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William Blythe Essay Competition — Howard Holderness

NAD + -aminoaldehyde dehydrogenase candidates for 4-aminobutyrate (GABA) and β-alanine production during terminal oxidation of polyamines in write School apple fruit. E-mail address: christophert@nutriag.com Department of Plant Agriculture, University of Guelph, Guelph, Ontario N1G 2W1, Canada Search for more papers by this author 1 Present address: NutriAg Ltd., 62 Arrow Rd, North York, Ontario M9M 2L8, Canada. Corresponding author. Fax: +1 519 767 0755. First published: 18 August 2015 Full publication history DOI: 10.1016/j.febslet.2015.08.005 View/save citation Cited by how to good academic essays Domus Academy (CrossRef): 0 articles Check for essay School updates. The last step of polyamine catabolism involves the essay Warminster School oxidation of 3-aminopropanal or 4-aminobutanal via aminoaldehyde dehydrogenase. In this study, two apple ( Malus x domestica ) AMADH genes were selected ( MdAMADH1 and MdAMADH2 ) as candidates for encoding 4-aminobutanal dehydrogenase activity.

Maximal activity and write Holderness catalytic efficiency were obtained with NAD + and 3-aminopropanal, followed by 4-aminobutanal, at pH 9.8. NAD + reduction was accompanied by the production of GABA and β-alanine, respectively, when 4-aminobutanal and 3-aminopropanal were utilized as substrates. Md AMADH2 was peroxisomal and writing a good article International Md AMADH1 cytosolic. These findings shed light on the potential role of apple AMADHs in how to write reflective 4-aminobutyrate and β-alanine production. 4-Aminobutyrate (GABA) is argumentive King’s, a ubiquitous, non-proteinaceous four-carbon amino acid that functions in metabolism and cell signaling. How To Holderness! GABA markedly accumulates in plants subjected to various biotic and abiotic stresses including anoxia, chilling, drought, salinity and mechanical damage[13] and is biosynthesized via two distinct pathways. GABA can be derived from glutamate via pH- and calmodulin-dependent glutamate decarboxylase activity in the first step of the GABA-shunt pathway[4,5]. Alternatively, oxidation of putrescine and spermidine can contribute to Warminster School GABA production[6]. In dicotyledonous plants, the copper-containing amine oxidases (CuAO; EC 1.4.3.22) are responsible for the conversion of putrescine to 4-aminobutanal (ABAL), spermidine to 1,3-diaminopropane and reflective Holderness School ABAL, and 1,3-diaminopropane to 3-aminopropanal (APAL)[7,8]. ABAL is in argumentative essay University rapid non-enzymatic equilibrium with Δ −1 -pyrroline, and the oxidation of these two metabolites is often considered to be catalyzed by the same enzyme, generally referred to as NAD + -dependent aminoaldehyde dehydrogenase (AMADH, EC 1.2.1.19)[6], leading to the biosynthesis of how to reflective Holderness, GABA.

The activity of plant AMADHs has generally been determined as substrate-dependent NADH production and there is only preliminary evidence, using excised tissue or crude protein extract, for the production of GABA from ABAL[9,10]. AMADHs are classified into purchase Warminster several different aldehyde dehydrogenase families (ALDH9, ALDH10, ALDH25, ALDH26 and ALDH27)[11,12]. ALDH9 enzymes are found in animals, fungi and bacteria, whereas ALDH10 enzymes are found in plants and bacteria, and ALDH25, ALDH26 and ALDH27 enzymes are found in bacteria. All plant AMADHs classified as ALDH10 display 7085% sequence identity to each other and 3539% sequence identity with bacterial, fungal or mammal orthologs from the other families[1315]. Genome analysis of several higher plant species has revealed two homologous AMADH genes per species, with the how to write reflective exception of cucumber, grape and maize[1416]. Historically, the AMADHs have been frequently designated as 4-aminoaldehyde dehydrogenase, 4-guanidinobutyraldehyde dehydrogenase, betaine aldehyde (BAL) dehydrogenase or trimethylaminobutyraldehyde dehydrogenase, depending on substrate preference. Missihoun et al.[16] reported the existence of two putative AMADH genes in Arabidopsis , AtALDH10A8 and AtALDH10A9 , although it was possible to generate and purify recombinant At ALDH10A9 only in argumentive King’s Ely sufficient quantities for demonstration of NAD + reduction in the presence of BAL, ABAL or APAL. Furthermore, these authors suggested that At ALDH10A8 and At ALDH10A9 are targeted to how to write reflective School leucoplast and peroxisome, respectively. Purchase School! Study of the how to reflective essay Holderness biochemical properties of plant AMADHs has become attractive because of the Moyles potential involvement of these enzymes in important physiological processes.

For example, Basmati or Jasmine rice, which lacks a functional ABAL dehydrogenase protein, acetylate ABAL (or its cyclic form Δ 1 -pyrroline), resulting in the accumulation of 2-acetyl-Δ 1 -pyrroline, a potent component of rice fragrance[17]. Plant AMADHs also catalyze the biosynthesis of osmoprotectants such as glycine betaine via the oxidation of BAL in reflective the chloroplast stroma[14,18]. Other research has described the accumulation of GABA in fruits, particularly apple, stored under controlled atmosphere conditions, which include low temperature, low O 2 and elevated CO 2 (03 C, 12.5 kPa O 2 , 220 kPa CO 2 )[6,19,20]. Previous research has demonstrated that the peroxisomal MdAO1 gene is highly expressed in King’s Ely apple fruit and write reflective the encoded protein prefers diamines such 1,3-diaminopropane, putrescine and gpa calculator cadaverine as substrates[7]. In the present study, two AMADH genes were identified and cloned from the ALDH10 gene family (designated as MdAMADH1 and MdAMADH2 ). These two genes were heterologously expressed and the resultant recombinant proteins were purified and biochemically characterized. Both proteins catalyzed the NAD + -dependent oxidation of APAL and ABAL to β-alanine and GABA, respectively, but BAL was a very poor substrate.

Transient expression of green fluorescent protein (GFP) fusions in Arabidopsis protoplasts revealed that Md AMADH2 is peroxisomal, whereas Md AMADH1 is cytosolic. Details of the cloning strategy and primer list are provided in theSupplementary Materials and Methods. 2.2 MdAMADH expression in various plant organs. Total RNA was extracted from the frozen powder of how to write reflective essay Holderness School, mature fruit, fully expanded leaves, and opened flowers as described previously[7]. One microgram of Dnase I (Ambion) treated total RNA was used for first-strand cDNA synthesis with Oligo( dT ) 20 and Superscript III (Invitrogen, Carlsbad, CA) at 50 C for 30 min followed by 55 C, according to the manufacturer's protocol. Quantitative real time PCR was conducted as described previously[7]. Bcpm London! Relative expression and data analysis were determined using the 2 −ΔCt method[21] and how to reflective essay Holderness the apple Elongation factor 1-α ( MdEF-1α , Apple Genome Database MDP0000294265), as a superior housekeeping gene that is constitutively expressed[22,23]. The efficiency of the how to academic primer pairs was determined and only pairs with 90105% efficiency were selected for further experiments. Two technical replicates were averaged for each biological replicate, and the mean of three biological replicates was determined for each organ. Primer sequences utilized in gene expression of MdAMADH1 , MdAMADH2 and MdEF-1α are listed inSupplementary Table S1.

2.3 Expression and purification of recombinant proteins. Escherichia coli strain BL21 cells (EMD Millipore) were transformed with standard transformation methods[24]. Recombinant protein expression was induced by adding 0.4 mM isopropyl β- d -1-thiogalactopyranoside to the Lysogeny broth medium supplemented with 50 μg mL −1 ampicillin after attaining an OD 600 of 0.5. The cells were harvested 4 h after induction and essay then stored at −80 C. Bacterial lysis and how to Domus purification of recombinant protein were performed as described elsewhere[7]. Fractions containing high protein levels were pooled and precipitated on ice by how to essay slowly adding ammonium sulfate, with gentle stirring, to 80% saturation.

Total proteins were visualized by SDSPAGE gel electrophoresis, followed by staining with Coomassie Blue R-250 using standard protocols. Immunoblot analysis was based on a semi-dry method using a mouse monoclonal IgG against the His probe (Santa Cruz Biotechnology, 1:1000) and an anti-mouse IgGAlkaline Phosphatase (Sigma, 1:10,000) as primary and secondary antibodies, respectively[7]. Bcpm Gpa Calculator London! The Alkaline Phosphatase Conjugate Substrate Kit (Bio-Rad) was used to visualize fusion proteins. Enzymatic activity was assayed as the substrate-dependent production of NADH[13] or GABA and β-alanine[25]. Details are provided in how to essay Holderness theSupplemental Materials and Methods. 2.5 Transient transformation of good essays Academy, Arabidopsis cell suspension protoplasts.

Protoplasts were prepared from how to write essay Holderness School, Arabidopsis thaliana cell suspension culture ecotype Col-0 as described previously[26]. N- and C-terminal GFP-fusion plasmids containing apple AMADHs were co-transformed with either mCherry peroxisome (pRTL2/Cherry-PTS1)[27], or cytosolic red fluorescent protein (RFP) (pRTL2-MCS-RFP-stop)[28] marker. Protoplasts were transformed using polyethylene glycol 4000 (Sigma) as described previously[29,30]. Confocal laser scanning microscopy was performed with an upright Leica DM 6000B microscope connected to a Leica TCS SP5 system. The excitation/emission scan settings were 488 nm/505 nm for the mGFP channel and 543 nm/610 nm for the RFP/mcherry channel. Modulation of laser light intensity and time-lapse scanning was performed using the Leica software (LAS AF). 3.1 Sequence comparison, conserved motifs and phylogenetic relationship of writing International School Eerde, MdAMADHs and essay Holderness School some characterized AMADHs.

In silico analysis of the argumentive King’s apple genome database revealed two putative members of the how to write essay School ALDH10 gene family. Full length genes were cloned using apple fruit cDNA, followed by rapid amplification of essay Warminster School, cDNA ends. These genes were designated by the common names MdAMADH1 (GenBank Acc. Reflective Holderness! No. KP218040) and MdAMADH2 (GenBank Acc. No. KP218041) for our purposes, but were recently classified by the ALDH nomenclature committee[11] as ALDH10A9 and ALDH10A8, respectively. MdAMADH1 and purchase Warminster School MdAMADH2 encode proteins of reflective essay, 55.0 and 54.6 kDa, respectively, and are 94.8% identical to each other. Md AMADH1 shares 82.6% identity with A. thaliana At ALDH10A8 and 79.1% identity with At ALDH10A9, whereas Md AMADH2 shares 82.8% identity with At ALDH10A8 and 80.5% identity with At ALDH10A9.

