N-myristoylation: An Overview

N-myristoylation: An Overview N-myristoylation Protein N-myristoylation is the covalent attachment of myristate, a 14-carbon fatty acid, onto the N-terminal glycine residues of protein substrates. It is transferred co- or post-translationally to a subset of proteins from a thioester form, myristoyl-CoA, catalyzed by N-myristoyl transferases (NMTs). (insert genes expressing nmt1 and nmt2, NMT recognizes a general consensus sequence for myristoylation (Gly-X-X-X-(Ser/Thr/Cys)) containing a N-terminal glycine, 3 other amino acids and a serine, threonine or a cysteine in the fifth position.) While this process is often observed co-translationally on nascent shorter protein substrates, post-translation myristoylation ensues during apoptosis on N-terminal glycine residue exposed after caspase cleavage of protein substrates.1 An increase in proteins hydrophobicity conferred by this modification allows for weak protein-lipid and protein-protein interactions, as well as for membrane targeting and function of proteins involved in signal tr ansduction cascades.2 NMTs served as therapeutic targets owing to their importance for the survival of human pathogens and their association with carcinogenesis.1 To globally profile NMT protein substrates, chemical proteomic approaches have been employed where small tags on fatty acids such as an alkyne (YnMyr) or azide (AzMyr) (Fig. X) were developed to probe myristoylated proteins via metabolic labeling.3,4 This probe was successfully used in a high-confidence profiling of the co-translational myristoylome in human and zebra fish.5 Although YnMyr remains to be the probe of choice owing to its minimal background labeling6, it was demonstrated to label proteins with other known lipid-modifications such as NÃŽÂ µ-myristoyl,7 S-palmitoyl8 and GPI-anchors9-compromising its specificity towards labeling of N-terminal myristoylated proteins. To circumvent the challenge of identifying the true NMT substrates, Tate et al. used an integrated chemical biology approach where selective inhibition of NMT with smal l-molecule inhibitors combined with YnMyr labeling and quantitative proteomics allowed for profiling of more than 30 known and novel protein candidates for N-myristoylation in blood-stage malaria parasite.9 (describe that the presence of inhibitor abolished the labeling of the true substrates, which should not be enriched in control samples) This technology was also applied to globally profile the N-myristoylome of other human pathogens such as in  Leishmania donovani,10 Trypanosoma brucei,11and recently Trypanosoma cruzi.12Theprofiling of a large set of N-myristoylated proteins with diverse cellular functions unravels the significance of this lipid modification in these parasites. Furthermore, this also validates NMT as a viable drug target in attenuating the virulence of these pathogens. Extending the same approach to HeLa cancer cells enabled the identification of more than 100 of both co- and post-translationally modified N-myristoylated proteins, majority of which were identi fied at endogenous levels for the first time.13 Indeed, this robust technique proved to be powerful in discriminating on-target proteins from off-targets in a proteome-wide analysis, resulting in the discovery of novel NMT protein substrates at high confidence. Although promising, the method described where NMT inhibitors were used may not be applicable to more complex systems where cell viability may be compromised, e.g. in the context of viral and bacterial infection. An alternative targeted approach tosimplify data analysis of enriched proteins employs isolating those that bear the N-terminal glycine requirement for N-myristoylation. This enabled the profiling of downregulated host N-myristoylated proteins upon infection with herpes simplex virus (HSV)14, as well as novel fatty-acylated proteins encoded by HSV. This same approach provided a more defined picture of the demyristoylating function of the bacterial effector IpaJ upon host cell invasion of Shigella flexneri, which was determined to contribute to its virulence.15   Palmitoylation Proteins S-palmitoylation is the attachment of a 16-carbon long fatty acid (as palmitate-CoA) to cysteine residues, which was first discovered by radiolabeling of virus-infected cells with [3H]palmitate.16 The formation of the thioester linkage is mediated by a family of protein acyl transferases (PATs) that bear a conserved Asp-His -His-Cys catalytic motif (DHHC-PATs), which can be removed by hydrolysis aided by acyl protein thioesterases (APTs).17 Owing to the reversibility of this modification, S-palmitoylation of proteins was thought to be dynamically regulated ,whereby a subset of proteins are transiently palmitoylated in a certain time point/cellular activity. (insert something) S-palmitoylation has been demonstrated to be an essential mechanism for protein stability, activity, and proper cellular localization.18 Recent advances in identifying palmitoylated proteins revealed not only its key role in regulatory mechanisms but as well as in host invasion and virulence of pathogen s. Large-scale proteomic profiling of S-palmitoylated proteins using metabolic labeling has been heavily dependent on employing the alkyne analogue of palmitic acid, 17-ODYA (Fig. X). This commercially available chemical reporter is suitable for these analyses as it has shown better specificity and has minimal background in labeling proteins that are ought to be acylated by shorter fatty alkyl chains.6 The subsequent click reaction with fluorophore- or biotin-azide then allows for in-gel fluorescence monitoring and biotin-pulldown strategy prior to LC-MS proteomic analysis of labeled proteins, respectively. In these studies, hundreds of palmitoylated proteins were identified with a wide range of functions, highlighting the importance of S-palmitoylation in a plethora of cellular mechanisms and pathways. For instance, the first report on using such strategy applied to mammalian cells identified around 125 candidate S-palmitoylated proteins at high confidence, including G proteins, recept ors and uncharacterized hydrolases.19 Using the same strategy in dendritic immune cells (DC2.4) identified more than 150 predicted S-palmitoylated proteins and revealed that palmitoylation of interferon-induced transmembrane protein 3 (IFITM3)20 and Toll-like receptor 2 (TLR2)21 is essential for their antiviral activity. A more recent study on Cryptococcus neoformans revealed that a single PAT, Pfa4, palmitoylates the fungal proteins required for parasite integrity and virulence-palmitoylating 72 proteins identified in a global-scale approach.22 A more quantitative approach to measure levels of palmitoylated proteins combines metabolic labeling with 17-ODYA and Stable Isotope Labeling with amino acids in Cultured Cells (SILAC). In virus-infected RPE-1 epithelial cells, selective repression was observed for host S-palmitoylated proteins, including interferon signaling regulators and members of the tetraspanin family.14 A novel set of HSV-encoded proteins palmitoylated by the host machinery were selectively and significantly identified, further