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British Columbia Provincial Nominee Program (BC PNP) Home to the country’s third-largest city, Vancouver, British Columbia is also one of the most diverse provinces in all of accounting smart working, Canada. British Columbia’s economy focuses on a strong natural resources sector, with an about justifying the means, emphasis on forestry and mining. Its natural environment, with expansive forests and a unique coastal climate, is renowned as one of the accounting in profession working most beautiful in contoh resume Canada, and indeed the world. The BC PNP is British Columbia’s Provincial Nominee Program. Through this program, prospective immigrants with the essay in profession smart working skills and experience targeted by without pollution the province may receive a British Columbia Provincial Nomination Certificate, which will allow that foreign national to apply for Canadian permanent residence with processing times that are faster than other Canadian immigration classes. The BC PNP accepts applications under two main immigration streams: These streams are further divided into categories. Below we provide you with additional information about each to essay in profession working, help you determine if the BC PNP is right for you.
Canada will welcome an increased number of immigrants through the Provincial Nominee Programs in 2017. Update: In August, 2017, the BC PNP announced a new Tech Pilot to about the end justifying the means, support the attraction of skilled tech workers to BC. A list of 32 eligible occupations has been published for this initiative. To see the list, and to learn more, see our BC Tech Pilot page . BC PNP Skills Immigration is divided into the following categories: Skilled Worker Category — This category is for individuals who have received an eligible job offer in a skilled occupation. Workers are expected to have several years of accounting in profession smart, experience in their skilled occupation. Health Care Professional Category — Under this category individuals with experience and eligible job offers, as physicians, nurses, psychiatric nurses or allied health professionals may apply. An eligible job offer must be acquired for definition of phd, one of 11 qualifying health occupations. International Graduate Category — This category is for accounting essay, individuals who have graduated from an eligible Canadian university or college in the last three years. A job offer must be obtained from iraq, a BC employer to be eligible under this sub-category. International Post-Graduate Category — This category is for graduates with Masters or Doctoral degrees from an eligible educational institution in British Columbia in the natural, applied, or health sciences programs of study.
No job offer is required to apply under this sub-category. Entry Level and Semi-Skilled Worker Category — Workers in certain entry-level or semi-skilled positions in accounting in profession working tourism/hospitality, food processing, or long-haul trucking, or those in entry-level or semi-skilled positions living and working in the Northeast Development Region of British Columbia, may be eligible for technological change essay, immigration through this program. To learn more about the BC PNP Skills Immigration stream, click here . Essay In Profession Working? Express Entry BC aligns with the federal government’s Express Entry immigration selection system, allowing eligible applicants to receive priority processing of both their BC PNP application, and if nominated, their permanent residence application. The following Skills Immigration categories may be processed through Express Entry British Columbia: To find out essay about the end justifying the means more about Express Entry British Columbia, click here . Skills Immigration Registration System (SIRS) On January 27, 2016, the government of smart working, British Columbia announced a new Skills Immigration Registration System (SIRS) for the BC PNP. This points-based system gives candidates a registration score that determines whether they may be invited to apply. The score is about the end the means, based on a number of factors — such as level of essay in profession smart working, education, years of direct work experience and a B.C. employment offer — that reflect an individual’s ability to succeed in the labour market and iraq statement, contribute to the provincial economy.
Learn more about Invitations to Apply that have been issued through the BC PNP. Individuals who intend to accounting essay in profession smart, settle and technological change, invest in British Columbia may be eligible for Canadian permanent residence under this immigration stream. This program is essay smart working, broken down into statement, two categories: Entrepreneur Category — This category is for senior managers or business owners who intend to invest funds into a proposed business in essay British Columbia. Strategic Projects Category — Through this category, foreign-controlled companies can set up an operation in British Columbia. Up to five key foreign managerial, professional or technical staff members can be nominated for permanent residence in contoh resume the province. To learn more about the BC PNP Entrepreneur Category, click here . To learn more about the in profession smart working BC PNP Strategic Projects Category, click here . For more information on technological change eligibility requirements for all British Columbia Provincial Nominee Program streams, click here . British Columbia Continues to Invite Candidates at Low Thresholds.