Comparison of the putative AMADHs from apple with characterized AMADHs from other plant species reveals that apple and Arabidopsis AMADH isoforms are very similar to each other, suggesting that the two members of the ALDH10 family in writing essay School various species are the result of independent gene duplication[15]. Sequence comparisons indicate that apple AMADHs possess three highly conserved catalytic residues (N162, E260 and C294, Md AMADH numbering), which form the active site in Ps AMADHs[31], and two conserved aspartate residues located at the entrance of the substrate channel (D110, D113), as well as Y163 and W288, which are considered essential for high-affinity binding of how to reflective essay Holderness, ω-aminoaldehydes such as APAL[32] (Supplementary Fig. S1). Together, these data suggest that Md AMADHs are typical AMADHs in persuasive argumentative essay Auburn University the ALDH10 family. 3.2 Expression analysis of how to write reflective essay, MdAMADHs.

The levels of MdAMADH1 or MdAMADH2 expression were similar across fruit, leaves and flowers (Fig. 1 ). However, the expression of MdAMADH1 was two to writing essay Court three times that for MdAMADH2 in all organs studied. Organ-specific expression of two MdAMADH genes. Transcript abundance in corresponding organs was quantified by real-time RT-PCR using gene-specific primers and normalized to a housekeeping gene, EF-1α . Write Reflective School! Data represent the mean ± S.E. of three biological replicates. 3.3 Biochemical properties and substrate specificity of writing International School, apple AMADHs. The cDNAs for MdAMADH1 , MdAMADH2 were individually expressed in E. coli and the recombinant His-tagged proteins purified to how to write essay School homogeneity, as indicated by Coomassie blue staining and immunoblot analysis of the soluble and affinity-purified fractions (Supplementary Fig. S2).

Enzyme activity was determined as the production of thesis Ely, NADH during the how to write reflective Holderness oxidation of APAL, ABAL or BAL; activity with BAL was detected only after increasing the protein concentration in the enzymatic reaction to ten times the level used for the APAL-dependent reaction (i.e., 100 nM). Initial measurements of maximal BAL-dependent activity were two to three orders of gpa calculator DLD College London, magnitude less than those for APAL and therefore BAL was omitted from the kinetic studies. Stability tests revealed that recombinant apple AMADH proteins retained their enzymatic activity for several months after ammonium sulfate precipitation and storage at how to essay Holderness School, −80 C. DLD College London! The activities of Md AMADH1and Md AMADH2 displayed a sharp pH optimum at pH 9.8, a similar pattern over write Holderness, the pH range 711, and 20% of their maximal activity at pH 8 (Fig. 2 ). School! The maximal APAL-dependent activities of Md AMADH1 and Md MADH2 with saturating NADP + were only 1020% of those with saturating NAD + (data not shown). Activity of Md AMADHs as a function of pH. Assays were run at essay Holderness, room temperature with HEPESGlycine buffer, 1 mM ABAL and School 1 mM NAD + . Activity is expressed as a percent of the activity at the pH optimum (100%: 1.2 and how to write essay 1.5 μmol −1 min −1 mg −1 for writing essay Moyles Court Md AMADH1 and Md AMADH2, respectively).

Data represent the mean of four technical replicates from a typical enzyme preparation. NAD + -dependent activities at pH 9.8 demonstrated that Md AMADH1 and Md AMADH2 display partial substrate inhibition for both APAL and ABAL (Table 1 ;Supplementary Fig. S3). The enzymes have approximately four to 18 times higher affinity and 65120 times higher catalytic efficiency for how to essay Holderness APAL than ABAL. APAL displays ten to 50 times stronger inhibition than ABAL across the two enzymes.

These findings are in general agreement with previous reports for Zm AMADH1a, Zm AMADH1b, Zm AMADH2, Ps AMADH1 and Ps AMADH2[13,31]. Persuasive Essay Auburn University! When APAL was provided at a subsaturating level to limit substrate inhibition and allow for sufficient catalytic rates, there was a three-fold range in affinity for NAD + across the enzymes (Table 1). The kinetic parameters for each Md AMADH were determined using a typical enzyme preparation and a reaction solution containing 50 mM glycineNaOH buffer at pH 9.8, 1 mM NAD + , 10% glycerol, 10 mM 2-mercaptoethanol and how to write reflective essay School various concentrations of ABAL or APAL. The data were fit to Moyles the appropriate MichaelisMenten equation using non-linear regression. How To Write Holderness! The kinetic curves are shown inSupplementary Fig. How To Write Academic Essays! S3.

V max (μmol min −1 mg −1 protein), K m (μM), catalytic efficiency ( k cat / K m ) and substrate inhibition constant ( K is , μM) are shown for each substrate or coenzyme, as appropriate. NAD + dependence was estimated in the presence of 40 μM APAL to minimize substrate inhibition; therefore, these parameters are apparent values. How To Write Holderness! To confirm the bcpm DLD College biochemical function of the Md AMADHs, ABAL or APAL was supplied to essay Holderness the recombinant proteins in vitro at concentrations approximately two times their corresponding K m values in argumentative University order to minimize substrate inhibition. The production of GABA or β-alanine was readily detectable within 30 s after initiating the reactions, and the accumulation of these products over time depended on enzyme concentration and write reflective School substrate preference (Fig. 3 ). Notably, the how to write good academic essays Domus estimated initial reaction rates were similar to those obtained by monitoring NADH production (Supplementary Fig. S3). Furthermore, the accumulation of Holderness, β-alanine rapidly reached a plateau, even though 5669% of the substrate remained, suggesting that enzymatic activity was subject to product inhibition (Fig. 3).

Production of GABA and writing a reflection essay School β-alanine by recombinant Md AMADH1 (A) and Md AMADH2 (B) incubated with ABAL or APAL. For assay of ABAL-dependent activities, the 5 mL reactions contained 50 nM protein, 1 mM NAD + and 200 ( Md AMADH1) or 300 ( Md AMADH2) μM ABAL, whereas for assay of APAL-dependent activities, the write essay School reactions contained 10 nM protein, 1 mM NAD + and 30 ( Md AMADH1) or 20 ( Md AMADH2) μM APAL. Data represent the mean of three technical replicates from a typical enzyme preparation. Closed and open circles represent GABA and β-alanine, respectively. The estimated rates of GABA production at 30 s were 0.96 and 2.16 μmol min −1 mg −1 protein for Md AMADH1 and Md AMADH2, respectively. King’s Ely! The corresponding rates of β-alanine production were 11.95 and 13.4 μmol min −1 mg −1 protein, respectively.

Substrate conversion after 15 min was 1217% and 3144%, respectively, for the ABAL- and APAL-dependent reactions. How To Holderness School! 3.4 Subcellular localization of apple AMADHs. Amino acid sequence analysis revealed that the Md AMADHs, as well as At ALDH10A9, Os AMADH1, Os AMADH2, Zm AMADH1a, Zm AMADH1b, Zm AMADH2, Hv AMADH1, Ps AMADH1, Ps AMADH2 and Sl AMADH2 proteins possess a C-terminal canonical peroxisomal targeting signal 1 (PTS1, SKL), whereas At ALDH10A8, Hv AMADH2, So BADH and Sl AMADH1 do not (Supplementary Fig. S1). At ALDH10A8 is reported to essay Moyles localize to the leucoplast[16], whereas So BADH and Hv BBD2 are plastidial and cytosolic, respectively[18,33]. The subcellular location of reflective Holderness, Sl AMADH1 has not been studied. To examine the subcellular localization of gpa calculator, apple AMADHs, Arabidopsis protoplasts were transformed with GFP-fusion constructs and organellar markers such as mCherry peroxisomal and RFP cytosolic were employed. The subcellular locations of the GFP-tagged AMADHs were determined by confocal laser-scanning microscopy. In all cases, both C- and N-terminal GFP fusion constructs were made to exclude the possibility that localization was dependent on the position of the GFP tag.

GFP- Md AMADH1 did not co-localize with the mCherry peroxisomal marker (Fig. 4ad), whereas GFP- Md AMADH2 displayed punctate structures and partially co-localized with the mCherry peroxisomal marker (il). This finding demonstrated that members of the apple AMADHs are present in two different subcellular compartments, the cytosol and the peroxisome, as described above for several plant species. This might indicate non-redundant functions for Holderness School the Md AMADHs at argumentative Auburn University, the protein level. Md AMADH1-GFP and how to reflective Holderness School Md AMADH2-GFP co-expressed with RFP cytosolic were exclusively localized in cytosol (Fig. 4eh and mp, respectively). Similar behavior has been observed with C-terminal GFP fusion constructs of DLD College London, Md AO1[7]. This suggests that the how to reflective Holderness targeting role of the C-terminus PTS1 is masked in the Md AMADH1-GFP and Md AMADH2-GFP fusion proteins by the GFP tag. Subcellular localization of Md AMADH1 and Md AMADH2 fusion proteins in Arabidopsis cell suspension protoplasts.

Confocal laser scanning microscopic images of a protoplast transformed simultaneously with plasmid expressing either GFP- Md AMADH1 (a), Md AMADH1-GFP (e), GFP- Md AMADH2 (i) or Md AMADH2-GFP (m) (i.e., the first column) and mCherry peroxisomal marker (b and j) or RFP cytosolic marker (f and n) (i.e., the second column). A merged image of the GFP and Moyles Court mCherry or RFP fluorescence images is shown in the third column (c, g, k and o) and co-localization of green and red fluorescence signals appears yellow. Write Reflective Essay School! Cell morphology was observed with transmitted light microscopy and is shown in fourth column (d, h, i and p). Images were taken 16 h after transformation. The experiment was repeated three times with similar results. Bars represent 10 μm. GABA can be derived via the oxidative deamination of King’s, polyamines in addition to the GABA-shunt pathway[6]. A pharmacological approach estimated that the terminal oxidation of putrescine accounts for how to write reflective essay Holderness School 39% of the GABA concentration in roots of soybean seedlings exposed to salt stress[34]. A Good Eerde! There is reflective essay, a paucity of information about the role of polyamine oxidation in essay Court School GABA biosynthesis in fruit[35]. Recently, Zarei et al.[7] described an reflective Holderness School apple fruit Md AO1 that is peroxisomal and persuasive argumentative has high catalytic efficiency for write School 1,3-diaminopropane and putrescine to produce APAL and ABAL, respectively. In the present study, we identified two members of the ALDH10 protein family which encode apple AMADHs.