suggesting that HSV exploits the palmitoylation pathway which contributes to its virulence. As palmitoylation is a reversible process, the dynamic cycling of palmitoylated proteins in mouse T-cell hybridoma cells was investigated using this quantitative approach in combination with a pulse-chase technique.23 Through the use of a serine lipase-inhibitor as the chase, palmitoylated proteins that undergo fast turnovers were distinguished from those that are stably modified. This indicates that a subset of this dynamic palmitoylation event is regulated by serine hydrolases, validating the fundamental regulatory mechanism of depalmitoylation for proteins with rapid turnovers. It is important to note that in this study, only the insoluble protein fractions were analyzed, as the soluble proteins were not amenable to metabolic probe incorporation.19 Given the dynamic nature of palmitoylation, the metabolic labeling strategy would allow labeling of only those that are palmitoylated at the time of probe treatment and were stably modified. An older approach, coined as acyl-biotin exchange (ABE), has the potential to capture the full complement of palmitoylated proteins. In this multistep procedure, the protein lysates are treated with hydroxylamine to selectively cleave the thioester bonds, followed by disulfide capture with thiol-containing biotin analogue, and subsequently enriched through a pulldown technique prior to LC-MS analysis. ABE was first utilized in tandem with semi-quantitative MudPit analysis on profiling the palmitome of Saccharomyces cerivisae.24 The 12 known and 35 new palmitoylated proteins identified presented the first evidence on the diverse specificities of PATs. The ABE method was further employed in profiling the palmitoylome in rat neurons,25 human T cells,26 and recently in poplar tree cells,27 establishi ng its applicability to a wide range of biological systems. Both ABE and metabolic labeling approaches combined with SILAC revealed their large complementarity in profiling S-palmitoylated proteins in Plasmodium falciparum.28 A total of more than 400 palmitoylated proteins were identified where 202 proteins were enriched in both methods. As expected, metabolic labeling identified a lesser number of proteins, reflecting the less complexity in this approach. A pulse-chase labeling using ABE in a quantitative approach with 2-BMP as the parasite PAT inhibitor revealed the identification of a range of stably and dynamically palmitoylated proteins. Indeed, this study demonstrated the importance of palmitoylation in multiple parasite-specific processes, specifically in drug resistance, asexual stage development, host cell invasion, and protein export. Both methods were also employed in investigating the dysregulation of palmitoylation in breast cancer cells by inducing Snail-overexpression- an event correlated with chemoresistance and metastasis.29 Results showed that some proteins were differentially expressed regardless of differential palmitoylation. Thus, Snail-overexpression compromises the dynamic palmitoylation of some proteins that may be involved in pathways that contribute to malignancy. Albeit most proteins are S -palmitoylated in their cysteine residues, others were reported to be O-palmitoylated30 and N-palmitoylated17, which are also labeled by 17-ODYA. To distinguish S-palmitoylated proteins from these other forms in Toxoplasma gondii, a method similar to ABE was employed which also takes advantage of the labilityof thioester bonds to hydrolysis.31 In this approach, the metabolic incorporation of 17-ODYA and enrichment is followed by hydroxylamine cleavage to profile S-palmitoylated proteins. This confirmed 282 hydroxylamine-sensitive proteins from 501 putative palmitoylated proteins enriched from the initial 17-ODYA labeling. This also revealed and validated that palmitoylation of AMA1, a protein essential for host-cell invasion, is not required on invasion but increases microneme secretion. Taken together, these studies presented underscore the utility of large-scale S-palmitome profiling in understanding the biological importance of this lipid modification. Applying these techniques to future palmitome analysis would further discover potentially novel protein functions and cellular mechanisms across different biological systems. Wright, M. H., Heal, W. P., Mann, D. J. Tate, E. W. Protein myristoylation in health and disease. J. Chem. Biol. 3, 19-35 (2010). Farazi, T. A., Waksman, G. Gordon, J. I. The Biology and Enzymology of ProteinN-Myristoylation . J. Biol. Chem. 276 , 39501-39504 (2001). Heal, W. P., Wickramasinghe, S. R., Leatherbarrow, R. J. Tate, E. W. N-Myristoyl transferase-mediated protein labellingin vivo. Org. Biomol. Chem. 6, 2308-2315 (2008). Heal, W. P., Wright, M. H., Thinon, E. Tate, E. W. Multifunctional protein labeling via enzymatic N-terminal tagging and elaboration by click chemistry. Nat. Protoc. 7,105-117 (2012). Broncel, M. et al. Myristoylation profiling in human cells and zebrafish. Data Br. 4, 379-383 (2015). Charron, G. et al. Robust Fluorescent Detection of Protein Fatty-Acylation with Chemical Reporters. J. Am. Chem. Soc. 131, 4967-4975 (2009). Liu, Z. et al. Integrative Chemical Biology Approaches for Identification and Characterization of Erasers for Fatty-Acid-Acylated Lysine Residues within Proteins. Angew. Chemie Int. Ed. 54, 1149-1152 (2015). Wilson, J. P., Raghavan, A. S., Yang, Y.-Y., Charron, G. Hang, H. C. Proteomic Analysis of Fatty-acylated Proteins in Mammalian Cells with Chemical Reporters Reveals S-Acylation of Histone H3 Variants. Mol. Cell. Proteomics 10, M110.001198 (2011). Wright, M. H. et al. Validation of N-myristoyltransferase as an antimalarial drug target using an integrated chemical biology approach. Nat Chem 6, 112-121 (2014). Wright, M. H. et al. Global Analysis of Protein N-Myristoylation and Exploration of N-Myristoyltransferase as a Drug Target in the Neglected Human Pathogen Leishmania donovani. Chem. Biol. 22, 342-354 (2015). Wright, M. H., Paape, D., Price, H. P., Smith, D. F. Tate, E. W. Global Profiling and Inhibition of Protein Lipidation in Vector and Host Stages of the Sleeping Sickness Parasite Trypanosoma brucei. ACS Infect. Dis. 2, 427-441 (2016). Roberts, A. J. Fairlamb, A. H. The N-myristoylome of Trypanosoma cruzi. Sci. Rep. 6,31078 (2016). Thinon, E. et al. Global profiling of co- and post-translationally N-myristoylated proteomes in human cells. Nat Commun 5, (2014). Serwa, R. A., Abaitua, F., Krause, E., Tate, E. W. OHare, P. Systems Analysis of Protein Fatty Acylation in Herpes Simplex Virus-Infected Cells Using Chemical Proteomics. Chem. Biol. 22, 1008-1017 (2015). Burnaevskiy, N., Peng, T., Reddick, L. E., Hang, H. C. Alto, N. M. Myristoylome profiling reveals a concerted mechanism of ARF GTPase deacylation by the bacterial protease IpaJ. Mol. Cell 58, 110-122 (2015). Schmidt, M. F. G. Schlesinger, M. J. Fatty acid binding to vesicular stomatitis virus glycoprotein: a new type of post-translational modification of the viral glycoprotein. Cell 17, 813-819 (1979). Linder, M. E. Deschenes, R. J. Palmitoylation: policing protein stability and traffic. Nat Rev Mol Cell Biol 8, 74-84 (2007). Smotrys, J. E. Linder, M. E. Palmitoylation of Intracellular Signaling Proteins: Regulation and Function. Annu. Rev. Biochem. 