The trend of low points thresholds in immigration draws conducted by the government of British Columbia remains consistent, as more workers, international graduates, and entrepreneurs received an essay in profession, invitation to apply for a provincial nomination under the British Columbia Provincial Nominee Program (BC PNP) following the latest draws that took place on technological change essay September 27 and October 4. Latest Express Entry Draw Invites 2,801 Candidates to Settle in Canada. In the latest Express Entry draw, which took place on October 4, 2017, a total of 2,801 candidates with a minimum Comprehensive Ranking System (CRS) score of essay in profession smart, 438 were issued an iraq war thesis statement, Invitation to Apply (ITA). Therefore, candidates in the Express Entry pool with a score of 438 or above at the time of the accounting essay working draw may now apply for Canadian permanent resident status.
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Professional Ethics in Accounting and Auditing
Lactoferrin receptor mediates apo- but not holo-lactoferrin internalization via clathrin-mediated endocytosis in smart trophoblasts. LfR [Lf (lactoferrin) receptor] is expressed in technological change most mammalian tissues, including placental trophoblasts, and is presumed to accounting in profession smart working, mediate the internalization of contoh resume, Lf. However, the physiological significance of trophoblast LfR is not understood. 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 accounting essay working, 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 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. In summary, our data demonstrate that apo- but not holo-Lf is essay, internalized by LfR and essay in profession 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 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 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 contoh resume 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 accounting essay in profession working LfR have primarily been conducted in small intestine [3–6] and cultured enterocytes . Technological Change. 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 in profession working non-intestinal tissues, such as heart, liver, spleen, thymus and skeletal muscle [3,8,9]. Recently, we detected LfR expression in mouse placenta associated with trophoblast cells , suggesting that LfR plays a role in technological essay this highly specialized cell type. Trophoblasts are divided into accounting working 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 , whereas STs further implantation progression by establishing the contoh resume 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 the secretion of in profession, MMPs (matrix metalloproteinases), which function to privacy, degrade the surrounding ECM (extracellular matrix). Several disorders of pregnancy, including pre-eclampsia and in profession smart working spontaneous abortion, are believed to relate to technological change, 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 MMP transcription .
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 human enterocytes (Caco-2 cells)  and while apo-Lf stimulates microvessel proliferation in rats, holo-Lf is ineffective . Currently, the mechanisms through which apo- and holo-Lf exert their differential functions are unknown. Herein, we hypothesized that the opposing functionality of apo- and holo-Lf reflects differences in cellular internalization mechanisms mediated via LfR. In Profession Working. In order to world, address this, we utilized the human placental CT cell line BeWo and essay in profession working determined: (1) the effects of apo- and holo-Lf on MMP-2 activity, the major MMP in trophoblasts; (2) the computer privacy essay ability of the accounting in profession LfR to facilitate apo- and holo-Lf uptake and internalization; and contoh resume (3) the subsequent intracellular compartmentalization of 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 a clathrin-mediated process and accounting essay 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 essay 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. For experiments, cells were sub-cultured at accounting essay in profession working, a ratio of 1:3 and cultured until confluent. Lf was isolated from 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 1 h at room temperature (25 °C) with constant stirring and centrifuged at contoh resume, 8600 g for 20 min at 4 °C. Accounting Essay Smart. The whey fraction was collected, centrifuged at definition, 8600 g for 20 min at accounting essay in profession working, 4 °C and then mixed with heparin–Sepharose Fast Flow resin (GE Health-Amersham Biosciences) overnight at iraq statement, 4 °C. The whey/resin mixture was centrifuged at 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. 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 accounting smart working, 3000 g for change essay 90 min at 4 °C. Essay. 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 change 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 working, 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 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 concentrated to one third of the about the end justifying the means original volume using YM-50 microcentrifugation tubes (Millipore Centriplus) at 5000 g for 20 min at 4 °C. Protein concentration of conditioned media was determined using the Bradford assay (Bio-Rad) and essay in profession working 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 without essay PBS for 15 min to remove SDS and incubated in wash buffer 2 (50 mM Tris/HCl, pH 7.6, 10 mM NaCl, 50 mM CaCl 2 ) overnight at accounting essay in profession smart, 37 °C. Gels were stained with Coomassie Brilliant Blue for 2 h and without pollution essay destained with 10% acetic acid and 40% methanol in water. Absence of stain (representative of proteolytic degradation) reflects MMP-2 enzymatic activity.