The pH optimum for argumentive thesis King’s Md AMADH1 and Md AMADH2 was 9.8. These findings are consistent with the write reflective Holderness pH optima (9.410.2) of essay Moyles Court School, plant AMADHs that have been previously characterized (i.e., Zm AMADH1a, Zm AMADH2, Sl AMADH2, Ps AMADH1, Ps AMADH2, Sl AMADH1, Zm AMADH1b and Hv BBD1)[13,33]. It has been proposed that the catalytic C294 ( Md AMADH numbering) preserves its thiolate form when pH is significantly above 8 to accelerate nucleophilic attack at write reflective Holderness, the substrate[13]. The apple AMADHs studied here favored NAD + as the cofactor over NADP + with saturating APAL. Previously, Kope#x10d;ný et al.[13] reported that the E188 residue (pea, apple or Arabidopsis AMADH numbering) is responsible for NAD + preference (Supplementary Fig. S1), indicating that the apple AMADHs are probably NAD + -dependent enzymes. The apple AMADHs had negligible activities with BAL, findings similar to those previously reported for Zm AMADH1b, Zm AMADH2, Sl AMADH2 and Ps AMADHs[13]. Notably, Muñoz-Clares and co-workers[14,15] demonstrated that residue position 441 (SoBADH numbering) must be occupied by purchase Warminster School A or C for efficient BAL oxidation. For example, the BAL-dependent activity of reflective Holderness, a So BADH A441I mutant is only 0.5% of that for purchase School the wild-type protein. This position (residue 444 based on numbering of the apple and Arabidopsis AMADHs) is how to reflective essay School, generally occupied by writing School Eerde an Ile residue in how to reflective essay Holderness School plant AMADHs (Supplementary Fig. S1); exceptions are Zm AMADH1a, Hv AMADH2 and So BADH, which have high affinity for purchase essay Warminster School BAL and where I is replaced by reflective essay Holderness C, C and A, respectively[14,15], suggesting that the apple ALDH10 family proteins studied here should not be very active, if at bcpm London, all, with BAL.

In the how to reflective essay Holderness School present study, we demonstrated that in vitro APAL was preferred over ABAL as a substrate (Table 1 andFig. Warminster! 3). However, the write compartmentation and pH optimum of the enzyme, as well as availability of the alternative substrates, would be important determinants in the functional role of the enzyme in vivo. Notably, the use of multiple substrates (i.e., substrate ambiguity) by an enzyme is becoming a better recognized phenomenon, which is thought to enhance the performance of cellular functions[36]. Subcellular localization studies of apple AMADHs were conducted using Arabidopsis cell suspension protoplasts. International! A majority of how to write reflective essay, plant AMADHs is targeted to the peroxisome, as indicted by the presence of the C-terminal PTS1, but there is a reflection essay Court School, also evidence for plastidial and cytosolic localizations[16,18,33].

The presence of canonical PTS1 (SKL) suggests that Md AMADH1 and Md AMADH2 would localize to the peroxisome. Surprisingly, the expressed Md AMADH1 fusion protein appeared to reside in the cytosol, whereas Md AMADH2 was partially targeted to the peroxisome (Fig. 4 ). Notably, both proteins carry identical canonical PTS1 (SKL) and residues in close proximity. Ma and Reumann[37] demonstrated that four and more acidic residues located in close proximity to SKL (10 amino acid upstream of SKL) are incompatible with peroxisomal targeting, altering SKL to a non-functional PTS1. How To Write Reflective Essay Holderness School! The apple AMADHs possess two acidic residues in close proximity. This may explain partial localization of essay Warminster School, Md AMADH2 in how to write reflective School the peroxisome, although the minimum number of persuasive argumentative essay Auburn, acidic residues that are required to hinder peroxisome targeting by SKL has not been determined[37]. Unpredictable masking of PTS1 surface exposure and competition between targeting signals are potential explanations for a non-functional SKL which could prevent Md AMADH1 from reaching the peroxisome.

Recently, it has been reported that peroxisomal proteins often exhibit dual localization and this phenomenon is more widespread than previously thought[38]. Cytosolic AMADH1 could be retained in the cytosol by various mechanisms such protein-protein interaction, protein conformational change in response to pH, or other post-translational modifications, and then released and targeted to the organelle via stress-specific signal perception[39]. This hypothesis would be in agreement with the write reflective Holderness School marked difference between the slightly acidic pH of cytosol and the strongly alkaline pH optima of AMADHs, and with the targeting of the N-terminus and full-length protein of At ALDH10A8 to leucoplasts and cytosol, respectively[16]. This point must be clarified by writing a good Eerde future study. How To Write Reflective Essay School! Supplementary data associated with this article can be found, in the online version, athttp://dx.doi.org/10.1016/j.febslet.2015.08.005. Supplementary data associated with this article can be found, in the online version, at argumentive thesis, http://dx.doi.org/10.1016/j.febslet.2015.08.005. Write Holderness! This research was supported by funds to BJS from the Individual Discovery Grants Program of the Natural Sciences and Engineering Research Council () of Canada.

The authors thank Dr. Argumentive Thesis King’s Ely! Gordon J. Hoover for conducting the amino acid analysis, Dr. Robert T. Mullen for providing the fluorescent protein-containing vectors, the ALDH nomenclature committee for designating systematic names for reflective School our apple genes, and writing Moyles an anonymous reviewer for providing excellent comments on how to, an earlier version of the manuscript. Supplementary data associated with this article can be found, in the online version, athttp://dx.doi.org/10.1016/j.febslet.2015.08.005. FEBS Letters 589 (2015) 1873-3468 2015 Federation of purchase essay Warminster School, European Biochemical Societies. ABAL; 4-aminobutanal; ALDH; aldehyde dehydrogenase; AMADH; aminoaldehyde dehydrogenase; AO; amine oxidase; APAL; 3-aminopropanal; BAL; betaine aldehyde; GABA; 4-aminobutyrate; GFP; green fluorescent protein; NAD(P) + ; nicotinamide adenine dinucleotide (phosphate); PTS1; peroxisomal targeting signal 1; RFP; red fluorescent protein; 4-Aminobutyrate; 4-Aminobutanal; Aminoaldehyde dehydrogenase; Peroxisomal targeting signal; Polyamine; Apple. Issue online: 18 August 2015 Version of how to write reflective, record online: 18 August 2015 Manuscript Accepted: 6 August 2015 Manuscript Revised: 4 August 2015 Manuscript Received: 25 May 2015. Supplementary information. Supplementary Materials and Methods, Figs. S1S3 and Table S1. Please note: Wiley-Blackwell is not responsible for essay Moyles the content or functionality of any supporting information supplied by the authors.

Any queries (other than missing content) should be directed to write School the corresponding author for the article. Ely! 1 B.J. Shelp , A.W. How To Write Reflective Essay Holderness School! Bown , M.D. McLean , Metabolism and functions of gamma-aminobutyric acid . Trends Plant Sci. , 4 , ( 1999 ), 446 452 . CrossRef | PubMed | Web of Science® Times Cited: 366 2 B.J. Argumentive! Shelp , W.L. Allan , D. Faure , Role of how to write reflective Holderness, γ-aminobutyrate and γ-hydroxybutyrate in how to write good Domus plant communication . Write Reflective Holderness School! F. Balu#x161;ka Plant-Environment Interactions ( 2009 ), Springer Heidelberg 73 84 . CrossRef 3 B.J.

Shelp , G.G. Bozzo , A. Zarei , J.P. Simpson , C.P. Trobacher , W.L. Allan , Strategies and tools for studying the metabolism and article International School Eerde function of γ-aminobutyrate in plants. Holderness School! II. International Eerde! Integrated analysis . Botany , 90 , ( 2012 ), 781 793 . How To Write Reflective Essay! CrossRef | CAS | Web of Science® Times Cited: 15 4 B.J. Shelp , G.G. A Good Eerde! Bozzo , C.P. Trobacher , G. Chiu , V.S.

Bajwa , Strategies and tools for studying the metabolism and write reflective Holderness School function of γ-aminobutyrate in plants. I. Pathway structure . Botany , 90 , ( 2012 ), 651 668 . CrossRef | CAS | Web of Science® Times Cited: 17 5 C.P. Academy! Trobacher , A. Zarei , J. Write Reflective Essay Holderness School! Liu , S.M. Clark , G.G. Bozzo , B.J. Shelp , Calmodulin-dependent and calmodulin-independent glutamate decarboxylases in bcpm London apple fruit . BMC Plant Biol. Reflective Essay Holderness! , 13 , ( 2013 ), 144 CrossRef | PubMed | CAS | Web of Science® Times Cited: 9 6 B.J. Shelp , G.G. Bozzo , C.P. Trobacher , A. Zarei , K.L.

Deyman , C.J. Brikis , Hypothesis/review: contribution of putrescine to 4-aminobutyrate (GABA) production in response to abiotic stress . Plant Sci. , 193194 , ( 2012 ), 130 135 . CrossRef | PubMed | CAS | Web of writing a good article School, Science® Times Cited: 41 7 A. Write Essay Holderness! Zarei , C.P. Trobacher , A.R. Cooke , A.J. Meyers , C.J. Hall , B.J.

Shelp , Apple fruit copper amine oxidase isoforms: peroxisomal MdAO1 prefers diamines as substrates, whereas extracellular MdAO2 exclusively utilizes monoamines . Purchase Warminster School! Plant Cell Physiol. , 56 , ( 2015 ), 137 147 . CrossRef | PubMed | Web of Science® Times Cited: 1 8 J. Planas-Portell , M. Write Reflective Essay Holderness School! Gallart , A.F. Tiburcio , T. Altabella , Copper-containing amine oxidases contribute to writing essay Moyles Court terminal polyamine oxidation in peroxisomes and apoplast of Arabidopsis thaliana . Write! BMC Plant Biol. , 13 , ( 2013 ), 109 CrossRef | PubMed | CAS | Web of Science® Times Cited: 22 9 H. Flores , P. Filner , Polyamine catabolism in academic essays Domus Academy higher plants: characterization of pyrroline dehydrogenase . How To Reflective Essay School! Plant Growth Regul. , 3 , ( 1985 ), 277 291 . CrossRef | CAS | Web of Science® Times Cited: 97 10 B. Weilgat , K. Kleczkowski , Putrescine in writing a reflection essay Moyles pea seedlings . Acta Soc. Bot. Pol. , 40 , ( 1971 ), 197 207 . CrossRef 11 V. Vasiliou , D.C. Thompson , C. Smith , M. School! Fujita , Y. Writing A Reflection Court! Chen , Aldehyde dehydrogenases: from write reflective essay Holderness School, eye crystallins to essay School metabolic disease and cancer stem cells . Chem. Biol. Interact. Reflective Holderness! , 202 , ( 2013 ), 2 10 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 19 12 H. Riveros-Rosasa , L. González-Segura , A. Julián-Sánchez , Á.G. Díaz-Sánchez , R.A. Muñoz-Clares , Structural determinants of substrate specificity in a good article School aldehyde dehydrogenases . Chem. Biol. Interact. , 202 , ( 2013 ), 51 61 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 11 13 D. Write Reflective Holderness! Kope#x10d;ný , R. Kon#x10d;itíková , M. Tylichová , A. Vigouroux , H. Moskalíková , M. Soural , M. #x160;ebela , S. Moréra , Plant ALDH10 family: identifying critical residues for substrate specificity and Ely trapping a thiohemiacetal intermediate . How To Essay Holderness! J. Biol.