73, 559-587 (2004). Martin, B. R. Cravatt, B. F. Large-scale profiling of protein palmitoylation in mammalian cells. Nat Meth 6, 135-138 (2009). Yount, J. S. et al. Palmitoylome profiling reveals S-palmitoylation-dependent antiviral activity of IFITM3. Nat Chem Biol 6, 610-614 (2010). Chesarino, N. M. et al. Chemoproteomics reveals Toll-like receptor fatty acylation. BMC Biol. 12,91 (2014). Santiago-Tirado, F. H., Peng, T., Yang, M., Hang, H. C. Doering, T. L. A Single Protein S-acyl Transferase Acts through Diverse Substrates to Determine Cryptococcal Morphology, Stress Tolerance, and Pathogenic Outcome. PLoS Pathog. 11,e1004908 (2015). Martin, B. R., Wang, C., Adibekian, A., Tully, S. E. Cravatt, B. F. Global profiling of dynamic protein palmitoylation. Nat Meth 9, 84-89 (2012). Roth, A. F. et al. Global Analysis of Protein Palmitoylation in Yeast. Cell 125, 1003- 1013 (2006). Kang, R. et al. Neural palmitoyl-proteomics reveals dynamic synaptic palmitoylation. Nature 456, 904-909 (2008). Morrison, E. et al. Quantitative analysis of the human T cell palmitome. Sci. Rep. 5, 11598 (2015). Srivastava, V., Weber, J. R., Malm, E., Fouke, B. W. Bulone, V. Proteomic Analysis of a Poplar Cell Suspension Culture Suggests a Major Role of Protein S-Acylation in Diverse Cellular Processes. Front. Plant Sci. 7, 477 (2016). Jones, M. L., Collins, M. O., Goulding, D., Choudhary, J. S. Rayner, J. C. Analysis of Protein Palmitoylation Reveals a Pervasive Role in Plasmodium Development and Pathogenesis. Cell Host Microbe 12, 246-258 (2012). Hernandez, J. L. et al. Correlated S-palmitoylation profiling of Snail-induced epithelial to mesenchymal transition. Mol. Biosyst. 12, 1799-1808 (2016). Zou, C. et al. Acyl-CoA:Lysophosphatidylcholine Acyltransferase I (Lpcat1) Catalyzes Histone Protein O-Palmitoylation to Regulate mRNA Synthesis. J. Biol. Chem. 286 ,28019-28025 (2011). Foe, I. T. et al. Global analysis of palmitoylated proteins in Toxoplasma gondii. Cell Host Microbe 18,501-511 (2015).

Natural and Sexual Selection of Vestigial and Wild Type Drosophila Melanogaster

Natural and Sexual Selection of Vestigial and Wild Type Drosophila melanogaster Abstract In this experiment, vestigial flies as well as wild type flies were used to create some diversity as well as test one of our hypotheses. Our hypotheses are as follows; 1) The wild type flies will have a greater relative fitness compared to the vestigial flies based on only sexual selection. The wild type has a relative fitness of 1 since it has fully functional wings. Vestigial Winged flies are not as sexually appealing since their wings are not functional and the wing movement is a vital part of their mating ritual. ) The vestigial fly will have a greater relative fitness when considering both sexual and natural selection. With the presence of natural selection (fly paper hanging from top of cage) posing a larger threat to the wild flies b/c they can fly better, this will allow vestigial flies to prosper better. To set up this experiment, two twenty-five gallon aquariums, 3 petri-dishes, 200 fli es, rotten bananas, and yeast were used. It was decided that there would be 80 vestigial flies and 20 wild type flies to total to an initial population of 100 drosophila.In the sexual selection circumstance, the vestigial drosophila’s carrying capacity is somewhere around 80 individuals, which is consistent with the wild type drosophila’s carrying capacity in this circumstance (Figure 4). In the sexual and natural selection circumstance the wild type drosophila have a carrying capacity at approximately 100 individuals (Figure 5). As both of the p-values from each circumstances in our experiment are greater than 0. 05, both hypotheses must be refuted. IntroductionThis experiment focuses specifically on Drosophila melanogaster, also known as the common fruit fly (Lynch, M. , et al. pp 645-663). For over 100 years scientists have been using them in experiments as they are referred to as a â€Å"model organism. † They were deemed a â€Å"model organism† since they are characterized as having a large reproductive capacity, a medium size, a short generation time, and are inexpensive to keep alive. Fruit flies only consume yeast growing on rotting fruit. Sixty-one percent of human genetic diseases have been found in fruit flies, they also share many similar genes with humans.The male fruit fly spends most of his time chasing after and singing to female flies with hopes of mating with her, they are successful about once a day (Stowe, K. 2008). The male fruit fly frequently sticks out one wing or the other which is accompanied by its vibration, producing a â€Å"love song† that can be recorded with specialized microphones (Hall, Jeffrey C. , pp 1702-1714). To distinguish male drosophila from females, there must be a distinctive black spot on the distal end of their abdomen for it to be a male, the females do not have this black spot.In this experiment, vestigial flies as well as wild type flies were used to create some diversity as we ll as test one of our hypotheses. Vestigial flies do not have well functioning wings, however wild flies do have functioning wings. Within this experiment, degrees of natural selection, sexual selection, and a combination of both were tested on the two types of flies. Natural selection is survival of the fittest. Sexual selection is the ability of organisms to choose their mates depending on their species. Relative fitness is the ability of an organism and species to survive and produce fertile, successful offspring.Purpose of our experiment is to determine the effect of natural selection on two subspecies coexisting under similar circumstances. Our hypotheses are as follows; 1) the wild type flies will have a greater relative fitness compared to the vestigial flies based on only sexual selection. The wild type has a relative fitness of 1 since it has fully functional wings. Vestigial Winged flies are not as sexually appealing since their wings are not functional and the wing moveme nt is a vital part of their mating ritual. 