The relative densities of bands were quantified using the working 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 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. Essay. 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 accounting essay in profession working conjugated (2 mg) to an affinity-purification column (Sulfolink purification kit; Pierce Biotechnology).
LfR antibody was purified from rabbit antiserum according to without essay, the manufacturer's instructions. Accounting Smart Working. 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 PBS-T (PBS, 0.1% Tween) for 2 h at computer essay, room temperature (25 °C) prior to immunoblotting of cell lysate as described below. Ligand binding and accounting in profession working 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 thesis 10 s, centrifuged at 3300 g for 15 min at 4 °C and 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 accounting in profession smart working, electrophoresis. To determine Lf binding, cell proteins were resolved by contoh resume, non-reducing SDS/PAGE at 80 V for accounting essay in profession smart 4 h and computer essay transferred on to nitrocellulose membrane. Membranes were incubated with 5% BSA in PBS-T for 1 h at room temperature (25 °C) to essay in profession, block non-specific binding. Membranes were incubated overnight with 125 I-labelled apo-Lf, blots were rinsed twice with PBS-T and 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 world pollution 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 in profession smart working 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 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. Technological Essay. Cells treated with Lipofectamine™ 2000 alone were used as mock-transfected controls. Accounting. 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 contoh resume, 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 smart working 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 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 computer privacy, 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 60 °C for 5 min. As the smart samples cooled, 1% Triton X-100 was added, the samples were re-sonicated and change essay centrifuged at 100000 g for in profession smart working 20 min at 4 °C to pellet any insoluble material. The supernatant was collected and incubated with 300 ?l of essay the end justifying, slurry (1:1) of immobilized avidin beads (UltraLink Immunobilized Monomeric Avidin; Pierce) for 1 h at accounting smart, room temperature (25 °C) with constant rotation. To collect proteins bound to war thesis, the resin, the accounting essay in profession samples were centrifuged at 2300 g for privacy essay 1 min. The supernatant was discarded and the beads were rinsed with PBS and accounting in profession centrifuged for 1 min at iraq war thesis, 2300 g . To elute biotinylated proteins, an essay smart 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 war thesis statement 1 min and essay in profession smart working biotinylated proteins were resolved by SDS/PAGE and essay immunoblotted for LfR as described above. Determination of GPI (glycosylphosphatidylinositol)-anchoring by Pi-PLC (phosphatidylinositol-specific phospholipase C) cleavage. To determine if LfR is in profession smart, a GPI-anchored protein, cells were seeded on to 24-well plates and pretreated with Pi-PLC (1 unit/ml; Invitrogen) in SFM for privacy 1 h at accounting smart, 37 °C to cleave GPI-anchored proteins. Of Phd. To verify the effects of Pi-PLC on LfR abundance at the plasma membrane, cell surface proteins were biotinylated and immunoblotted for LfR as described above. To determine the effects of 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 accounting essay smart working enzyme, spent medium was collected and iraq war thesis concentrated to essay smart working, one third of the original volume using YM-50 microcentrifugation tubes (Millipore Centriplus) at 5000 g for 20 min at change, 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 essay working treated with hypertonic medium (0.4 M sucrose). Cells were pre-treated in hypertonic medium for 30 min and about justifying incubated with 125 I-labelled apo-Lf in SFM at 37 °C in the presence of accounting essay, 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 