Chem. , 288 , ( 2013 ), 9491 9507 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 7 14 Á.G. Díaz-Sánchez , L. Article School Eerde! González-Segura , C. Mújica-Jiménez , E. Rudiño-Piñera , C. Montiel , L.P. Write School! Martínez-Castilla , R.A. Essay Warminster School! Muñoz-Clares , Amino acid residues critical for specificity for reflective Holderness betaine aldehyde of the plant ALDH10 isoenzyme involved in the synthesis of gpa calculator DLD College London, glycine betaine . Plant Physiol. , 158 , ( 2012 ), 1570 1582 . Write Holderness! CrossRef | PubMed | CAS | Web of thesis King’s, Science® Times Cited: 19 15 R.A. Muñoz-Clares , H. Riveros-Rosas , G. How To Holderness School! Garza-Ramos , L. Argumentative Essay! González-Segura , C. Mújica-Jiménez , A. Julián-Sánchez , Exploring the how to write reflective essay Holderness evolutionary route of the acquisition of betaine aldehyde dehydrogenase activity by plant ALDH10 enzymes: implications for the synthesis of the osmoprotectant glycine betaine . BMC Plant Biol. Thesis Ely! , 14 , ( 2014 ), 149 CrossRef | PubMed | CAS | Web of write Holderness, Science® Times Cited: 2 16 T.D. Missihoun , J. International Eerde! Schmitz , R. Klug , H.H.

Kirch , D. Bartels , Betaine aldehyde dehydrogenase genes from how to reflective School, Arabidopsis with different sub-cellular localization affect stress responses . Planta , 233 , ( 2011 ), 369 382 . A Reflection! CrossRef | PubMed | CAS | Web of Science® Times Cited: 37 17 L.M. Bradbury , S.A. Gillies , D.J. Brushett , D.L. Waters , R.J. Henry , Inactivation of an aminoaldehyde dehydrogenase is responsible for fragrance in rice . Plant Mol. Biol. , 68 , ( 2008 ), 439 449 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 61 18 P. How To Reflective Essay Holderness! Weigel , E.A. Weretilnyk , A.D.

Hanson , Betaine aldehyde oxidation by spinach chloroplasts . Plant Physiol. , 82 , ( 1986 ), 753 759 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 109 19 J. Lee , D.R. Rudell , P.J. Davies , C.B. Watkins , Metabolic changes in 1-methylcyclopropene (1-MCP)-treated ‘Empire’ apple fruit during storage . Metabolomics , 8 , ( 2012 ), 742 753 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 29 20 C.P. Trobacher , S.M. Clark , G.G. Bozzo , R.T. Mullen , J.R. DeEll , B.J. Shelp , Catabolism of GABA in essay Moyles Court School apple fruit: subcellular localization and biochemical characterization of two γ-aminobutyrate transaminases . Postharvest Biol.

Technol. , 75 , ( 2013 ), 106 113 . CrossRef | CAS | Web of Science® Times Cited: 12 21 K.J. Livak , T.D. Schmittigen , Analysis of relative gene expression data using real-time quantitative PCR and the 2(−delta delta C ( T )) method . Methods , 25 , ( 2001 ), 402 408 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 39019 22 P. Pouparda , L. Parisib , C. Campiona , S. How To Reflective Essay School! Ziadia , P. Writing A Reflection Essay! Simoneaua , A wound- and ethephon-inducible PR-10 gene subclass from apple is differentially expressed during infection with a compatible and an incompatible race of Venturia inaequalis . How To Essay Holderness! Physiol. Essay! Mol. How To Holderness! Plant Pathol. , 62 , ( 2003 ), 3 12 . CrossRef | CAS | Web of argumentive, Science® Times Cited: 22 23 T. Czechowski , M. Stitt , T. Altmann , M.K. Write Reflective School! Udvardi , W.R. Persuasive Essay Auburn University! Scheible , Genome-wide identification and testing of superior reference genes for transcript normalization in Arabidopsis . Plant Physiol. , 139 , ( 2005 ), 5 17 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 1134 24 T. Maniatis , E.F.

Fritsch , J. Sambrook , Molecular Cloning: A Laboratory Manual . Write Essay Holderness School! 2nd ed. Persuasive Essay! ( 1989 ), Cold Spring Harbor Laboratory Cold Spring Harbor, New York 25 W.L. Allan , B.J. Shelp , Fluctuations of γ-aminobutyrate, γ-hydroxybutyrate, and related amino acids in Arabidopsis leaves as a function of the light-dark cycle . Can. J. Bot. , 84 , ( 2006 ), 1339 1346 . How To Write Reflective School! CrossRef | CAS | Web of Science® Times Cited: 20 26 A. Zarei , A.P. Körbes , P. Younessi , G. Montiel , A. Champion , J. Memelink , Two GCC boxes and AP2/ERF-domain transcription factor ORA59 in jasmonate/ethylene-mediated activation of the argumentive Ely PDF1.2 promoter in Arabidopsis . Plant Mol. Biol. , 75 , ( 2011 ), 321 331 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 58 27 S.L.

Ching , S.K. Gidda , A. Rochon , O.R. van Cauwenberghe , B.J. Shelp , R.T. Mullen , Glyoxylate reductase isoform 1 is localized in the cytosol and not peroxisomes in plant cells . J. Integr. Plant Biol. , 54 , ( 2012 ), 152 168 . Reflective Holderness! Wiley Online Library | PubMed | CAS | Web of Science® Times Cited: 10 28 J.M. Shockey , S.K. Gidda , D.C.

Chapital , J.C. Kuan , P.K. Gpa Calculator London! Dhanoa , J.M. Bland , S.J. Rothstein , R.T. Mullen , J.M. Dyer , Tung tree DGAT1 and how to write Holderness DGAT2 have nonredundant functions in purchase Warminster School triacylglycerol biosynthesis and are localized to different subdomains of the write reflective essay endoplasmic reticulum . Plant Cell , 9 , ( 2006 ), 2294 2313 . CrossRef | CAS | Web of Science® Times Cited: 212 29 M. Axelos , C. Gurie , L. Mazzolini , C. Bardet , B. Lescure , A protocol for bcpm gpa calculator London transient gene expression in write essay Holderness Arabidopsis thaliana protoplasts isolated from cell suspension cultures . Plant Physiol. Biochem. , 30 , ( 1992 ), 123 128 . Web of University, Science® Times Cited: 267 30 J. Schirawski , S. Planchais , A.L. Reflective Essay Holderness School! Haenni , An improved protocol for writing School the preparation of reflective essay School, protoplasts from an established Arabidopsis thaliana cell suspension culture and infection with RNA of turnip yellow mosaic tymovirus: a simple and argumentive King’s Ely reliable method . J. Virol. How To Write Holderness! Methods , 86 , ( 2000 ), 85 94 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 28 31 M. Tylichová , D. Kope#x10d;ný , S. Moréra , P. Briozzo , R. Lenobel , J. Snégaroff , M. #x160;ebela , Structural and functional characterization of plant aminoaldehyde from Pisum sativum with a broad specificity for natural and synthetic aminoaldehydes . King’s Ely! J. Mol. Biol. , 396 , ( 2010 ), 870 882 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 24 32 D. Kope#x10d;ný , M. How To Write Holderness School! Tylichová , J. Snegaroff , R. Kon#x10d;itíková , H. Popelková , M. #x160;ebela , Carboxylate and aromatic active-site residues are determinants of high affinity binding of how to essays Domus, ω-aminoaldehydes to plant aminoaldehyde dehydrogenase . FEBS J. , 278 , ( 2011 ), 3130 3139 . How To Write Reflective School! Wiley Online Library | PubMed | CAS | Web of thesis Ely, Science® Times Cited: 11 33 T. Fujiwara , K. How To Reflective Essay School! Hori , K. Ozaki , Y. Yokota , S. Mitsuya , T. Ichiyanagi , T. Hattori , T. Argumentive Thesis Ely! Takabe , Enzymatic characterization of peroxisomal and how to write Holderness cytosolic betaine aldehyde dehydrogenase in barley . Physiol.

Plant. , 134 , ( 2008 ), 22 30 . Wiley Online Library | PubMed | CAS | Web of Science® Times Cited: 33 34 S.G. Xing , Y.B. Jun , Z.W. Hau , L.Y. Liang , Higher accumulation of #x263;-aminobutyric acid induced by salt stress through stimulating the activity of diamine oxidase in Glycine max (L.) Merr. roots . Plant Physiol. Biochem. Writing A Good School! , 45 , ( 2007 ), 560 566 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 58 35 K.L. Deyman , C.J. Brikis , G.G. How To Write School! Bozzo , B.J. Bcpm Gpa Calculator London! Shelp , Impact of write Holderness, 1-methylcyclopropene and controlled atmosphere storage on polyamine and 4-aminobutyrate levels in ‘Empire’ apple fruit . Front.

Plant Sci. , 5 , ( 2014 ), 144 CrossRef | PubMed | Web of Science® Times Cited: 4 36 C. Pandya , J. Farelli , D. Dunaway-Mariano , K.N. Allen , Enzyme promiscuity: engine of evolutionary innovation . J. Biol. Chem. , 289 , ( 2014 ), 30229 30236 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 15 37 C. Argumentive Thesis King’s Ely! Ma , S. How To Write Essay School! Reumann , Improved prediction of peroxisomal PTS1 proteins from genome sequences based on experimental subcellular targeting analyses as exemplified for protein kinases . J. How To Good Academic Essays Domus Academy! Exp. Bot. , 59 , ( 2008 ), 3767 3779 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 24 38 J. Ast , A.C. Write! Stiebler , J. Freitag , M. Bölker , Dual targeting of how to good academic essays, peroxisomal proteins . How To Write Reflective! Front. Physiol. Bcpm Gpa Calculator London! , 4 , ( 2013 ), 297 CrossRef | PubMed | Web of Science® Times Cited: 17 39 M.C.

Cheng , E.J. How To Reflective Essay Holderness! Hsieh , J.H. Chen , H.Y. Chen , T.P. Lin , Arabidopsis RGLG2, functioning as a RING E3 ligase, interacts with AtERF53 and negatively regulates the plant drought stress response . Plant Physiol. , 158 , ( 2012 ), 363 375 . CrossRef | PubMed | CAS | Web of Science® Times Cited: 41. Articles related to the one you are viewing. 2017 Federation of European Biochemical Societies.

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SEO Case Study: 11,065% More Organic Traffic in how to School, 6 Months. Do you want to learn to bcpm gpa calculator rank #1 for high-traffic keywords in how to write essay, ANY niche? Pretty cool, right? This a monster 7,000 word SEO case study, so I've including a table of contents below so you can easily navigate between sections. SEO Case Study: The Step-By-Step Process Michael Karp Used to rank #1 and Generate 20,314 Organic Pageviews (With a Single Post) A little background. The first post I created (and the thesis focus of this SEO case study) for UAV Coach was called How to Fly a Quadcopter — The Ultimate Guide: It's an in-depth 4,400-word guide to essay Holderness safely piloting a remote control quadcopter.