2) The vestigial fly will have a greater relative fitness when considering both sexual and natural selection.With the presence of natural selection (fly paper hanging from top of cage) posing a larger threat to the wild flies b/c they can fly better, this will allow vestigial flies to prosper better. Materials and Methods To set up this experiment, two twenty-five gallon aquariums, 3 petri-dishes, 200 flies, rotten bananas, and yeast were used. The bananas chosen to be an accelerant for the growth of the yeast and  were frozen so they would be easier to cut. The yeast was used because the drosophila melanogaster prefer this as a food source.The vestigial and wild type flies were sexed (to determine their sex), sorted, and counted. An initial population size of 100 total flies was decided so that it would be easier to determine the phenotypic percentage of the total population. Fly paper was placed in one of the sets of cages to impose a m ethod of natural selection as well as the sexual selection which is being solely tested by the other set of cages. It was decided that there would be 80 vestigial flies and 20 wild type flies to total to an initial population of 100 drosophila.Next, the flies were anesthetized flies using Fly Nap. The flies were counted out to reach desired ratio, sexing the flies making sure there are equal amounts of males and females to be sure there is ample individuals to allow successful mating. The fly’s food was prepared by taking a frozen rotten banana, cutting it in half, mashing up the banana meat, and mixing yeast into it. The bananas were then cut into halves horizontally (six halves total in each cage) and split down the middle length wise leaving the peel on the banana, revealing the yeast-banana mush and placed into the petri dishes.The petri dishes were then arranged into a horseshoe shape in the bottom of the aquarium. Next, the allele frequencies were determined by using th e hardy Weinberg equation (p2+2pq+q2). This experiment will prove our hypotheses sufficiently if carried out correctly by changing the total populations of the subspecies of each fly. Since the vestigial flies do not fly they will be affected by sexual selection only. However, the wild type flies will be affected by sexual and natural selection.The outcome that is predicted is that the relative fitness of the vestigial flies will stay low in the sexual selection circumstance and they will prosper in the sexual and natural selection circumstance since they do not fly and are not able to come near the fly paper posing as the natural selection. Also, the wild type flies will remain at a high fitness level in the sexual selection, but a low fitness level in both sexual and natural selection circumstance. Table 1: General Data from Sexual Selection| Generation| Sexual Selection Only| Total Drosophila| Wild Type| Vestigial| | 0| 20| 80| 100| 1|   |   |   | 2| 6| 12| 18| 3| 338| 50| 388| 4| 201| 21| 222| 5| 428| 35| 463| table 1: shown in this table is the initial data of the total count of flies captured for counting in each species each week/generation in the sexual selection group. | Results Table 2: General Data from Natural and Sexual Selection| Generation| Natural and Sexual Selection| Total Drosophila| Trapped Wild Type Flies| | Wild Type| Vestigial| | | 0| 20| 80| 100| 0| 1|   |   |   |   | 2| 16| 59| 75| lots| 3| 23| 10| 33| lots| | 86| 1| 87| lots| 5| 114| 21| 135| Lots| table 2: shown in this table is the initial data of the total count of flies captured for counting in each species each week/generation in the group that was exposed to natural selection as well as sexual selection. | Table 3: Anova: Single Factor, natural selection |   |   | Source of Variation| SS| df| MS| F| P-value| F crit| Between Groups| 63202. 5| 1| 63202. 5| 3. 513094| 0. 09775| 5. 317655| Within Groups| 143924. 4| 8| 17990. 55| | | | | | | | | | | Total| 207126. 9 | 9|   |   |   |   | | | | | | | |Table 3: Shown in this table is the ANOVA calculation which determines the probability that our null hypothesis will be found true. In this case, the proability of that happening is approximately 0. 09 or 9%. This number is low enough to allow us to accept our hypothesis. | Table 4: Anova: Single Factor natural and sexual selection|   | Source of Variation| SS| df| MS| F| P-value| F crit| Between Groups| 774. 4| 1| 774. 4| 0. 486447| 0. 505277| 5. 317655| Within Groups| 12735. 6| 8| 1591. 95| | | | | | | | | | | Total| 13510| 9|   |   |   |   | | | | | | | Table 4: Shown in this table is the ANOVA calculation for the Natural and Sexual selection grouping of drosophila and the possibility of our null hypothesis occurring. In this case, the value is approximately 0. 5, or 50%. This proves that our hypothesis is not completely reliable. | In â€Å"generation 0,† for the sexual and natural selection set of flies, the total sampl e size is 100 flies consisting of 20 wild type (10 male, 10 female), and 80 vestigial (40 male, 40 female), there were no flies stuck on the fly paper yet.The allelic frequency: p=0. 048, p= 0. 961. For the sexual selection set of flies, the total sample size is 100 flies, consisting of 20 wild type (10 male, 10 female), and 80 vestigial (40 male, 40 female), the allelic frequency is p= 0. 78, and q=0. 22. The dominant trait is denoted by the letter â€Å"p† (wild type), and â€Å"q† denotes the recessive (vestigial). After five generations were allowed to pass, the final total of drosophila in the sexual selection group included 428 wild type and 35 vestigial out of 463 total drosophila(Table 1).After five generations were allowed to pass, the final total of drosophila in the sexual and natural selection group included 114 wild type and 21 vestigial out of 135 total drosophila (Table 2). There is an obvious and drastic decrease in the amount of flies after the five ge nerations passed when comparing the two separate groups. Also, it must be pointed out that the set of data does not have values for the first generation because when the flies were counted, the collection of flies to count was unsuccessful.Next, the p-values of each set of data were calculated by using the ‘ANOVA: Single factor’ function in Microsoft Excel. The p-value of the sexual selection only set of data came out to be 0. 097, or 9. 7% that our null hypothesis that the vestigial flies would be more fit than the wild type flies would come true (Table 3). In Table 4, it is shown that the calculated p-value for the sexual and natural selection conditions of the vestigial and wild type flies produced a value of 0. 1, or 51% that our null hypothesis would come true. Figure 2 shows the data collected in the sexual selection circumstance of both the vestigial and wild type drosophila as well as the total population as to compare visually the drastic contrast between the t wo subspecies. The wild type flies remain more successful than the vestigial flies. Figure 1: In this figure, it is shown visually the differences in the numbers of flies counted per each subspecies of drosophila while under only sexual selection conditions.Figure 3 shows the Natural and sexual selection circumstance’s effects on the populations of both vestigial and wild type flies as well as the total number of flies in the enclosure. The Vestigial flies start off more prosperous than the wild type flies, but then the wild type drosophila regain their success and start reproducing at incredible rates. Figure 2: In this figure, it is shown visually the differences in the numbers of flies counted per each subspecies of drosophila while under both natural and sexual selection.Figure 4: In this figure, the relative fitness and carrying capacity of each subspecies is shown. Figures 4 and 5 have been included to compare fitness levels of each of the fly subspecies. The scale on t he y-axis has been multiplied by 10 to better understand the incredible contrast between the species. In both circumstances, the vestigial drosophila starts out with a higher relative fitness then reaches an equilibrium level with the wild type flies and they switch dominance of their relative fitness.Figure 5: In this figure, the relative fitness and carrying capacity of each subspecies is shown. Discussion This curvature of the graphs and intersections happen because when the subspecies reach their carrying capacities, they must make a change so that they can survive as a community. In the sexual selection circumstance, the vestigial drosophila’s carrying capacity is somewhere around 80 individuals, which is consistent with the wild type drosophila’s carrying capacity in this circumstance (Figure 4). In the sexual and natural selection ircumstance the wild type drosophila have a carrying capacity at approximately 100 individuals (Figure 5). As both of the p-values fr om each circumstances in our experiment are greater than 0. 