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1 NEW COMPUTATIONAL METHODS FOR 3D STRUCTURE DETERMINATION OF MACROMOLECULAR COMPLEXES BY SINGLE PARTICLE CRYO-ELECTRON MICROSCOPY Dissertation zur Erlangung des mathematisch-naturwissenschaftlichen Doktorgrades "Dr. rerum naturalium" an der Georg-August-Universität Göttingen vorgelegt von Martin Schmeißer aus Hamburg Göttingen 2009
2 Members of Thesis Committee: Prof. Dr. Holger Stark Prof. Dr. Ralf Ficner (First Referee) (Second Referee) Prof. Dr. Erwin Neher 2
3 Affirmation I hereby declare that this thesis has been written independently and with no other sources and aids than quoted. Göttingen, Martin Schmeißer 3
4 Publications and Software Publications Parts of this thesis have been published, submitted to publication or presented at conferences: Schmeisser, M., B. C. Heisen, M. Luettich, B. Busche, F. Hauer, T. Koske, K.-H. Knauber and H. Stark (2009). "Parallel, distributed and GPU computing technologies in single particle electron microscopy." Acta Crystallogr D Accepted Manuscript. Schmeisser, M., B. C. Heisen, H. Stark, D. Moser and A. S. Frangakis (2008). Parallel processing with graphics cards. Workshop. New Algorithms in Macromolecular Crystallography and Electron Microscopy. J. P. Abrahams. Leiden, NL. Schmeisser, M. and H. Stark (2009). Method and device for recording and processing projection images. PCT/EP2009/ M.-P.-G. z. F. d. W. e.v. Germany, European Patent Office. Luettich, M., B. Sander, B. C. Heisen, M. Schmeisser, B. Busche and H. Stark (2008). 3D Structure Determination of Dynamic Macromolecular Complexes by single particle cryo- EM. New Algorithms in Macromolecular Crystallography and Electron Microscopy. J. P. Abrahams. Leiden, NL. Hauer, F., N. Fischer, U. Kothe, M. Schmeisser, M. V. Rodnina and H. Stark (2009). An extended ribosomal L7/L12 stalk and ribosomal expansion segments on the large ribosomal subunit of Thermotoga maritima as revealed by 3D cryo-em. To appear in Journal of Structural Biology. 4
5 Software The following software has been developed within the context of this thesis: AmiraImagic: A plug in for import and manipulation of Imagic data in the Mercury Amira Visualization package. 3DMC: A tool for three-dimensional mass centering of models. ImportExport: A tool for storing and administration of EM Projects and Data in a relational Database. ClustWatch: A tool for administration of a plurality of computing nodes, administration of computing time, resources and tasks. SmartTray: Framework for volunteer and cluster computing Supervised Alignment Tools: Tools for MRA to multiple sets of references including CCC based partitioning of results. 3DView: A 3D viewer to compare, visualize and analyze a plurality of 3D models EulerLib: A library for storing, calculating and manipulating projection directions ClustCMD: Command Line Tools for ImportExport, Clustwatch and SmartTray for scripting use 5
6 Table of Contents Affirmation... 3 Publications and Software Publications... 4 Software... 5 List of Figures... 9 List of Tables List of Abbreviations ABSTRACT ACKNOWLEDGEMENTS Introduction Methods for structure determination of macromolecular assemblies Single particle electron microscopy Crystallographic methods Fundamentals of light microscopy Optical transfer function Resolution Electron microscopy Electron properties Electron optics The Fourier transform Image formation Phase contrast Phase-contrast-transmission function (PCTF) Single particle analysis Image acquisition Preparation of data sets for single particle analysis Alignment Multivariate statistical analysis and clustering of aligned images Iterative approach Assignment of projection directions Definition of Euler angles Established methods for assignment of projection directions Angular reconstitution... 52
7 1.5.4 Tilt series Random Conical Tilt (RCT) Orthogonal tilt Reconstruction techniques Fourier based methods Direct methods Iterative methods Refinement via projection matching once a start structure is determined Determination of obtained resolution by Fourier-Shell-Correlation (FSC) Improvement of processing speed for algorithms used in EM Need for speed How to improve computational speed? Divide et impera Hardware architectures and their implications for parallel processing If you can't have a faster CPU use more of them Shared memory computing Farming GPU computing Historical - Limitations alleged by Amdahl's law Data parallel programming Change your code Software standards for shared memory and distributed computing Software standards for GPU programming Aim of the work Materials and methods Software Biochemical methods Chemicals Buffers and solutions Laboratory materials Special equipment Bacterial samples and plasmids EM imaging conditions Preparation of EM samples