Keyword Rankings and Search Volume. Keyword: how to writing a reflection essay Court fly a quadcopter. Keyword: how to fly a drone. Keyword: how to fly a quadcopter in how to reflective Holderness School, a circle. These rankings bring hundreds of visitors to the site every day:

In the first 6 months, the argumentive Ely post brought in how to reflective, 21.4% of UAV Coach's total traffic . Not only is the post bringing in how to write essays Domus Academy, loads of new traffic. Needless to say, Alan was pretty happy with the results: While I’ve worked with hundreds of companies on their marketing and sales strategies, it wasn’t until working with Michael that I saw such rapid growth in organic, search engine traffic. How To Write Essay School. Doubling our results month-over-month has not only helped us to essay grow our email list and write reflective essay School, generate revenue through our affiliate partners. Argumentive Thesis Ely. It’s helped us to transform our mere industry blog into a real, thriving business. Founder, UAV Coach. The Repeatable 6-Step Formula Used to Rank Content in ANY Niche. You're about to learn the how to reflective Holderness EXACT process I use to create killer content, outrank global brands like Mashable and essay School, deliver thousands of targeted visitors to my client's websites. Here's the short and sweet version: 1. Keyword research and how to essay Holderness, competitive analysis.

2. Content creation. 5. Content promotion. 6. Link building. You’re probably thinking: But Michael, that’s so simple.” Step #1: Keyword Research and Competitive Analysis.

Since the drone space is a relatively new niche, news articles are getting the most links and shares. We wanted to create evergreen content targeted to beginner pilots, and the best way to find these topics was through keyword research. Assess traffic potential (based on monthly search volume) Rank faster by targeting less competitive keywords Maximize organic traffic by targeting keyword variations your competitors are missing Align keywords with search intent at different stages of the buyer journey. Head over to the Google Keyword Planner and a reflection essay Moyles Court School, type in some head keywords: Scroll to the related keywords section and order the write reflective essay Holderness data from highest to lowest search volume:

Next, download the argumentative University data: And review each keyword to find a relevant, evergreen search term to target: This is where I decided to focus on the term “how to fly a quadcopter.” It had decent monthly search volume (720) It's what beginning pilots would be searching for It was a topic I could research easily. BUT the main reason . It had a TON of related long tail keyword variations. Plug your target keyword back into the Keyword Planner and scroll down to the other keywords it generates. You can also use the write reflective essay Holderness School Keyword Options filter to only show results closely related to your search term:

These are great terms to sprinkle throughout the essay University post and boost the relevancy of the content. Learn to fly a drone (140) Beginner quadcopter (480) How to fly a drone (880) Cool, right? You're just getting started. The Google Keyword Planner is reflective essay School good for finding close variations of your seed keyword.

First, perform a Google search for your target keyword and scroll down to the Searches related to. section. This little area is a goldmine for long tail keywords. Pro Tip: Take one of the argumentive Ely keywords from the “Searches related to…” area, pop that into Google, and check out the “Searches related to…” results for that keyword. Reflective Essay. Rinse and repeat until you have a long list of keywords the Google Keyword Planner doesn’t show you. Enter your keyword into Google and look at the other terms Google suggests: These are the most popular terms people are searching for in Google that are related to your seed keyword.

Manually entering keywords into bcpm London Google can be very tedious and time-consuming. But then when you enter your keyword plus another letter, Google Suggest will spit out a different list of suggestions: Ubersuggest will scrape all this data and do all the heavy lifting so you don't have to how to write reflective Holderness spend hours manually entering keyword combinations like quadcopter a , quadcopter b , etc.. The tool will give your hundreds of suggestions. Most of the suggestions won't be a good fit for your site, but a few will.

BONUS STRATEGY: DIG DEEPER WITH SEMRUSH. Enter your target keyword into bcpm SEMrush: From the Keyword Overview report, click to view the full Related Keywords list: Instantly you get access to 951 related keywords, along with search volume, competition level, search trends and a direct link to view the write reflective essay SERP page for each keyword: Click on how to good academic essays Domus Academy, each term to get an how to write Holderness School, overview report showing you hundreds of related terms for each of those keyword. From the list of related keywords I can see best quadcopter gets 5,400 monthly searches. If want to learn more about SEMrush you can read my monster review here, and get a one month free trial here . Seeing the benefits. Unfortunately, this is where many SEOs take off their hat, grab a mojito and call it a day. You need to pay attention to: Domain authority of sites already ranking for the keyword Type and quality of content ranking in the SERPs. By doing competitive analysis, you can save a lot of persuasive essay Auburn, time and effort that would’ve been wasted trying to rank for impossible keywords.

How to analyze your competition (in 15 minutes or less) First, install the MozBar Chrome extension and reflective, do a search for your target keyword. Go back to writing a reflection Court the search results. You should see some useful data below each search result: The Mozbar extension shows: Page Authority Domain Authority Number of links pointing to the page. Each of these can help you determine how hard or easy it is to rank for that keyword. This looks like a promising keyword to quickly rank for.

The title The URL The heading tags Alt text. This is how to reflective essay School where things could have gone wrong for writing a reflection Moyles School, me in a hurry. If these pages had solid backlink profiles and high page/domain authorities, I would have wasted a lot of time creating, promoting, and building links to this article with very little return. Content length Multi media use (images, video, infographics, gifs) Keyword density Content type (list post, expert roundup, interview, case study, how-to etc..) How up-to-date is the content Social media optimization. The entire article was only 303 words (compared to how to reflective Holderness School the 4,000+ I wrote for purchase School, UAV Coach’s). High traffic potential Relatively weak competition Opportunity to create a dominant piece of evergreen content Perfect keyword for our target audience. I had a great list of solid keywords and how to essay Holderness School, a solid foundation for argumentative Auburn, the rest of the SEO campaign. When creating content, there’s one mentality everyone should adopt: It includes in-text links for easy navigation: Note: This is a smart move for any long-form content, because Google will often take some of these links and write essay, put them in the search result. This can help your post stand out in the SERPS and increase click-through rates (now a ranking signal within Google's algorithm).

More in-depth More up-to-date Provides a better user experience Actionable. Important: Try to beat your competition on every level - length, current information, design. On-page factors are the how to write academic Domus Academy elements of a webpage that influence search engine ranking. 1. What your page is about. 2. How To Write Reflective Holderness. What keywords you want to rank for. 3. How valuable your content is. You want to make sure your content is writing a good well-optimized for essay School, your target and related keywords. The correct on-page SEO doesn't take a long time to set up. Keyword placement (title, headers, body, URL, alt text) Keyword variation (LSI) Multi media Internal links External links Load speed. Here's how to essay School successfully optimize your content: Your target keyword needs to be placed in these areas:

If you’re using WordPress, it’s easy to place your keyword in the SEO title, URL, and meta description. URL: Make sure you have a clean permalink structure. Google has stated the first 3-5 words in the URL is weighted as a search ranking factor. If possible, include exact match keyword at the beginning of the URL. First 100 words: Include your target keyword somewhere in the first paragraph of your body copy: Heading tags: Make sure your blog post title is wrapped in write essay Holderness School, an H1 tag. Google views the H1 tag as your post headline. It helps Google better understand the topic of your post. If you’ve got in-text navigation links, it’s also smart to writing a good School Eerde include it there: Google cannot read images, so make sure to include your target keyword in the image alt text so Google can understand what they are about. And finally, place keyword variations a few times throughout the write reflective School body content.

LSI keywords are synonyms that Google uses to determine a page's relevancy (and sometimes quality). Even if the Google Keyword Planner doesn’t show any search volume for your keyword variations, you should still place these in your content for argumentive, relevancy and to make sure you’re not over-optimizing for write reflective essay, exact match keywords. And in-text navigation links: Text can only do so much. Not linking out to external resources is article International a common SEO mistake. Many people fear that doing so will cause their site to leak authority, when in fact linking out to how to reflective essay related content is how to write good academic essays a relevancy signal Google use's to identify the how to reflective topic on King’s Ely, your site. Go through your content and find a few places to link out to the most relevant articles ranking for how to reflective School, your keyword. Internal links can funnel traffic and page rank from other high-traffic authority posts on your site. Google has gone on gpa calculator London, record saying that page load speed is an on-site ranking factor.

Your on-page SEO is fully optimized and ready to hit the front page of Google. Now, it's time to make sure you're converting all that new traffic into email subscribers. Step #4: Turn traffic into email subscribers. Building an write, email list should be your #1 priority. 2. Set up delivery channel. 3. Argumentive King’s. Set up automatic delivery. A lead magnet is a bribe offering something of how to reflective, value to a reader in exchange for their email address. Free ebooks Free trials Checklists ?Mindmaps Infographics Short videos. In this video Robbie showed you how to turn old content into a high-converting lead magnet in writing a reflection essay, 13 seconds. We decided to take a leaf out of Robbie's book and use this same tactic for the quadcopter guide.

2) Optimize your website to capture leads. Optimizing your website involves strategically placing opt-in forms around your site. Popups Scroll mats In-content Two-step lightboxes Header bars Lead boxes. As you can see, our value proposition is joining the reflective essay community and receiving a free PDF of the thesis guide. And it converts at write essay School, 5.12%: We then used the SumoMe? Leads app to create a leadbox that delivers the PDF inside the content. An opt-in box pops up: It converts 20-50+% of the people clicking on the button: Check out essay Moyles Court School Robbie’s latest list building tutorial and this 12,000-word guide to online lead generation and learn more ways to optimize your website to capture email addresses. 5) Automatically deliver lead magnet to reflective Holderness new subscribers. I've got good news and bad news.

2. Deliver the article School lead magnet. The goal of this page is to how to reflective Holderness make sure people are clicking the a good School link in the confirmation email you send them. This will ensure they are double opted in. Ranking content highly in Google is usually a long tail process. Social media automation Established social media accounts Quora Forum marketing Scoop.it.

Let's drive some traffic. First, I started off with one of the biggest time-to-reward strategies: Then click inside the “What do you want to share? ” box: A window will pop up. Write a tweet (or another type of social post) and include a link to your article at the end: Including a hashtag can help extend your post's reach. Choose a date and time that same week to share the article. Create another post, change up the text, and schedule it again for sometime that week. Pro Tip: Use th?is ?strategy to find the how to essay Holderness School best time to schedule your social media posts and bcpm gpa calculator, get more traffic. 2) Established social media accounts.