05, both hypotheses must be refuted. It is suggested that this experiment is conducted again using a better enclosure for the flies. During the semester, countless flies were flying around the lab which could account for the numbers of flies being inaccurate. There are possible errors in the sexing as well as the counting of the flies.Only a few flies crawled into the tubes with apple cider vinegar, and didn’t allow for every fly to be accounted for. The temperature of the lab was often quite cold, therefore this could account for some of the flies dying. As temperature was not a factor we were testing in this experiment, the temperature should be kept at a constant level. This experiment is important on a global level because of the closeness in the similarities of the genes between the drosophila and humans (Lynch, M. , et al. pp 645-663).This experiment shows us how, if humans were meant to survive strictly on nat ural and sexual selection, that mutations would play a part in the selection of mates. References Hall, Jeffrey C. Jun 1994. The Mating of a Fly. Science 264 No. 5166: pp 1702-1714. Lynch, M. , J. Blanchard, Houle, D. , T. Kibota, Shultz, S. , L. Vassilieva, Willis, J. Perspective: Spontaneous Deleterious Mutation. Evolution 53 No. 3: pp 645-663. Stowe, K. A. , L. L. Hester, and Vieyra, M. L. 2008. Biology 101 Lab Manual. Hayden- McNeil Publishing Inc. , Plymouth, Michigan, USA.

A Shipwreck of Faith - Free Essay Example

Sample details Pages: 5 Words: 1580 Downloads: 3 Date added: 2019/03/21 Category Religion Essay Level High school Tags: Faith Essay Did you like this example? In Daniel Defoes book Robinson Crusoe, the main character endures many battles along his adventures while entering adulthood. As Robinson Crusoe journeys across the seas, God makes Crusoe enduress many trials. First, starting from when he sets off to sea, disobeying his parents wishes in the process, to encountering many shipwrecks, and lastly the opportunity of sharing his growing faith with others. By Crusoe encountering these many obstacles, it goes just to show how having a weak foundation in faith can reveal many unwanted paths. Christianity plays a major role in this well-known piece, as this is the biggest obstacle Crusoe is struggling with right from the jump. Crusoes father had a life plan set for him ever since he was little. He wanted his son to become a great lawyer, as stated, my fatherdesigned me for the law(Defoe 9). This was not in Crusoes heart obviously, as it was set for the ocean. Crusoe completely ignores what his father was trying to accomplish for his son, and set sail for the great adventure that the seas had to offer, showing us our first Christianity reference. In the Bible, the story of the prodigal son is a story of how two brothers both get share of their fathers inheritance. The father warns them to be careful of how they use such money, but only one son listens, as the other ignores his father and adventures what the world has to offer (New International Version, Luke 15.11-13). Don’t waste time! Our writers will create an original "A Shipwreck of Faith" essay for you Create order Crusoe is unaware of his actions as he has not developed a deep and prolonging faith within the religion of Christianity itself. In the beginning, he is what most Christians would describe as a lukewarm Christian. What this means in simpler terms is that he knows what Christianity is and says he believes in the customs and aspects of the religion, but does not live his life according to what is being described within the Bible itself. With the current situation that Crusoe is found within the beginning of the story, this takes a toll on his future escapades as his weak mind and faith have him struggle in the battles soon to come. As time moves on, Crusoe soon finds himself shipwrecked with nowhere to go. He is captured by Turkish pirates, who make Crusoe and many others their slaves. Slavery was very popular back in the time of Robinson Crusoe, so it was not looked as a terrible matter as seen today. Still, for Crusoe, he was going through a lot mentally, as one went from slave owner to slave is a drastic change in roles. Seeing this happening, readers could suggest that this was an act of God humbling Crusoe, and showing how little he was compared to everything else that was surrounding him every day, that life could change so quickly in an instant. While captured, he meets another slave, Xury. Later when Crusoe and Xury are rescued by a Portuguese sea captain, Crusoe makes a deal with the captain, to set Xury free on only one condition: he converts to Christianity (Xury in Robinson Crusoe par. 3). With this deal set between the captain and Crusoe, this demonstrates Crusoes first attempt at trying to share his steady growing faith within meeting new people. As God humbled Crusoe, it brought to the surface a different kind of sailor, who wants to show others what life is actually all about. Defoe never went into anymore detail of what happened with Xury and his call to action towards a new religion found upon by both of the slaves, but this was a very large step at the development towards Crusoes journey in the religion. Growing up with his parents, especially his father, Crusoe had a well-established education, not only for the base education taught in most schools, but within the profession his father wanted for him. Stated in the book, my father and mother about their being so positively determined against what they knew my inclinations prompted me to (Defoe 6). The profession of being a lawyer is pretty rough, not only going through and living the job out in your everyday life, but finding the differentiations between everyday people and others who differ in social class. Lawyers have to find certain kinds of tactics, how to use them, and how to apply them to certain people who might think or do things contrasting from the normal way of everyone else. With this in mind, Crusoe was probably taught this by his father, which could help him with his developing faith by putting thoughts to action. As the old saying goes, actions speak louder than words. A big example of this could be seen when Crusoe and Friday are on the cannibalistic island together. As Crusoe rescued Friday from the cannibals, he takes Friday under his ownership and shows him the basics of living, his language, and soon Crusoes second conversion of faith in someone (Robinson Crusoe Theme par. 1).. As Crusoe teaches Friday, Friday