8 2.4.1 Preparation of holey carbon film coated EM grids Preparation of EM samples for negative stain electron microscopy Preparation of cryo EM samples Results Towards 3D structure determination of Acetyl-CoA Carboxylase from Cornyebacterium glutamicum Purification of AccBC-AccD1 complex from C. glutamicum strain Δpyc paacbc p-h6d Negative stain EM of Acetyl CoA-Carboxylase complex AccBC-AccD Optimization of purification of AccBC-AccD1 complex from C. glutamicum strain Δpyc paacbc p-h6d1 for EM RCT negative stain EM of Acetyl CoA-Carboxylase complex AccBC-AccD1 after optimized purification Getting ready for the next hardware generation The SmartTray Foundations for GPU computing Getting ready for the next generation of instrumentation New methods for computation of missing cone free starting models Connectivity graph Informative value of connectivity graphs Influences of misclassification and alignment errors on the connectivity graph and assigned projection directions Separation of conformational differences Influence of alignment and tilt errors on reconstructed starting models Discussion Bibliography Curriculum Vitae
9 List of Figures Figure 1-1 Diffraction Figure 1-2 The convolution of an object and an aberrated PSF Figure 1-3 The PSF for a perfect optical system Figure 1-4 Airy disks for different pupil diameters Figure 1-5 The PSF of a typical human eye depending on the present pupil diameter Figure 1-6 The Transfer of spatial frequencies through an optical system Figure 1-7 Visualization of the Rayleigh criterion for resolution Figure 1-8 Aberration correction in light optics Figure 1-9 Spherical aberration correction in electron optics Figure 1-10 Zernike like phase plates in electron optics Figure 1-11 Transformation of phase contrast to amplitude contrast Figure 1-12 The pctf at different defoci Figure 1-13 A biological macromolecule in different conformations Figure 1-14 Projection views generated by conformation Figure 1-15 Projection views generated by conformation Figure 1-16 Particles in the specimen Figure 1-17 Example of raw data Figure 1-18 Preparation steps for further digital images processing Figure 1-19 Translational and rotational degrees of freedom in 2D Alignment Figure 1-20 References for the "reference free" alignment scheme Figure 1-21 Alignment and class averages Figure 1-22 Iterative alignment and classification Figure 1-23 Angular reconstitution Figure 1-24 Random Conical Tilt Figure 1-25 Orthogonal Tilt Reconstruction Figure 1-26 Projection matching Figure 1-27 A healthy FSC Figure 1-28 Examples of unhealthy FSC plots Figure 1-29 The von Neumann Architecture Figure 1-30 Flynn s Taxonomy Figure 1-31 GPU Hardware Architecture Figure 1-32 Amdahl s law Figure 1-33 Gustafson-Barsis law
10 Figure 2-1 Cryo microscope CM200 FEG Figure 2-2 The "Sandwich" preparation technique Figure 3-1 Acetyl-CoA Carboxylase Gel Filtration Plots Figure 3-2 Spot Scan of fractions between 45 and 55 ml from A-CoA gel filtration Figure 3-3 Typical 2D MSA based A-CoA class averages from A-CoA Figure 3-4 Comparison of typical 2D A-CoA class sums Figure 3-5 Spot scan of gel filtration peak ml from A-CoA gel filtration Figure 3-6 Tilted Pairs from A-CoA Figure 3-7 2D class sums from A-CoA obtained after GraFix Figure 3-8 Comparison of 2D class sums from A-CoA Figure 3-9 Artificial 3D models from A-CoA Figure 3-10 Artificial models of the likely overall organization of AccBC-AccD1 complex.. 96 Figure D class sums from A-CoA, cryo in holes Figure 3-12 The Workflow of the SmartTray Framework Figure 3-13 A list of Nodes running the Smart Tray is kept on the Master server Figure 3-14 FEI Titan Cryos cryo electron microscope Figure 3-15 Schematic illustration of tilt and class sum relations of projection images Figure 3-16 Definition of spherical polar coordinates Figure 3-17 The first untilted image (UT1) used as the starting point for the algorithm Figure 3-18 Alignment untilted side Figure 3-19 Tilt from untilted to tilted side Figure 3-20 Alignment on untilted side, tilt from untilted to tilted side Figure 3-21 Alignment untilted, tilt from untilted to tilted side, alignment tilted side Figure 3-22 Tilt from untilted to tilted side, alignment on tilted side Figure 3-23 Tilt untilted to tilted side, alignment tilted, tilt tilted to untilted side Figure 3-24 Alignment untilted, untilted to tilted, alignment tilted, tilt tilted to untilted. 110 Figure 3-25 First step of the new approach Figure 3-26 Building up of a connectivity graph and assignment of projection directions. 113 Figure 3-27 Relationships utilized for connectivity graph and projection directions Figure 3-28 Sample connectivity graph obtained Figure 3-29 Validation of circular subgraphs Figure 3-30 Connectivity graph for perfect classification Figure 3-31 Connectivity graph systematic miss classification Figure 3-32 Connectivity graph cross miss classification