If you’ve got social media accounts with established followings, it’s important to share your content there as well. Quora is reflective essay Holderness School a massive QA website. Do a search in the top bar for Moyles, a topic related to your niche: This will bring up a list of threads that contain your keyword: Click through to any threads where your content could help answer a question. In-depth answers are trusted more and typically work better than only posting your link. Just like Quora, forums are packed with people discussing different topics. “Keyword” + “powered by vBulletin” Go to one that seems relevant. Then look at these stats to see if it’s active:

Discussions Messages Views Replies. Higher engagement = great traffic potential. Write a description about your page that compels people to how to write reflective essay Holderness School click through. Persuasive Argumentative. Then link to it. As you become an active member, you will also establish a reputation as an authority. 5) Submit content to Scoop.it pages. This tactic doubles as a content promotion and link building strategy. You can suggest a piece of content to write School the owner, and if they like it they’ll share it on School, their page. These pages also get frequented regularly by industry enthusiasts who want to stay up to how to reflective essay Holderness School date with the best information and share it with the platform's built-in social media integration. The default result is a list of “ Scoops ” or specific posts, but you want “ Topics ”:

This will bring up a list of pages related to your keyword. If it’s not there, the host has decided not to writing a reflection essay School accept suggestions. Move on to another page. That’s it! The page owner will review your post and decide whether or not to feature it. 2. Directory submissions. 3. Fake forum account link building. 4. Weekly roundups.

5. How To Write Reflective Holderness School. Resources/links pages. Yes, lots of people have denounced this strategy, but I still do it. A solid DA 58 link: But here’s what happens with blog comments: 2. Being myself, providing a lot of value in how to write good essays Domus Academy, the comment, and linking to my article. Next, I went to write reflective essay School a bunch of directories around the purchase Warminster School web and submitted a direct link to the article. It will most likely go through a review process before being accepted. 3) Fake Forum Account Link Building. Forums are some of the most authoritative sites on the web, especially if they’ve been around for a long time. Weekly roundups are articles that curate the best content of that week into a single post.

“Keyword” + “link roundup” “Keyword” + “weekly link roundup” “Keyword” + “best articles of the week” Then click “Search Tools” > “Any Time” > “Past Month” to find actively updated roundups: Once you’ve found some roundups, head over to the site and find the write reflective essay School publisher’s contact information. Then send them an write Domus Academy, email like this: If the publisher likes your content, they’ll include it in their next roundup:

This got us some great contextual links on some high authority sites: This is the strategy where we got most of our links. It’s the one that really propelled the how to write essay School guide up to the top of the first page. “Keyword” + “intitle:resources page” “Keyword” + “intitle:links page” Unlike weekly roundups, you don’t need to filter the results based on date. How To Write Domus Academy. Since these pages are evergreen, it doesn’t matter when they were published. This strategy landed us a lot of excellent links on highly relevant sites in our niche. And on write reflective Holderness, some high authority sites: Including, a DA 96 link from MIT! (Couldn’t believe that one.) How UAV Coach Hit the #1 Position in Google.

After that link building campaign, I didn’t build any more links to the quadcopter guide. (But notice that it didn’t happen until we promoted our content to a good article International School Eerde get exposure to Holderness it in good academic Academy, the first place.) This shows just how much Google trusts the page, sees it as a top resource, and actively wants people to read it. A Key Takeaway From This SEO Case Study. This is a scalable SEO strategy you can use to rank in ANY niche. You probably won’t rank #1 for every article you publish. Write Reflective Holderness. That’s the nature of argumentive thesis, SEO. I’ve collected the best resources (articles, guides, tutorials, etc) I’ve found that break down each part of the how to essay Holderness content promotion and ranking process you just read: 2. Content creation. 4. Writing International School. List building. 5. Content promotion.

6. Link building. What do you think of these SEO strategies? Have any of your own to share? *Feature image created by how to write reflective essay Holderness, Dario Moriconi from Noun Project. Robbie, this is the best SEO case study i have ever read. The way you illustrates your time consuming research in a post is a reflection Moyles Court much appreciated. You are an inspiration.

Keep it up man! Thanks Fahad, means a lot! However, in this case most of the how to write reflective credit should go to Michael. Thanks for the kind words Fahad. Glad you enjoyed it. Looking forward for great tutorials/case studies here:) Great work Micheal.

Thanks Fahad! Glad to have you as a reader. I’m working on a new case study as we speak. Love reading case studies like these. Thanks so much Michael and how to good academic, Robbie. I’ve noticed that Quora has really picked up in popularity and is a good place to invest your time in. It’s also a great way to ask questions to get feedback or input to use as a research tool, something that I’m currently doing for how to essay Holderness School, some upcoming posts. The step by step process you used for International School Eerde, keyword research was really insightful too. Write School. Thanks for an epic post. Awesome, glad you enjoyed the article Shae! Quora is argumentative essay Auburn definitely a major untapped resource.

It can be leveraged for traffic in write essay Holderness, pretty much every niche out writing essay Court there. Thanks Shae. Robbie and I love putting them together. Totally agree with you on Quora. Like any traffic generation strategy, the more time you put into it the more you’ll get out of write reflective essay Holderness School, it. Thanks for stopping by!

Awesome article! I used the exact steps in my projects. Gpa Calculator DLD College. This tactic provides an excellent result. Thanks, Michael and Robbie for this blog post. I’d like to add that you can get backlinks by stealing them from how to reflective School, your competitors. Check out who is linking to your competitors and try to place your links there. SEMrush and Ahrefs are the DLD College London best tools.

Cheers Michael! Glad you enjoyed the article. Great point about stealing competitor backlinks. Write Essay. Actually wrote an argumentive thesis, article on that exact topic last year: http://www.robbierichards.com/seo/steal-competitors-backlinks-rankings/. Prob needs some updating. That’s a great strategy Michael. If our rankings start dropping it’s one of the ones I’ll turn to first. a rare thorough content maketing case study.. on the level of backlinko (the standard). I know its beyond the scope of this post but paid traffic (ie: reTargetting pixeling on write essay Holderness School, fckbk, ggle-display) and email-sms-snailmail drip sequences for how to good Domus, reEngagement on both link partners and consumer followup would be a wonderful addition. keep up the steller work.

Hey Amul, always nice to be compared with Brian Dean. I’ve read and how to write reflective essay School, learned a lot from how to write Academy, his content. How To Essay Holderness School. I’m experimenting with some paid ad strategies right now on a good International School, personal and client projects. So, I’ll try and put something together in the near future. Any specific area you’re most interested in – retargeting, facebook ads, google display, email automation? I’m hands on with this stuff every day. Reflective Essay School. But, I’d like to focus on one area at writing article International School, a time Cheers! In Spain we say “plas, plas, plas,” when we applaud a artwork. Congratulations for the excellent article. Haha thanks Laura!

Glad you enjoyed the article. This is how to reflective School phenomenal! Thank you. Writing Article. My goal is to get back to producing content on how to write reflective essay Holderness, my own site and driving traffic there. You did a terrific job of walking us through how to do it.

Thanks Jen! Glad it sparked a little motivation. Writing A Good Eerde. Let us know how you fare with it all. Love to here how it goes. Hey Jen, I’m in reflective Holderness School, the same boat. Michael and Robbie, As a marketer, following along with both of your guy’s blogs I’ve learned a lot from these long posts. It’s always cool to see how comprehensive they are that a beginner could jump in, but they also teach more advanced marketers a thing or 2. It’s one thing to talk about a tactic, but totally different to put your money where your mouth is and show how it worked in a case study. Keep it up guys. Always super helpful.

Cheers, Travis! Means a lot. Totally agree, the real value comes through application and sharing the results and lessons learned. Much more actionable (and valuable) that way. Thanks for the comment Travis. Couldn’t agree more. What I’d like to see are more case studies published in other niche’s, in the form of personal stories, triumphs, etc.

I think they’re really underutilized. Untapped keywords is very, very key. Can’t emphasize enough how much this is such a high ROI activity. Having said that, these are all great ideas, many of which I haven’t seen consistently trumpeted around other parts, quite original. Absolutely, Alex! Untapped keywords are usually the easiest to rank for writing, and generally convert better. How To Write Essay School. Why make this SEO thing harder than it has to be, right?! Exactly.

I’m very big on front-loading the work. Spend time finding untapped keywords, collect backlink opportunities, content promotion opportunities, a list of people to writing article Eerde reach out how to write reflective essay to. If you gather all of this before even publishing your content, you’re almost guaranteed to writing essay Court have a successful post. Hi. Many thanks for how to reflective School, your article, a great read and I’ve taken away a couple of things that I hope to apply. I have a quick question, you mention that within your content that you include external links to persuasive essay University high value related content. I just wondered if you use a nofollow for those links or not? Thanks again.

My external links are followed for sure. Thanks for the kind words Lucilla. And I leave them followed as well. Incredibly inspiring case study Robbie. I think thats probably the most detailed case study I have ever read.

Some really good takeaways. Cheers, Paul! But, in this case, most of the School credit goes to my friend Michael. Thanks Paul. Glad you got a kick out of it. And a BIG thanks to Robbie for persuasive argumentative essay University, letting me showcase it here. Wow what an amazing case study Robbie, So much take aways from this post. Write Essay Holderness. I can’t thank you enough for this one! Definitely gonna use it for my future posts. Let the argumentative essay University traffic journey begin:)

Hey Yordi, glad you enjoyed the post. Follow along with it and I’m sure you’ll see some great results! Come back soon and let us know how you go. Also, don’t forget to check out Michael’s infographic. Essay Holderness. Great quick checklist to follow.

Amazing case study and you again encourage all marketers. We have launched new website that is write academic essays Domus Academy about “Web hosting” and I will sure implement this strategy into School my task. Thank you so much for the great strategy. Awesome, Sarfraj! Let me know how you go. Awesome case study and a great strategy which can be done by any SEO guy without spending much. Thanks Ruben! That’s the beauty of this case study, Michael really does break everything down in a way that pretty much anyone can take the process and apply it quickly to their own blog. A brilliant article, I can say that it’s a timely content for me. Purchase Essay. I follow the 2nd part of this case study, But now understand I’ve to reflective essay work on the 1st part too.

Grateful to you Robbie. Cheers! Let me know how go. Thanks for writing Court, such a wonderful proven track you have share with us, I really mean it. I appreciate Michael too because of their major role in this. Now I’ve a reason, visible track most important things is result oriented approach, now I’m going to work in a way which mean something I hope I’ll get my desired result soon. Follow the process Aslam and write, I’m sure you’ll see some great results. I love reading long and detailed post. Well, I must say that this one of the Warminster School detailed post about seo case steady. Loved the Holderness School minute details.

This clearly explains the time you took to how to good academic Academy publish this. Many hours went into this post. But, the positive feedback makes it all worth it. Excellent case study. A great reminder of the power of directories, resource and how to Holderness, round up links. Thanks Sam! Yes, those link sources are often overlooked, but provided a lot of powerful backlink opportunities. Resource pages in particular, as evident in Michael’s case study. This case study is the best thing i have read this week. How To Write. The way you are guiding shows your command on Inbound Methodology but also your sincerity to give some thing worthy to write essay School your readers. I am truly impressed by a reflection, your efforts and how to Holderness, i am expecting more informational posts in future.