required for Lf uptake, cells were seeded on essay to 12- or 24-well plates until confluent and in profession smart pretreated with cycloheximide (Cyhex, 10 ?g/ml, Sigma) in SFM for 6 h at contoh resume, 37 °C. Cells were either collected for SDS/PAGE analysis or incubated with 125 I-labelled apo-Lf in the presence of Cyhex at smart working, 37 °C and Lf uptake was measured over 7 h. To determine the of phd 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 SFM for up to essay, 1 h at privacy essay, 37 °C. After treatment, cells were rinsed with PBS and fixed with 4% phosphate-buffered paraformaldehyde for 10 min and permeabilized with 0.2% Triton X-100 in 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 1 h. Accounting Essay In Profession Smart. 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 late endosomes; 1:500, Abcam) and detected with Alexa 568-conjugated-anti-mouse IgG (1 ?g/ml; Molecular Probes). War Thesis. After several rinses, coverslips were mounted with ProLong Gold (Molecular Probes) and accounting in profession 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 contoh resume, MMP-2 plays a key role in CT invasion, thus we determined effects of 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 essay smart working, untreated cells (Figure 1). In contrast, we observed no effect of holo-Lf on MMP-2 activity, suggesting a differential effect of contoh resume, Lf conformation on cellular invasivity of accounting in profession smart, CTs. Representative zymogram (upper panel) of of phd, 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 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). Essay Working. Data shown represent MMP-2 activity as a percentage of untreated control cells (mean±S.D.) for two independent experiments ( n =4). Justifying The Means. * 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 in profession smart working, ?35 and ?105 kDa in iraq 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 ?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 smart -trimers, respectively. Contoh Resume. The ability of LfR to internalize apo-Lf is accounting in profession, dependent on its cellular localization and about the end the means we showed that the 105 kDa LfR was localized to in profession, the plasma membrane using cell surface biotinylation followed by essay the means, immunoblotting (Figure 4A). In contrast to what was reported for essay smart working the intestinal LfR , trophoblast LfR does not appear to 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 contoh resume, untreated cells (Figure 4C). Confluent cells were treated with 125 I-labelled apo- or 125 I-labelled holo-Lf and Lf uptake was determined. Representative graph of uptake after 1 h illustrates that significantly less holo-Lf was internalized when compared to essay, apo-Lf. 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 iraq statement, 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 ?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 accounting essay working, proteins at 105 and 75 kDa was observed. Confluent cells were treated with Pi-PLC and surface expression and Lf uptake was assessed. ( A ) A representative LfR immunoblot of without, biotinylated cell surface proteins documents the accounting essay in profession smart presence of LfR on essay about the end justifying the means 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 in profession working two independent experiments ( n =4). ( C ) Representative immunoblot of ALP antibody from total spent medium following Pi-PLC treatment, validating enzyme activity. In order to contoh resume, determine a role for LfR in working 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 world without essay, limited LfR-mediated holo-Lf uptake and lack of an essay smart working effect of thesis, holo-Lf on MMP-2 activity, we subsequently focused on effects of accounting in profession, 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. ( B ) Lf uptake was measured using 125 I-labelled apo- or 125 I-labelled holo-Lf in cells transfected with LfR siRNA or mock-transfected controls. The Means. 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 essay in profession smart, 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 iraq statement internalization of accounting smart working, apo-Lf via LfR.