11 Figure 3-33 Models from RCT and new method reconstruction Figure 3-34 Influence of alignment and tilt inaccuracy Figure 3-35 Quality of models with known alignment and tilt inaccuracy Figure 3-36 Reconstructed models quality
12 List of Tables Table 2-1 Software used within this thesis Table 2-2 Chemicals used within this thesis Table 2-3 Buffers and solutions used within this thesis Table 2-4 Laboratory materials used within this thesis Table 2-5 Special equipment used within this thesis Table 2-6. Bacterial samples and plasmids used within this thesis Table 2-7 Magnification settings used for recording of data sets Table 2-8 Magnifications errors for magnifications from Table Table 2-9. Effective magnifications resulting from Table
13 List of Abbreviations 2D Two-dimensional 3D Three-dimensional Å Angstrom (1Å = 1-10 m) ACo Acetyl Coenzyme A ART Algebraic Reconstruction Technique CCC Cross Correlation Coefficient CCD Charge Coupled Device CPU Central Processing Unit cryo-em Cryogenic Electron Microscopy DFT Discrete Fourier Transformation FFT Discrete Fast Fourier Transformation FSC Fourier Shell Correlation GPU Graphical Processing Unit HAC Hierarchical Ascending Classification HPC High Performance Computing ICA Independent Component Analysis MPI Message Passing Interface MRA Multi Reference Alignment MSA Multivariate Statistical Analysis NA Numerical Aperture OTF Optical Transfer Function OTR Orthogonal Tilt Reconstruction PCA Principal Component Analysis PCTF Phase-Contrast Transfer Function PSF Point Spread Function RCT Random Conical Tilt SIMD Single Instruction Multiple Data SIMT Single Instruction Multiple Threads SIRT Simultaneous Iterative Reconstruction Technique SISD Single Instruction Single Data TEM Transmission Electron Microscopy 13
14 14
15 ABSTRACT New computational methods for 3D structure determination of macromolecular complexes by single particle cryo-electron Microscopy In cryo-electron microscopy (cryo-em) single-particle reconstruction, projection images of many first order structural identical copies of the specimen of interest are combined to recover the underlying three-dimensional object. The analyzed particles are embedded in vitreous ice in a hydrated state at many random orientations and projection images are recorded using a transmission electron microscope (TEM). The heterogeneity of twodimensional projection image data resulting from the co existence of different conformational or ligand binding states of a macromolecular complex remains a major obstacle as it impairs the validity of reconstructed density maps and limits the progress towards higher resolution. Nevertheless, single particle cryo-em is the method of choice for structure determination of large macromolecular complexes that are difficult to purify in the amounts and quality needed for X-ray crystallization. In contrast to other structure determining techniques, single particle cryo-em is not an ensemble technique. Images of individual macromolecular complexes are recorded that can potentially be used to detect structural variations within a single dataset which is highly important for structure determination of large and dynamic macromolecular assemblies. However, computational tools to sort large datasets of images into subpopulations representing images that belong to a unique 3D object are still missing. In structural analysis of macromolecular assemblies with a high degree of structural dynamics, separation of conformations is a challenging task because for each particle image there are six degrees of freedom (translations and rotations), an unknown number of conformations and a high level of noise that have to be considered simultaneously. Classical random conical tilt reconstruction, utilizing additional experimental information by recording of tilted image pairs generates single class sum starting models, limited by a missing cone of information and a low Signal to Noise ratio originated at the limited number of class members and thus projection views used for reconstruction. A new method is presented, based on acquisition of tilted image pairs under cryo conditions facilitating classification of both untilted and tilted raw images. In these double-classified tilted