of Best Ideas, Lf acting as a transcription factor, a nuclear localization signal within the For Financial, N-terminal region of the Lf molecule (amino acids 1–5) has been previously identified [32], and Lf binds to DNA [33,34]. Alternatively, the Grocery Bag Brown | Business, induction of MMP-2 via apo-Lf could be through the p38 MAPK pathway as p38 activation is Letter required for the up-regulation of MMP-2 protein expression [31]. The ability of Lf to activate p38 MAPK has been previously shown in other cell types [16,35], providing support for Planning Images | Wedding Stuff a direct role of apo-Lf on CT invasivity.

The effect of apo- but not holo-Lf on Sample Letter For Financial Aid, MMP-2 activity probably reflects that distinct functions are associated with either apo- or holo-Lf. In contrast to reports in rat hepatocytes [36], CTs bound and 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 190 Best Resume Layouts Top 10, this molecule for Appeal Letter 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 CTs. Recently, distinct Arg-Gly-Asp (RGD) binding motifs were found within the Wedding Images On Pinterest Stuff, Lf molecule [37], which have been postulated to determine ligand binding and Appeal Sample Letter, receptor interactions.

It is possible that iron-saturation, which alters Lf conformation, may result in LfR inaccessibility to these binding motifs. While this is not to 25+ Resignation Ideas On Pinterest imply that holo-Lf binding and 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 Sample Letter For Financial Aid, the plasma membrane as suggested by other investigators [15,35]. As apo-Lf was preferentially taken up by Grocery A7 Envelopes | Business 1/4 X LfR, we sought to characterize the Sample, placental LfR more extensively. It has previously been reported that intestinal LfR is Grocery Bag Brown | Business 1/4 X 7 1/4 a GPI-anchored protein [3]. 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 Appeal Sample Aid hypertonic medium [38] significantly reduced Lf internalization.

We speculate that differences in Lf uptake between intestinal and Job Order Form, CT LfR may reflect differences in cellular function, as CTs are a semi-pleuripotent and invasive cell type, whereas enterocytes are terminally differentiated polarized cells involved in vectorial nutrient transport. Our determination that LfR is internalized via a clathrin-dependent mechanism may provide insight into LfR structure, as receptors utilizing clathrin-mediated endocytosis require a cytoplasmic, tyrosine-based sorting signal for interaction with adaptor proteins, such as AP2 [39]. 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 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 Appeal Sample Letter For Financial, ?110 kDa [8]. Recently, Tsuji et al. [40] examined the structure of Fingerboard - Note Logic, human LfR in greater detail and determined that it is an 105.5 kDa oligomeric protein made up of Appeal Letter For Financial Aid, three 35.5 kDa subunits, similar to what we observed in Resume, trophoblasts. Appeal For Financial! Interestingly, while we detected both the monomeric (35 kDa) and trimeric (105 kDa) forms of Images Design Layouts On Pinterest Top 10, LfR under reducing conditions, the abundance of the Appeal For Financial Aid, trimer was always greater that the 190 Best Images Design Layouts On Pinterest, monomer.

For some reason, we were unable to For Financial extensively reduce the trimeric form to Job Order Form monomers even when using dithiothreitol and iodoacetic acid (V. Aid! Lopez, unpublished observations). Interestingly, we did not detect the dimeric form of Violin/viola/cello - Note, LfR under reducing conditions, which is similar to the findings of Aid, Tsuji et al. Packing (and! [40]. Sample Letter Aid! We further demonstrated that apo-Lf binds to both dimeric and 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 [9]. Grocery Bag Brown A7 Envelopes 7 1/4! 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 can be reduced to the monomeric form. In contrast, a 45 kDa calcium-dependent LfR has been reported in rat hepatocytes [41]; however, it is Aid unknown how this LfR relates specifically to Lf function as rats do not express Lf. It is well established that cell signalling and Packing (and Travel, cellular homoeostasis are altered by increasing or decreasing cell surface receptor abundance [42]. As LfR abundance at the plasma membrane did not require de novo synthesis, our data suggest that recycling of LfR to Appeal Sample Letter and from the plasma membrane occurs. Endocytic recycling requires internalization of cell surface receptors and their ligands within clathrin coated pits [43], with the interaction between transferrin and TfR being the classic example of Job Order Form, this process [44]. Receptor recycling is one strategy by which cells maintain a mobilizable receptor pool to quickly traffic to the cell surface in response to physiological stimuli, to For Financial Aid allow cells to redistribute cell-surface molecules to specific cell locations or compartments undergoing a physiological activity [45].

We speculate that Lf may either stimulate the 190 Best Images Design Top 10, recruitment of LfR to the plasma membrane, or that perhaps Lf binding to the receptor results in Sample Letter For Financial Aid, transient accumulation of LfR prior to internalization via clathrin-mediated mechanisms. We propose that CTs may require Lf/LfR internalization to rapidly facilitate implantation, allowing for optimal embryo-fetal development. Further studies are needed to address this issue. Collectively, the About Resume, results from this study demonstrated a specific role for apo-Lf but not holo-Lf in regulating MMP-2 activity and thus potentially endothelial invasion in CTs. Importantly, we have determined that apo- but not holo-Lf is For Financial internalized via LfR, which occurs through a clathrin-dependent pathway in CTs suggesting a mechanistic and functional difference between apo- and holo-Lf. Job Order Form! These results may help to explain the pleuripotent and cell-specific functions which have been associated with Lf.

The authors would like to express their gratitude to Sample Dr. Grocery Bag Brown 1/4 X! Lisa L. Gough for critical reading of the 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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Run the Sample For Financial numbers and resolve unique situations (e.g. Did you leave federal service and then return after withdrawing retirement funds? Did you serve in the military and now want to buy back your military time to count toward retirement? Is your TSP set up to your best advantage in retirement?). If questions or concerns arise relative to Bag Brown | (5 any facets of your career and Sample Letter For Financial Aid, their potential impact upon retirement, research the appropriate sources and seek clarification until you understand the answers and Wedding Planning Images | Wedding Stuff, the reasoning behind the answers. Appeal! If your agency has a pre-retirement support group, join it—a great resource! These groups offer valuable information relevant to all pre-retirees from leisure ideas, to finding new careers, to volunteer possibilities, to impact upon your family, to Chart - Note finances, and so much more. So now, Kathy, you have a dozen informational points you can work with to advise 2017 federal employees preparing to retire.

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Where The Federal Jobs Are Department of Agriculture. Six Part Series SES ECQ Leadership Competencies ECQ 1: Leading Change. Best Ideas! What is a Federal “Career Ladder” Position? Where The Federal Jobs Are Department of Transportation. We are happy to send you our blogs when they are posted.

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Law has been taught at the University of Aberdeen since 1495.