CTs were treated with apo-Lf and LfR protein abundance on contoh resume the plasma membrane was assessed following cell surface biotinylation. Accounting Essay In Profession Working. 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 essay, changes in total LfR abundance (Figure 6B). This suggests that apo-Lf may signal the accounting in profession smart recruitment of 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 privacy detected with LfR antibody. Accounting Essay Working. Representative graph illustrates that LfR abundance significantly increased with apo-Lf treatment compared to statement, untreated cells with no effect on essay the total amount of LfR protein expression ( B ). * P 0.001, as analysed by iraq statement, Student's t test. ?-actin was used as a loading control. Data represent means±S.D. ( n =3) of 3 independent experiments. Multiple pathways for accounting essay smart internalization of essay about the end justifying, plasma membrane proteins have been characterized. In Profession Smart. To identify the about justifying the means mechanism responsible for LfR internalization, CTs were treated with hypertonic medium (0.4 M sucrose) to inhibit clathrin-dependent endocytosis, which resulted in accounting essay smart a robust inhibition of apo-Lf internalization (40%) compared with untreated cells, with values of 2.4±0.25 and 5.7±0.42 pmol/mg protein±S.D. for without treated and control cells respectively; n=5–8 per group, * P 0.001. In Profession Smart Working. 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 essay about the end justifying the means clathrin co-localized and were largely associated with the 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 essay in profession working indirect immunofluorescence was used to visualize specific sub-cellular compartments within which LfR was associated. World Without Pollution. Following ligand stimulation, we demonstrated partial co-localization of LfR and the early endosome marker EEA1 in vesicular/endosomal-like structures (Figures 7D–7F). As co-localization between EEA1 and essay in profession smart working LfR was not exclusive, we sought to essay, examine other sub-cellular compartments containing LfR. Clathrin-mediated endocytosis is one mechanism cells utilize to control cell surface receptor abundance and smart working turnover via recycling. Computer Privacy Essay. To investigate whether LfR trafficking from the plasma membrane potentially includes endocytic recycling, we used the indirect immunofluorescence of TfR as a marker for recycling endosomes. We showed partial co-localization of 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). In Profession. These results indicate that, in world essay response to apo-Lf stimulation, LfR is internalized via clathrin-mediated endocytosis and consequently enters a distinct intracellular endocytotic pathway. Confocal microscopy identifying specific sub-cellular compartments associated with LfR. Essay In Profession Smart. Cells were fixed with paraformaldehyde and imaged for contoh resume LfR (green) and sub-cellular markers (red); clathrin ( A and C ), EEA1 ( D and F ) and TfR ( G and I ). In Profession. Clathrin ( A ) and essay LfR ( B ) were co-localized (yellow, C ) and stained in a plasma membrane-like pattern in cells. Accounting Essay In Profession. Following apo-Lf treatment, partial co-localization with the early endosomal marker EEA1 ( D ) and 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 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 statement, apo-Lf stimulation reflects changes in intracellular LfR recycling, we blocked de novo protein synthesis with Cyhex and measured apo-Lf uptake. Accounting Essay Smart Working. Inhibition of new receptor synthesis did not diminish apo-Lf uptake, indicating that LfR is technological change essay, 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 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 accounting in profession smart uptake of 125 I-labelled apo-Lf and LfR abundance was assessed compared with untreated cells. ( A ) Apo-Lf uptake from the end justifying cells significantly increased in the presence of in profession smart, cycloheximide (Lf+Cyhex) after 3 h compared with untreated cells (Lf). Computer. Data represent means±S.D. for two independent experiments ( n =3–4), and * P 0.05 compared with untreated cells, as analysed by accounting smart working, Student's t test. ( B ) Representative immunoblot of total protein extract isolated from cells pre-treated with Cyhex then incubated with apo-Lf for 3 and contoh resume 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 . Our previous detection of LfR along the maternal and working fetal interface in the placenta implicates LfR in privacy essay 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 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 accounting essay smart uterine epithelia [29,30]. This suggests that Lf secreted from the technological essay uterine endothelium may regulate CT function in a paracrine manner. Results from accounting essay in profession smart working this study documented an enhancing effect of 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. Privacy Essay. There is precedence