pairs in combination with their experimental tilt angular relationships of the classified particles, angular relationships relative to each other are conserved. This information can be utilized to compute multi class sum starting models without a missing cone. Additionally, the information obtained by classification and tilt relationships can be used to separate particles in different conformations into individual models. The novel strategy is intended to gain insight into the 3D structures and structural diversity of so far poorly characterized dynamic assemblies. Increased computational demands, originated at the increased number of projection images to process in order to analyze conformational differences, are addressed and strategies to overcome these using recent developments in information technology such as Multicore-, Parallel- and GPU- processing are presented. Each parallel processing paradigm alone can improve overall performance, while combining all paradigms, unleashes the full power of today's technology. The increased computational performance thus makes certain applications feasible that were virtually impossible before. 15
16 ACKNOWLEDGEMENTS I would like to express sincere appreciation to my thesis committee: Professor Dr. Ralf Ficner, Professor Dr. Erwin Neher and Professor Dr. Holger Stark. Thanks for giving me the opportunity to be part in this interesting research and thanks for your guidance and inspiration. In addition, I would like to thank Burkhard Heisen in special and also Niels Fischer, Florian Hauer, Björn Sander, Prakash Dube and Dietmar Riedel for interesting and valuable discussions. Further thanks go to Professor Dr. Klaus Weber for immeasurable biochemical experience and Kathy Gelato for native English skills. I thank Tobias Koske, Karl-Heinz Knauber and Mario Lüttich for hardware and software support and the fulfillment of my sometimes extraordinary demands and ideas. Thanks also go to Gordon Freeman, Percival Cox, John Dorian and Christopher Turk for interdisciplinary discussions. I would also like to thank all members of the Stark, Riedel and Lührmann labs for providing biochemical assistance, courtesy in many ways and their pleasant company. Last but not least I would like to thank my dog Bonzo for bearing company and keeping my feet warm during the writing of this thesis as well as the rest of my family who made it possible for me to present this work. 16
17 1 Introduction If you want to understand function, study structure. (Crick 1988) Structure without function is a corpse, function without structure is a ghost. (Vogel and Wainwright 1969) To understand biological processes apart from the functional analysis, the determination of the structure of proteins and macromolecular complexes is a key issue. There are several well established methods for the determination of protein structures. 1.1 Methods for structure determination of macromolecular assemblies Single particle electron microscopy Especially for structure determination of large, possibly asymmetric and biochemical and or conformational homogeneous complexes that are difficult to purify in the amounts and quality needed for X-ray crystallization, single particle electron microscopy is the method of choice (Frank 1996; Frank 2002; van Heel 2002; Stark and Luhrmann 2006; Leschziner and Nogales 2007). It is not an ensemble technique and it does not rely on the formation of protein crystals. The only prerequisite is the presence of a sample containing the purified protein of interest in a moderate concentration. Today this method routinely delivers threedimensional structures of macromolecular complexes at 4-20 Angstrom resolution. Even for large, asymmetric macromolecular complexes, a visualization of e.g. α-helices (Golas, Sander et al. 2003), conformational changes upon ligand binding (Golas, Sander et al. 2005) or trnas bound to translating ribosome (Stark, Orlova et al. 1997) can thus be realized. The instrument used, a transmission electron microscope (TEM), delivers two-dimensional projection images of macromolecular assemblies. Many structurally identical copies, in first order congruent in three dimensions, exist in the specimen analyzed. The three-dimensional structure of an object can be reconstructed from a series of two-dimensional projections along different directions. The mathematical theory for determining a function from its projections was originally derived by Radon (Radon 1917) leading to the development of several reconstruction algorithms used in e.g. tomography and electron microscopy, as reviewed in (Budinger, Gullberg et al. 1979). In electron microscopy, bright field weak phase contrast images are projections of the specimen oriented randomly within the sample, resulting in different projection directions. A series of micrographs can thus be used for a 17