soon starts to understand a lot of Crusoes customs, but about his relationship with God. (RobinsonCrusoe par. 33) While this representation goes on, it all ties back to that original saying. Crusoe might have taught Friday about Christianity, but he also did it in action as well, most of the time unintentionally. Friday was rescued by Crusoe, but in the text he was seen calling Crusoe names of power like master (Friday in Robinson Crusoe par. 1). With this being stated, it is being depicted that Friday is under authority or power of Crusoe, much like a slave. As a slave, Friday does whatever Crusoe orders him to. Friday soon gets into this mind set later on, and since he is Crusoes property he wants to follow everything that Crusoe does, including religion. Not only was the religion of Christianity being taught, but Friday wanted to learn more and develop these thoughts to show his master that he respects him who leheartedly. This could be seen as a big leap of faith, not only by Friday, but by Crusoe himself. He is starting to develop a sturdy relationship between him and God, by bringing others to know Him. This is big for Crusoe because it is showing signs of maturity and trust as he is growing through the time spent away from home and stranded on the island. As Robinson Crusoe escapes the island and finds his way back home, he soon discovers that his plantation has made him a reasonably good amount of money. After he obtains his money from the plantation in Brazil, he pays everyone who has cared for him, starting from the Portuguese captain who rescued him and Xury, to one of his widow friends and two sisters. As this is done in the story, this could look to be another religious aspect in Christianity. Tithing is a very important matter in most churches, as it is seen as giving back to the one who has given. This is seen often, as in Christianity it is believed that the love and sacrifice that has been given to the followers of Christ is unpayable, but giving what they have to help the church is portrayed as a temporary way of doing so. This could be compared to Crusoes side of the story, as many have helped him and others along the way of his countless journeys and explorations all without any kind of payment or return at the time. As h e receives his profit, he instantly pays back the Portuguese captain who rescued him from shipwreck. He then returns to England to then repay a widowed friend and his two sisters, another example of Crusoe giving back to the ones who gave to him in the beginning. With evidence from the text and representations that back the religious aspects of Christianity, there is no doubt that Crusoe was struggling with a lack in faith. What Crusoe went through, starting from the beginning seen defying his parents wishes, to encountering new experiences through troubled times, and finally ending it with building a strong foundation in the faith of God shows that it was a tough ride for him all around. Many would say Crusoe has seen it all, but of course he sets back sailing again at the end of the story, adventuring even further to see what more the world has to offer. It is clearly shown in the story that the theme is constant all the way towards the end; No matter how tough the obstacles will become, God is always the guide towards a strong outcome. Crusoe constantly tried and tried again after every attempt to do things by himself, but as soon as he let God take the reins everything started to look brighter again in Crusoes future. As we face troubles and hardships, Crusoe should be a reminder to all that without a strong foundation in faith, it will lead us into many unwanted paths that for some, may be too difficult to endure. Works Cited Defoe, Daniel. Robinson Crusoe. 1719. Print. Shmoop Editorial Team. Robinson Crusoe Theme of Religion. Shmoop, Shmoop University, 11 Nov. 2008, www.shmoop.com/robinson-crusoe/religion-theme.html. RobinsonCrusoe A Born-Again Christian?, www.wordbasedcounseling.org/Articles/RobinsonCrusoe.htm. Shmoop Editorial Team. Xury in Robinson Crusoe. Shmoop, Shmoop University, 11 Nov. 2008, www.shmoop.com/robinson-crusoe/xury.html. Shmoop Editorial Team. Friday in Robinson Crusoe. Shmoop, Shmoop University, 11 Nov. 2008, www.shmoop.com/robinson-crusoe/friday.html. New International Version. Biblica, 2011. BibleGateway.com, www.biblegateway.com/versions/New-International-Version-NIV-Bible/

The Crisis Of The Subprime Loan Scandal - 1222 Words

The overarching problem of profiteering is the foundation of many of the moral problems that came from the exploitation of homeowners during the subprime loan scandal. In the financial industry, Lewis (2010) defines the awareness of top executives and bank loan officers that participated in giving out these loans, More so, an interview with Steve Eisman reaffirms the fully conscious role that loan officers played in allowing individuals without good credit ratings to take these loans. In many cases, the banking industry argued that it was the fault of those that took the loans, but it was actually the facilitation of these loans by loan officers that laid the unethical and immoral foundation of this scandal: Whenever Wall Street people†¦show more content†¦Another aspect of Desmond’s (2016)† research is based on the perspective of the slumlord culture that also exploits low-income people in the city of Milwaukee. A focus on the behaviors of local landlords also exposes the lack of social responsibility on the part of predatory property owners that only seek to extract money from the most impoverished sectors of society. Another aspect of the unregulated financial markets reveals the predatory nature of landlord policies, specially in the case of Tobin in Desmond’s (2016) sociological examination of real estate culture in urban Milwaukee. On one hand, Tobin would be lenient with renters in his trailer park when they were temporally unable to pay rent, but on the other hand, a lack of written contracts also defines the underhanded method of manipulation and control that he wielded over his renters. Much like the banking institution loan officers that never che cked the credit background of a subprime loan recipient, so does Tobin ignore contractual and legal obligations when managing his tenants in the trailer park: Tobin’s negotiations with tenants were rarely committed to writing, and sometimes tenants remembered things differently from the way Tobin did. A tenant would say she owed $150 and Tobin would say it was $250 or $600. Tobin once forgot tht a tenant paid a year’s worth of rent in advance after winning a worker’s compensation claim. Trailer park residents had a word for this: being â€Å"Tobined.† InShow MoreRelatedThe Subprime Loan Crisis : An Analysis Of The Ethical Shortcomings1495 Words   |  6 PagesThe Subprime Loan Crisis An Analysis of the Ethical Shortcomings Ten years ago the US housing market was booming and with a constant rise in prices there didn’t seem to be an end in sight. 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In the years leading up to 2008, buying and selling mortgages became a very

The Failure of Rome’s Economy and the Fall of the Roman...