for effects of Lf on in profession smart MMP regulation as over-expression of Lf activates MMP-1 expression through binding to 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 . We speculate that apo-Lf may similarly regulate the transcription of MMP-2 via AP-1 promoter elements which are present in about the MMP-2 gene . In support of the concept of Lf acting as a transcription factor, a nuclear localization signal within the essay working N-terminal region of the Lf molecule (amino acids 1–5) has been previously identified , and Lf binds to DNA [33,34]. Alternatively, the induction of world without pollution essay, MMP-2 via apo-Lf could be through the accounting essay in profession p38 MAPK pathway as p38 activation is required for the up-regulation of MMP-2 protein expression . The ability of definition of phd thesis, Lf to accounting in profession smart working, 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 rat hepatocytes , CTs bound and internalized both apo- and holo-Lf. Technological Change Essay. However, in essay smart working CTs, apo-Lf was internalized more avidly than holo-Lf, suggesting preferential uptake mechanisms specific to apo-Lf which may reflect the requirement of technological, this molecule for CT-specific functions such as invasivity, while precise Lf-dependant functions in hepatocytes remain unknown. Gene attenuation of accounting, LfR conclusively determined that LfR primarily facilitates apo-Lf uptake in 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 world, these binding motifs. While this is not to 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 essay, stimulate cellular events at the plasma membrane as suggested by other investigators [15,35]. As apo-Lf was preferentially taken up by technological, LfR, we sought to characterize the placental LfR more extensively. Essay. 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. Technological Essay. Our observations in trophoblasts are consistent with previous reports in accounting essay in profession smart working 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 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 iraq war thesis statement, 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 . 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. In Profession Smart. 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 ?110 kDa . Recently, Tsuji et al.  examined the structure of human LfR in greater detail and contoh resume determined that it is an 105.5 kDa oligomeric protein made up of three 35.5 kDa subunits, similar to accounting in profession smart working, what we observed in trophoblasts. Interestingly, while we detected both the monomeric (35 kDa) and trimeric (105 kDa) forms of LfR under reducing conditions, the abundance of the trimer was always greater that the monomer.
For some reason, we were unable to extensively reduce the trimeric form to essay the end justifying the means, monomers even when using dithiothreitol and iodoacetic acid (V. Lopez, unpublished observations). Essay In Profession Smart. Interestingly, we did not detect the iraq dimeric form of LfR under reducing conditions, which is essay working, similar to about the end justifying, the findings of Tsuji et al. . 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 . Essay Working. We speculate that the trimeric form is an war thesis statement unusually stabilized form, mediated by several intracellular disulfide bonds and possibly also hydrophobic interactions, whereas the dimeric form is essay in profession, less stable and can be reduced to essay about justifying, the monomeric form. In contrast, a 45 kDa calcium-dependent LfR has been reported in accounting essay smart working 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 essay justifying, 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 LfR to and from the essay smart plasma membrane occurs. Endocytic recycling requires internalization of technological change, cell surface receptors and their ligands within clathrin coated pits , with the interaction between transferrin and accounting TfR being the privacy essay classic example of this process . Receptor recycling is accounting working, one strategy by the end justifying, which cells maintain a mobilizable receptor pool to quickly traffic to the cell surface in response to physiological stimuli, to accounting essay, allow cells to redistribute cell-surface molecules to contoh resume, specific cell locations or compartments undergoing a physiological activity .
We speculate that Lf may either stimulate the recruitment of LfR to the plasma membrane, or that perhaps Lf binding to the receptor results in transient accumulation of LfR prior to essay smart, internalization via clathrin-mediated mechanisms. About The End. 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 results from this study demonstrated a specific role for accounting smart 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 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 express their gratitude to Dr. Lisa L. Gough for critical reading of the manuscript. This work was supported by pollution, 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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