18 three-dimensional reconstruction (DeRosier and Klug 1968) if the projection directions of individual projection images can be identified Crystallographic methods Although arguably the best standard method for structure determination at atomic resolution is X-Ray crystallography, it still depends on the crystallization of the protein to study. Not only crystallization but also data analysis is impeded if the protein studied is either flexible or undergoes biochemical or conformational changes. In these cases if a crystal can be formed at all, the analysis of diffraction patterns makes the intensities and thus the waves amplitudes accessible but not the waves phases. Another crystallographic method is electron crystallography which depends on the existence of two-dimensional protein crystals. These can be analyzed using an electron microscope. In diffraction mode, diffraction patterns that can be recorded as in X-Ray Crystallography represent the amplitudes of the diffracted electron waves. The phases can be accessed by using the electron microscope in imaging mode. The information can be combined for structure determination, thus solving the phasing problem for electron microscopy, but the formation of a two-dimensional crystal is still necessary. 1.2 Fundamentals of light microscopy There are several similarities between light microscopy and transmission electron microscopy. Therefore, fundamentals of light microscopy are introduced first. Due to the wave-particle duality, depending on the problem analyzed, either geometric or wave optics can be used for Figure 1-1 The image produced when imaging a point like object is the so called point spread function (PSF). Due to diffraction a blurring occurs in reality. 18
19 description in light microscopy. Geometrical optics describes the propagation of light, ignoring the wave nature of light, as illustrated in the left image of Figure 1-1. In reality, a vast number of factors are responsible for the variations of the dimension and geometry of the image of a point light object produced by an optical system (Abbe 1873). First of all, the influence of diffraction is analyzed. The point spread function (PSF) describes the image produced when imaging a point-like object as shown in the right image of Figure 1-1. As a microscope is a linear spatial invariant optical system, its imaging properties are completely defined by its PSF. b( r) o( r) h( r ) In case of incoherent image generation, the image b(r ) produced by a linear spatial invariant system is given by the convolution of the object function o(r ) and the intensity PSF h(r ) as illustrated in Figure 1-2. o( r ) h( r r ) dr Equation 1-1 Figure 1-2 The convolution of an object in this case the capital letter E and an aberrated PSF in this case the PSF of a human eye. Figure modified from (Yoon, Jeong et al. 2004) The PSF for a lens of lower opening angle is the Airy disc, which is the Fraunhofer diffraction pattern for a circular pupil as shown in Figure 1-3. In an ideal system it only depends on the diameter d of the pupil (Figure 1-4) which is illustrated in Figure 1-4. Figure 1-3 The PSF for a perfect optical system is the Airy disc, which is the Fraunhofer diffraction pattern for a circular pupil. 19
20 Figure 1-4 Airy disks for different pupil diameters d. In reality there is no ideal optical system. In reality there are imperfections in the optics that influence the behavior of the PSF. When imaging with an aberrated PSF, the image will be aberrated. An example is given in Figure 1-5. Figure 1-5 The PSF of a typical human eye depending on the present pupil diameter. This corresponds to the present adjustment of the iris. Figure modified from (Yoon, Jeong et al. 2004) Optical transfer function Apart from the PSF, the optical transfer function (OTF) is often used to characterize the properties of an optical system. The image generated by the system is given by the convolution between the object and the PSF like shown in Figure 1-2. The frequency spectrum of the 20