The failure of Rome’s economy contributed majorly to the fall of Rome. The Roman Economy during the late Republic and Early Empire was based heavily on Agriculture and Commerce. Agriculture in ancient Rome was not only a necessity, but was idealized among the social elite as a way of life. Cicero had considered agriculture to be the best of all Roman Occupations (Sarudy). There had been a lot of trading between the provinces of the empire, and all regions of the empire were largely economically interdependent. Egypt was also important in providing wheat to Rome. Shipments of Egyptian wheat may have amounted to 20 million modii (an Ancient Roman measurement) or more annually. Twenty million modii of wheat was nearly enough for up to half†¦show more content†¦Emperor Domitian, who ruled from 81 A.D. to 96 A.D., had left the Roman Empire in a nearly balanced economy during the greater part of his reign. It is estimated that Domitian’s annual income was more than 1,200 million sestertii (Roman Currency) which over one-third would presumably have been spent maintaining the Roman army. The other major of major expense was the extensive reconstruction of Rome (Domitian). An Emperor that greatly impacted the Roman economy was Nero, who ruled from 54 A.D. to 68 A.D. When complaints arose that the poor were overly taxed, Nero attempted to repeal all indirect taxes. The Senate convinced him that this action would bankrupt the public treasury. In result, Nero cut taxes from 4.5% to 2.5% (Bluejayblog) Government taxes were also ordered to become public. In order to lower the cost of food imports, Nero declared that merchant ships were tax-exempt. After Rome burned in 64, Nero enacted a public relief effort to restore Rome. For a few construction projects, he created the large Domus Aurea and attempted to have a canal dug at the Isthmus of Corinth. 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Why Is Strategy Important to Business Free Essays

Strategy is determining where you are now, where you want to go, and how to get there. It’s the art of developing and implementing specific actions and decisions that will help a business to achieve goals and objectives as set out by the owners. Through the strategy process the overall direction of the company is set including opportunities and threats in the outside world and the internal strengths and weaknesses of the business. We will write a custom essay sample on Why Is Strategy Important to Business? or any similar topic only for you Order Now The aim of a business strategy is to ensure that the threats posed by the external environment are minimized and also to strengthen or minimize the effects of internal weaknesses. The opportunities and strengths are combined to ensure maximum productivity is achieved. The more the external environment changes the more opportunities there are likely for a company, but the more threats can be a problem if not prepared for. Without a business strategy, it would be difficult for an owner to realize the opportunities available. Strategic management in contrast with business policy is a technical approach having a five step model. Strategic management is a set of managerial decisions and actions that determine the long run performance of a company. Strategic management does not have to be just a part of the top level management. In fact, organizations should focus on strategic management at all levels of the management and the entire hierarchy of the company should be involved both in planning and implementation phases. Strategic management does not have to be just a part of the top level management. Organizations should focus on strategic management such that all levels of the management and the entire hierarchy of the company should be involved in both planning and implementation phases. The basic theory of strategic management involves the action of creating and implementing plans and decisions for the benefit of the organization and to help it achieve its objectives. Strategic management analyzes and scan the environment therefore future problems and opportunities could be well anticipated, allows a firm’s top management to anticipate changes and provide track and control for the enterprise, provides clear cut mission, also gives vision and objectives of enterprise to all the employees, and allows a firm to take decisions on the basis of long range forecast. Your vision must be future-oriented it has to paint a picture that attracts employees through the use of visual imagery. This is what gets people hooked into buy-ins, followership, and implementation of the plan. So it means you just have to think outside the box of today and describe the world of the future. With the current situation of the economy the fact that some business establishments have stopped their operation and closed has caused big drop in small and large corporations. Having back up plan would be the idea should the original plan does not succeed businessmen could have alternative options or backed up plans for the continuity and success of the business. Since businessmen are knowledgeable of the business processes, they know the importance of having back up plans. Without a contingency plan, and should the original plan fail, businessmen and the business could experience a lot of trouble. The Mission statement is important because it describes the reason for the existence of an organization and what it hopes to achieve. While mission statements vary from organization to organization and represent the distinctness of each one, they all share similar components. Before writing a mission statement, leaders in the organization must have an idea of what is in store for the future. This vision is the foundation for the mission statement. Most statements include descriptions of the organization’s target market, the geographic domain, their concern for survival, growth and profitability, the company philosophy, and the organization’s desired public image. In addition, it is a tool that conveys the goal, organizational structure and strategy, legitimacy, values, participation and ownership among employees, leadership, responsibility to the community, ethical priorities, and commitment to the public and stakeholders. The organizational decision making process involves proper and efficient implementation of strategic plans and methods to achieve desired business objective. Often one difficulty facing an organization is that multiple divisions are involved in the overall decision making process. Making a decision can have different implications for each respective division. With the businesses of today facing shortened product lifecycles, organizations are facing intense competitive pressures to build market share to stay ahead of rivals. Process changes, the introduction of new and improved disruptive technologies are forcing companies to adapt new business goals and objectives in shortened timeframes, this alone places stresses on all organization. References Karger (1991). Strategic Management: The Key to Corporate Success Pearce II, J.A., Robinson, Jr., R.B. (2008). Strategic management. (11th ed.). New York, NY: McGraw-Hill. Mission and Vision Statements http://www.referenceforbusiness.com/management/Mar-No/Mission-and-Vision-Statements.html#ixzz1LGZLzvbI How to cite Why Is Strategy Important to Business?, Papers