Plenty more to come Kashif! Credit goes to Michael on this post. We try to academic make everything 100% actionable for our readers. Write Reflective Holderness School. Glad to see that coming through in the comments. Robbie,Just finished an epic guide it’s really great and persuasive argumentative essay, actionable.I feel comfortable while reading the post and also apply for own site as well. Again thanks for how to Holderness, sharing such an amazing piece of content. Thanks Bilal! Glad you found it helpful. Argumentive Thesis King’s Ely. Let me know how the strategies work for you. Hello Robbie, thanks for your step-by-step guide in ranking website, I will use the strategy for my website http://www.pandapaperroll.com right away, hope it can work out.

Cheers. Awesome Frank! Let me know if you have any questions along the way. Cheers! Excellent article. This is something I honestly wish I would have put to market first.

If I may add one aspect of honorable mention, it’s what I call “binding keywords”. How To. We find binding keywords between buyer persona’s or core business pillars that act like a rudder for content. The idea behind the method is using Home Depot as an gpa calculator, example. By applying branch logic, if someone looks for a paint brush, we assume they have a painting project on the horizon. We would offer them content regarding paint and paint rollers, however we would label “paint” as a binding term and how to essay School, delineate content according to purchase Warminster the content that fits into write Holderness School the same bucket. Further, we determine which buyer personas fill the need for the specified bucket of content, then apply cross linking between content offers AND workflows, segmenting to multiple lists. Assuming we view the process like a grid, this allows us to gpa calculator DLD College engage all three axis of content engagement, rather than two. Reflective Essay. This is wonderful content. Writing Article International Eerde. Thanks for sharing. Thanks for the insight, Brad! Much appreciated.

I take a similar approach when planning my content strategy for clients and personal projects. I refer to it as siloed content. Placing content into related buckets and then mapping the intent of those keywords different stages in the buyer journey. I was just running through links from how to, my old company and re-read this article. What is truly amazing, is your forethought in persuasive essay Auburn University, the article, which holds water 1.5 years later.

I’ve shown this article to multiple clients as an example of how to keep their content on track for their designated verticals and write essay Holderness School, content publishing. Thanks again for the great content. Awesome Bradley! It’s an article School Eerde, oldie, but a goodie Glad it’s been helpful. Thx so much for all of the tips. I found this article on how to, VCB and glad I read it through (in doses). I had never throught about essay Warminster directory submissions for my blog posts and just submitted one to give it a try. And, I went back to scoop.it which I hadn’t used for a while.

I love the weekly or roundup idea, but I have not yet been successful in finding that for my area (hardwood flooring). How To Reflective Essay Holderness School. But, I have hooked up w/ a few realtors doing some of these. Awesome, glad you’re finding the case study helpful Debbie! Let us know if you have any questions. Thanks Michael and write good academic Domus Academy, Robbie for this epic post. How To Write Reflective Holderness School. The key take away for argumentive thesis King’s, me is the ease with which you turn what could be called “boring niche blog posts” into lead magnets.

Many niche bloggers rarely collect emails but you have made email marketing seem very simple.Your conversion rates are much higher than IM industry average of write reflective essay School, 3% conversion. That is incredible. Argumentive Thesis. You have lit some light bulbs in me, now over to Scoop it which I have neglected for so long. I got a link from R-TT after reading backlinko (Brian Dean). Will surely check your other directory links. Keep up the write Holderness good work. Awesome, glad you got some motivation/ ideas from the post Tom!

Let me know how it goes. Here is persuasive essay a short tutorial to reflective really ramp up your list building efforts: http://www.robbierichards.com/email/popup-upgrades/ Robbie, great article, and that UAV site is really next level. Impressive work dude, thanks for sharing. This is an epic post with so much to digest. There are so many gems here I feel like a fossicker who has just discovered buried treasure. I’ve downloaded the International School Eerde ebook and will devour every morsel. Thanks Robbie for how to reflective essay, featuring Michael’s richly detailed tutorial. Thanks Kim!

Glad you enjoyed the purchase essay Warminster post. How To Write Holderness. Let us know how you go with it all. I have only One Question if u dont mind … I heard and Learned from many sites around that Link Building Strategies had been changed over writing a reflection Court School a year or two. How To Write Reflective Essay. Its now a days called PBN Era. If some people may not have enough money to buy a PBN or make it then the argumentive King’s Ely Online World is a waste of time for them ? Hey Usman, definitely not a waste of how to write essay, time. PBNs are just one way of building links and authority.

But, be careful, if done incorrectly you can do more harm than good. Purchase Essay School. Check out Matt Woodward’s PBN course if you want to go that route. He has a lot of success with that strategy. Wow, I was searching for a pragmatic guide from keyword research to write reflective Holderness School building links and I stumbled upon this. I believe I can really implement this strategy and rank for a good International, my desired keywords. I just got hit by a Google Manual Spam action and need to get out of it. This guide will surely help me to excel in my niche. Great post, you’ve earned yourself a hardcore subscriber.

Cheers Sohaib! Glad you enjoyed the post. Essay Holderness. Mind if I ask how you got the manual penalty – duplicate content? Links? Thin content. Actually it was an old site with reviews of 500 words each.

I guess those days are over. I am now building sites from scratch with 2000+ reviews. and yeah I am eagerly waiting for your December post :-). Happy New Year Rob. Have a Blast. New case study going live on writing, Thursday, Sohaib. Robbie, This is what actually I’s looking for.

Believe me, You just changed my whole mind #128512; Now, I think, I can pay with the how to write essay School million Dollar niches #128512; And oh yes, I’m going to DLD College take a challenge today. I will update with every month (Hope, you will see me). Write Holderness School. I’m going to take action end of this year. Look forward to thesis Ely hearing about your progress, Tonmoy #128521; Solid post pal, it definitely shows that relevancy is write reflective essay everything these days, whether it’s onpage or offpage. I’m also starting to do more again with blog comments, forum posts, high TF category pages on web directories and resource page links, in addtion to PBN links #128521;

Cheers Dennis! I’ve had a lot of people mention they were focusing more on purchase Warminster School, the PBN route. Very interesting. I’ve spoken to how to Matt Woodward about it a little. He swears by it, and after seeing some of his results, I can understand why.

Very good article. Congratulations! Thanks guys! Glad you enjoyed the article. I hope you will find the time to write more articles per month because they are terrific. Stefano from Italy. Thanks Stefano! I appreciate that.

I have actually set a goal to write at least 300 words a day for writing a good International, the next 12 months straight. How To Essay Holderness School. This should help me double or triple the content production. Good stuff, I read this article a few weeks ago. Purchase Warminster. I have got a question, I see that you are doing article submission and posting comment. My question is write reflective Holderness School are there any SEO value in Court, it or you are just doing for eyeballs?

Comment posting is essay Holderness School definitely more of an “eyeballs” strategy. Most links in the comment section these days are nofollow so they won’t pass much link equity. Persuasive Argumentative. However, commenting properly on large sites can definitely get you some targeted referral traffic. Wow, Great! I’ve really enjoyed your post. How To Write Reflective Essay. I’ve some questions please: • Did you run any social media ads for that post? • Did you build any backlinks from your competitors backlinks? I am eagerly waiting to essay Auburn hear from you. Again, thanks for sharing the great experience. No social media ads. No link building (yet)

Your screenshots and explanation are just awesome. How To Reflective Holderness School. This is the best case study i read in recent days. Great post, but this spammy window! Hey Karol, thanks for stopping by the blog. Writing A Reflection Court. Just to clarify, by “window”, are your referring to the opt-in popup form? Great post/case study Michael! love the in depth look at how to, what you actually did to argumentive Ely achieve results instead of the School vague aerial view other guides give. Gpa Calculator DLD College London. I guess that’s why Robbie shared it here! Personally I like to use Long Tail Pro for my keyword research. How To Reflective Essay Holderness. It makes it a lot easier and University, more efficient to find related keywords that aren’t too competitive.

Cheers Dominique! I’m starting to write reflective Holderness test out Long Tail Pro as well. Action requires knweoldge, and School Eerde, now I can act! In what case do you have to write essay follow Tiered backlinking. Please suggest, we are operate a digital marketing company. Quadrigo, I don’t really worry about tiered link building. I found your case study through some other guide on persuasive Auburn, a blogging website and I can say it’s one of the best case studies I ever read. Giving out the how to write Holderness School URL and the exact strategies is one of things that people don’t do. But you did it. Bcpm Gpa Calculator London. I took few lessons from write essay Holderness, this post. thanks.

i was amazed by persuasive essay Auburn University, the content you created. i’am an experienced marketer from a big marketing Brand in Holland but, this post really gave me some insight. Just wanted to thank you for this. Cheers BJ! Glad you enjoyed the post. Wow, this is a great resource!

I already have the Moz Chrome Extension, but I have never tried Ubersuggest. I’m going to head over there and play around with it. Wow, this is such an write reflective essay Holderness, amazing article, the info and tips are pure gold, a million thanks for sharing this Robbie! Cheers mate! Glad you enjoyed it. wow, i was searching for SEO Case study on Google and landed to this page, this is one of the best Guide, thanks for sharing. Hi Robbie and Micheal, This is exceptional, really useful case study.

I had to bookmark for later reading. Quora is sure an invaluable resource place for uncovering your audience pain; I started a blog recently and so far I have had just five posts on a reflection Moyles Court School, it. All posts are direct answers to users questions asked on Quora. And from School, what I can tell, the result as been fair to say. The post are getting trickle traffic from writing a good International Eerde, search engine and some significant impression. Consider that this is a blog less than 3weeks old with no links pointing to it. Thanks Micheal, you’ve share something really good with us. how much backlinks you made for single post? I’ve built some articles that have over 200 RDs.

Wow! Robbie, Hats off to you and your posting. This is write reflective essay School just amazing. Never seen such detail post ever as to writing a good Organic Link Building. This is how to reflective essay so precise and descriptive that every SEO builder will be benefited. Thank you so much for how to good Domus Academy, this great post and reflective essay Holderness, awesome knowledge sharing. Thanks Harry. Credit to Mike Karp. Brilliant case study Robbie, even though it was published in 2015 which is writing a reflection Moyles a while in write reflective essay Holderness, the world of SEO, there is still loads to learn there. Cheers for sharing the good content! Terrific case study.

It would take several days or perhaps a week for me to create page like this one. Will certainly try to copy few things in my niche. Thank you very much. Robbie, really amazing case study, ever step is actionable, but I have a few questions, how do you write the article? can I outresource the article? how can I do that? This particular article was written by how to write good essays Domus, a friend of write reflective essay Holderness, mine – Michael Karp. He is essay a great writer, and how to write reflective School, available for King’s, hide. You can find Michael on Twitter here @_MichaelKarp.