21 image F( b( r )) is given by the product of the object s frequency spectrum and the Fourier transform of the PSF F( h( r )) which is called OTF. F( b( r )) F( o( r ) h( r )) F( o( r )) F( h( r )) Equation 1-2 Therefore the OTF represents the spectrum of spatial frequencies that are transmitted by the system. If the OTF is zero or nearly zero for some spatial frequencies, these frequencies are barely transmitted. This means that details of the object which lie in this spatial frequency range cannot be resolved. The OTF can be interpreted as a measure of the reduction of contrast from object to image which is illustrated in Figure 1-6. Contrast is the difference in intensity for two different objects. Figure 1-6 The transfer of spatial frequencies through an optical system is described by the OTF. Frequencies where the OTF equals zero cannot be transmitted. Structures in this spatial frequency range cannot be resolved. 21
22 1.2.2 Resolution A reasonable definition of resolution describes the constraints necessary to distinguish an object from its neighboring object. Figure 1-7 shows an illustration of the Rayleigh-criterion which states that two objects can be discerned when the maximum of the diffracted image of one object correlates with the first minimum of the diffracted image of the other object. Figure 1-7 Visualization of the Rayleigh criterion for resolution. This is fulfilled when the object to neighbor distance d is: 0.61 sin Equation 1-3 where λ is the wavelength of the light used, n the refractive index and α, the half cone of opening angle of the objective used. The denominator sin in Equation 1-3 is called numerical aperture (NA). Resolution of a light microscope is limited by the wave length of the light used as shown in equation 1.3 due to diffraction. This limits the light microscopes resolution to a resolution of 200 nm. Although methods have been developed recently, circumventing this limitation in the case of laser fluorescence microscopy e.g. in (Denk, Strickler et al. 1990; Klar, Jakobs et al. 2000), these methods rely on fluorescent dyes, possibly in combination with antibodies for image generation. The dimension of these dyes chromophores alone lies in the magnitude of proteins, feasible for studies of e.g. cell organelles. The study of protein structures yet is still unpromising using these techniques. 22
23 1.3 Electron microscopy In electron microscopy, instead of photons, electrons are used for imaging. The electron wavelength depends on the particle velocity an thus the acceleration voltage applied and is much smaller than the wavelength of light. Electrons are focused by electromagnetic fields used to form electron lenses. These features combined result in the property that electron microscopy is limited due to the construction of the electron microscope imaging system, not due to a limitation by diffraction in contrast to light microscopy. The properties of electrons and electron optics are now introduced in detail Electron properties Charged particles or point charges experience the so called Lorentz force: A where and A and therefore t v A Equation 1-4 In the presence of electromagnetic fields, where q is the charge, is the electrical field vector, is the velocity vector of the charge and is the magnetic field vector. The lens effect of an axially symmetrical electric field as well as that of a rotational symmetric magnetic field can be derived from Equation 1-4. In case of an axially symmetrical electric field, the equipotential surfaces are figures of revolution. They intersect the axis at right angles, and in the neighborhood of their vertex they are approximately spherical. If the space between these surfaces is imagined filled with a medium of a refractive index proportional to the field strength, a succession of lenses is obtained, somewhat distorted outside, but of the correct shape near the axis. As a combination of aligned lenses is again a lens, the imaging effect of the electric field is immediately evident. The explanation of the magnetic type of 23
24 lens is somewhat more complicated. The Lorentz force exerted by a magnetic field on a moving electron is at right angles both to the direction of the field and to the direction of motion. The effect is a twisting or equipotential surfaces rotation of the electron trajectories around the axis, an effect which has no counterpart in ordinary optics. The magnetic vector potential is zero at the axis and increases approximately linearly with the radius r. This means that the equivalent potential decreases outside the axis and that the magnetic field has a repellent effect, i.e., it drives the electrons back, toward the axis. Therefore, a magnetic lens is always a condensing lens. Of the numerous defects which lenses can have, only three are of interest in microscopy: the spherical aberration, the chromatic aberration, and the axial coma. Spherical aberration, the most important of the lens defects in electron microscopy, means that an axial point is imaged as a point strictly