The Vision Essay Research Paper Most people free essay sample

The Vision Essay, Research Paper Most people are disbelieving about psychics and psychic powers. In the book The Vision by Dean Koontz, there arises a existent convincing psychic Mary, who has visions of slayings that are yet to go on. But, a new turn to the narrative causes Mary to see a different sort of vision. Murders more ghastly than of all time. More hard to see. Harder to prosecute. All these factors cause the reader, and perchance Mary to inquire who are the 1s who truly care for her. Can the liquidator perchance be person she loves? Or possibly a haunting truth about the yesteryear. The narrative takes topographic point in assorted locations of modern twenty-four hours California. Some of the narrative takes topographic point in Los Angeles, but the most momentous portion of the narrative takes topographic point in a small town called King # 8217 ; s Point. The town is on the Pacific Coast Highway, and expensive houses dot the shoreline. Refering to the visions, Dean Koontz vividly describes the scene of each of them, as they take topographic point. For illustration, he takes the reader to one of the scenes of a slaying. A little beauty store in Santa Ana, California. He forces the reader to visualize the assorted facets of a normal beauty store, such as, the outside. The neon visible radiations, the thenar trees, the jade-plant hedges, and the money-scented air. He informs the reader of the aroma of the shampoo, pick rinse, Cologne, and sweat. He tells how the floor was covered in hair, and the violet colour of the walls, and the plush purple rug. He describes the sound of the hair drier and the gunfire in which the liquidator shot the teller. As one can see, the writer exhaustively describes the scene. The chief character is of class, the psychic, Mary Bergen. She is the writer of a syndicated newspaper column about psychic phenomena, and the 1 who pursues the visions in which the liquidator creates. The true individuality of the liquidator is non clear until the terminal of the book. Max Bergen, Mary # 8217 ; s hubby, and Alan Tanner, Mary # 8217 ; s brother, each attempt to assist Mary prosecute her visions to catch the slayer, and to liberate Mary # 8217 ; s life of the atrocious emphasis that encompasses her. But Max and Alan don # 8217 ; t acquire along really good. Alan feels that Mary could hold picked a better adult male to get married, because he believes that all Max is after is Mary # 8217 ; s money, and that Max doesn # 8217 ; t truly recognize how delicate she is. Max knows how Alan feels, but evidently he disagrees. Max is reasonably a strong adult male, six inches taller, and forty lbs heavier than Alan. Although Max had promised Mary that he would neer physica lly fight another individual, he feels a strong demand to contend Alan, but knows that won # 8217 ; t halt him from being so chesty. Alan on the other manus, can easy carry people with his sweet voice, and delighting visual aspect. There is besides Dr. Cauvel, Mary # 8217 ; s head-shrinker, and Lou Pasternak, one of Mary # 8217 ; s old friends. Calcium uvel urgently tries to associate Mary’s visions to the yesteryear. Pasternak, an alcoholic journalist, helps Mary and Max seek to happen the slayer, and halt him. Mary Bergen, the well-known psychic, has unfortunate visions of liquidators killing their victims. One twenty-four hours, a awful vision appears with no warning. And from so on, these visions are even more ghastly than her usual visions, and they ever prevent Mary from seeing the slayer # 8217 ; s face. This puzzles Mary, so she goes to her head-shrinker Dr. Cauvel, to seek some replies. He tries to dig into her yesteryear and unveil some truths. She was abused as a kid by one of her neighbours, who purportedly killed all of her brother Alan # 8217 ; s pets. And her male parent died when she was truly immature. Mary doesn # 8217 ; t clearly recall any of the maltreatment she experienced. All she can truly retrieve is the flutter of a batch of wings, like those of a bird. She frequently has visions of merely the wings, and it is an mystery which badly frightens Mary. All she knows, is the wings relate to her maltreatment. Sing that Mary has blocked this portion of her yesteryear ou t, Cauvel believes that her maltreatment is what caused her to get down holding these visions. That same twenty-four hours, a vision comes to her. As she tries to prosecute the vision and see the slayers face, 100s of glass Canis familiariss that the Doctor had collected, flew of the shelves, and at Mary. Scared and confused, Mary subsequently looks for comfort in Max. He of class amenitiess her, and attempts to assist work out the enigma. He takes her to King # 8217 ; s Point where the following slaying is to take topographic point and to run into with Lou, who helps them by holding a sitting to reply some of Mary # 8217 ; s inquiries. A Ouji board assists in replying these inquiries. But, there are a few inquiries asked by the reader, such as: Is the slayer really person who is near to Mary? The Vision by Dean Koontz, is a really exciting book that will most decidedly maintain one turning the pages. It has a batch of suspense, and most of all, a batch of enigma. The writer does a good occupation in allowing the reader cognize what is traveling on at all times. He does so in such an interesting mode, which keeps one beggary for more. The book itself has a really attractive screen which besides gets one # 8217 ; s attending. This book is recommended to about anyone, but largely to those who like slaying enigmas. In reading this book, one will happen that the chief character Mary, goes through many hazards but neer seems to give up. She persists with her visions until she feels that justness has been served. Even though she has some people against her, chiefly the sceptics, she besides has people who love her assisting. This merely shows that even though one might experience like the universe is against them, they really have friends who love them plenty to assist them work out all their jobs. It merely takes continuity and forbearance. 31a