Dude, great case study. This shows that the steps above work in write reflective essay Holderness, every niche and you only need hard work to make it possible. THis is the longest article that I ever read! :O seriously! anyway what you said about good academic essays Domus Academy kw research and etc. are true, but to make very very long content! i think its not to important, as long the content are unique and giving differentiation and value to our reader is write essay Holderness School more than enough , cause I heard some ppl giving suggestion its better to Auburn make 3-4 pages, rather than dump all article in one page! I think they right, because it’s to long. It can work to create a multi page post like Brian Dean did with his keyword research guide. It also depends on the niche.

But, for the most part you’re not going to reflective essay School get the good academic essays Domus high rankings with anything under 1500-2000 words. Especially if looking at more top-mid funnel keywords. Ohh, I’m new SEOER, I was searching for SEO Case study and landed to this page, this is one of the best Guide, I will try it. thanks for how to write essay Holderness, sharing. Let me know how it goes, Felix! Robbie, this is awesome the SEO case study, the good academic essays Domus best SEO case study I have ever read.

The way you explanations, hat off!! man, your time consuming research in a post is much appreciated. You are an inspiration. Keep it up man! Cheers mate! Appreciate the how to essay feedback.

Makes the late nights worth it. Hi Michael and gpa calculator DLD College London, Robbie. Thanks for sharing such an amazing guide. I was searching for a step by step SEO case study. Write Reflective. I stumbled here to find two amazing blogs. Got to learn about keyword research, on-page SEO etc from this awsome article.

I was researching about creating my own SEO strategy for argumentive, all the blogs I handle. It would be great if any one of how to Holderness School, you can write a full step by step SEO template or system which people can follow as a protocol. Thanks for the kind words. Funny you mention that. I’m actually working on a course concept right that aligns very closely with what you just described. I’ll let you know when I have that finished.

Hey Robbie, really enjoyed with this case study and uncovered couple of tips which I have never tested for content promotion (scoop) and Link building (Submiting On resources; weekly round ups). Purchase Essay Warminster School. WoW great tips, gonna test them out, I hope it will work more or less for me as well. Thanks for how to write Holderness, sharing these Tips. Keep up the gpa calculator Great work if you have any other blog post similar to how to reflective essay School it about the same topic let me know, would love to dive in that. Thanks Konstantine! You might want to check out the latest blog post too. All my life I’ve been creating content but not able to drive quality traffic, thanks to this comprehensive guide of a reflection essay School, yours Robbie. How To Reflective Holderness. I will practice the way how you create content and will definitely replicate this case study in no time. Again hats off to Ely you for sharing this stuff and I’ll starting doing my research now so that I could create a good quality content.

Cheers Vinson! Glad you’ve found it helpful. How To Essay School. Reach out if you have any questions! Glad to help. Hi.

Many thanks for your article, a great read and I’ve taken away a couple of things that I hope to apply. Write Academic Essays Domus. I have a quick question, you mention that within your content that you include external links to high value related content. Yes – I link out to helpful resources where it adds value. What’s your question? Cool case study.

I really like the content creation part of the article. – Your content should be better than all #1 page contents. Haha cheers mate! Appreciate that. Hi, It’s a brilliant article. How To Write Reflective Essay Holderness School. Great work. Thanks a lot. The SEO Case study is really amazing thank you so much for sharing this information it’s useful for me. Impressive stuff Robbie! I think using Buffer for social media posting offers slightly extra seo weight than ordinary posting.

Worst thing about SEO is that it takes a lot of time and your post shows there’s truth in that! Thanks Joe. Curious to hear more about your point of using buffer to get extra SEO weight from social? Hey, Robbie – Thanks for putting together a great article. Bcpm DLD College. Love the write essay Holderness topic of drones as well! I have never seen such an excellent article written on Digital Marketing. Writing International Eerde. As a beginner, reading through this material was a great learning. How To Write Reflective Essay Holderness. It increased depth of my knowledge and really inspired me.

Thanking you for an excellent piece. Glad it was helpful, Arpit! One of the argumentative Auburn University best SEO Case Study I’ve ever read. Thanks for sharing and revealing your site mate. I’ve got some ideas, Gonna Implement them. Awesome – thank you! Let me know how you go #128521; Printed out this bad boy! Thank you for creating such an amazing resource for the community to learn from.

P.S.: I believe this is 11 thousand words long. Thanks Idris! You’re prob right #128539; It’s a monster! Hope you get some value from it.

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The grade your teacher has scrawled at how to write reflective essay Holderness, the end is nowhere near what your essay deserves. In fact, it’s pretty average. And the comment at the bottom reads something like, ‘Some good ideas, but you didn’t answer the question!’. If asked a question about Keats, you should write about Keats. If this has ever happened to you (and it has happened to thesis, me, a lot), you’ll know how deeply frustrating it is – and write reflective, how unfair it can seem. This might just be me, but the exhausting process of researching, having ideas, planning, writing and re-reading makes me steadily more attached to the ideas I have, and the things I’ve managed to put on purchase, the page. Each time I scroll back through what I’ve written, or planned, so far, I become steadily more convinced of its brilliance. What started off as a scribbled note in the margin, something extra to think about or to how to essay, pop in if it could be made to fit the argument, sometimes comes to be backbone of a whole essay – so, when a tutor tells me my inspired paragraph about argumentive Ely Ted Hughes’s interpretation of mythology isn’t relevant to how to reflective essay Holderness School, my essay on Keats, I fail to see why. How To Good Academic Domus Academy! Or even if I can see why, the thought of taking it out is wrenching. Who cares if it’s a bit off-topic?

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The final question asks you to respond to a quotation. Students tend to purchase essay Warminster, find these sorts of questions the reflective Holderness most difficult to answer, but once you’ve got the hang of them I think the title does most of the work for you – often implicitly providing you with a structure for your essay. The first step is breaking down the quotation into a good article its constituent parts- the different things it says. I use brackets: ( Within Macbeth ’s representation of the how to write essay Holderness School witches, ) ( there is profound ambiguity ) about the ( actual significance ) ( and power ) of ( their malevolent intervention ) Examiners have a nasty habit of picking the most bewildering and terrifying-sounding quotations: but once you break them down, they’re often asking for something very simple. This quotation, for persuasive Auburn example, is asking exactly the same thing as the other questions. How To Holderness School! The trick here is making sure you respond to all the different parts. You want to make sure you discuss the following: Do you agree that the status of the good witches’ ‘malevolent intervention’ is ambiguous?

What is its significance? How powerful is it? James I, the King of England and Scotland at the time Macbeth was written, famously wrote ‘Daemonologie’, which encourages the practice of how to reflective essay Holderness School, witch-hunting. Having worked out exactly what the question is argumentive King’s Ely, asking, write out a plan (which should be very detailed in a coursework essay, but doesn’t have to be more than a few lines long in how to write reflective essay, an exam context) of the material you’ll use in each paragraph. Make sure your plan contains a sentence at the end of each point about how that point will answer the question. A point from my plan for argumentative Auburn University one of the topics above might look something like this: To what extent are we supposed to believe in the three witches in Macbeth ? Hypothesis: The witches’ role in Macbeth’s downfall is write School, deliberately unclear. Their claim to reality is uncertain – finally, they’re part of an School Eerde, uncertain tragic universe and write reflective essay School, the great illusion of the theatre. At the time Shakespeare wrote Macbeth , there were many examples of a reflection essay Court School, people being burned or drowned as witches There were also people who claimed to be able to exorcise evil demons from people who were ‘possessed’. Catholic Christianity leaves much room for the supernatural to write, exist This suggests that Shakespeare’s contemporary audience might, more readily than a modern one, have believed that witches were a real phenomenon and did exist. My final sentence (highlighted in red) shows how the material discussed in the paragraph answers the a good article International School Eerde question.

Writing this out at how to write essay School, the planning stage, in argumentative Auburn, addition to clarifying your ideas, is a great test of whether a point is reflective Holderness School, relevant: if you struggle to write the sentence, and make the connection to the question and good essays Domus Academy, larger argument, you might have gone off-topic. Step Three: Paragraph beginnings and endings. This 16th century English illustration shows a witch feeding her familiars. The final step to making sure you pick up all the reflective possible marks for ‘answering the question’ in writing essay Court School, an essay is ensuring that you make it explicit how your material does so. This bit relies upon getting the beginnings and write Holderness School, endings of bcpm DLD College, paragraphs just right. To reiterate what I said above, treat your reader like a child: tell them what you’re going to say; tell them how it answers the question; say it, and then tell them how you’ve answered the reflective essay Holderness School question. This need not feel clumsy, awkward or repetitive. The first sentence of thesis King’s Ely, each new paragraph or point should, without giving too much of how to write essay School, your conclusion away, establish what you’re going to discuss, and purchase essay Warminster, how it answers the question. The opening sentence from the how to write reflective School paragraph I planned above might go something like this:

Early modern political and religious contexts suggest that Shakespeare’s contemporary audience might more readily have believed in witches than his modern readers. The sentence establishes that I’m going to discuss Jacobean religion and witch-burnings, and also what I’m going to a reflection Court School, use those contexts to show. I’d then slot in all my facts and how to write Holderness School, examples in the middle of the paragraph. Bcpm Gpa Calculator DLD College London! The final sentence (or few sentences) should be strong and decisive, making a clear connection to the question you’ve been asked: Contemporary suspicion that witches did exist, testified to how to reflective School, by witch-hunts and exorcisms, is writing a good article, crucial to our understanding of the witches in Macbeth. To the early modern consciousness, witches were a distinctly real and dangerous possibility – and the witches in the play would have seemed all-the-more potent and terrifying as a result. The best way to write reflective School, get really good at making sure you always ‘answer the question’ is to write essay plans rather than whole pieces.

Set aside a few hours, choose a couple of essay questions from past papers, and for each: Write a hypothesis Write a rough plan of what each paragraph will contain Write out the first and last sentence of each paragraph. You can get your teacher, or a friend, to look through your plans and give you feedback. If you follow this advice, fingers crossed, next time you hand in an essay, it’ll be free from red-inked comments about argumentive King’s irrelevance, and instead showered with praise for the precision with which you handled the topic, and how to Holderness, how intently you focused on answering the question. It can seem depressing when your perfect question is just a minor tangent from the article International School Eerde question you were actually asked, but trust me – high praise and good marks are all found in how to write essay Holderness, answering the persuasive argumentative question in front of you, not the one you would have liked to see. Teachers do choose the questions they set you with some care, after all; chances are the question you were set is the more illuminating and rewarding one as well. 40 Responses to “Focus and Precision: How to Write Essays that Answer the essay Holderness Question” August 21, 2014 at 8:22 am, Kristen Webster said: I have been reading your articles on better essay writing and argumentative essay Auburn University, I am wondering whether you can provide an example of a well written essay please? August 21, 2014 at 11:59 am, ORA Admin said: We haven’t produced any sample essays ourselves.

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