speaking only while the imaging rays are infinitesimally close to the axis. These are called paraxial rays, and their intersection with the axis is the paraxial or Gaussian image. At larger angles α, the intersection moves away from the Gaussian point by a distance which is called the longitudinal spherical aberration, and which in first approximation is proportional to α². A point will be imaged as a disk of a diameter proportional to α³. This diameter is called the transversal spherical aberration. These characteristics can be applied to use electrons in an imaging system comparable to light microscopy (Ruska and Knoll 1931) and was used by Ernst Ruska in Berlin in 1931 to build the first electron microscope (Knoll and Ruska 1932). Starting with a 17-fold magnification, the electron microscope potential has been constantly increased and nowadays delivers resolutions ~ 0.5 Angstrom in Transmission mode. Due to the electrons much smaller wavelength, in EM a much better resolution than in light microscopy can be achieved. The electron wavelength λ depends on the acceleration voltage used following De-Broglie equation: Equation 1-5 where h is Planck's constant, m the mass and v the particles velocity. For an electron this leads to: 24
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2006-06-27 at 11:12:00 am #15867
Dell takes high road to Scotland
DELL, the US computer giant, is to "significantly ramp up" its operations in Glasgow and create jobs in Scotland rather than in Ireland, where it has its European manufacturing base.Since it opened last year the company's Glasgow centre has handled UK-only sales and technical support calls from medium and large business clients, as well as the public sector.This is now to include European, Middle East and African (EMEA) business.It means that a planned steady build-up of staff of around 400 now to 850 by 2008 will easily be surpassed at the Glasgow base at the city's former Imperial Tobacco factory, which has been supported by up to £7.5m of Regional Selective Assistance.Gartner analyst Malcolm Hancock said he expects Dell to fully tap into a "surplus" of highly skilled staff in Scotland who have become casualties of a once flourishing Silicon Glen.Dell is Ireland's biggest IT employer, but Hancock claimed: "A readily available skilled workforce nearby is a big mover." Ireland has to import around 80,000 people a year to fill jobs. "So one can see where Dell is heading for future business needs."A low rate of corporation tax in Ireland has proved a prime mover in the past for the likes of Dell and Microsoft to locate major operations there.However, the Irish Business and Employers Confederation's third national survey claims that the cost of doing business in Ireland is now "spiralling out of control and threatening the viability of both indigenous and multinational companies."According to the IBEC, quoted in Business Limerick, the Irish economy lost 7,700 jobs nationally in the first four months of 2006.Dell director (software and peripherals) Michael Buck said: "A significant ramp-up is to occur in Glasgow which we are growing fast, especially when it comes to technical support expertise."I can confirm that whilst the focus during the past year has been UK-only, from now on EMEA is to be included in our strategy. I cannot say yet by how much exactly, but further new jobs are to come in Glasgow on top of what we have previously announced."Texas-based Dell is currently under pressure worldwide following a sharp drop in its first quarter profits this year, and it has seen its share price drop by 40% in the past year.Buck said that they were involved in a constant drive to cut costs and gain premium prices for their vast range of pcs, notebooks and servers plus a highly lucrative printer market, which Dell entered three years ago.Dell is keeping the launch of its new laser printers range – due out this week – firmly under wraps, so as not to give dominant player Hewlett-Packard any competitive advantage.Hancock said the firm was spending $180bn globally in this sector, of which one third is European based. The UK accounts for a fifth of the European figure of $60bn.

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text:
A hot new television series
- Mario
Camp Chef
- Chef Kawasaki
Screaming Boos
- Luigi
- Zelda
- Paratroopa
- Koopa Troopa
- King Dedede
- Link
- Bowser Jr.
- Pikachu
- Lucas
- Lanky Kong
- Diddy Kong
Killer Goombas
- Jigglypuff
- Peach
- Bowser
- Donkey Kong
- Daisy
- Kirby (43rd place)
- Yoshi
- Falco
- Shy Guy
- Birdo
- King Boo
Rabid Spinies
- Gliscor
- Krystal
- Slippy
- Fox
- Olimar
- Mr. Game and Watch
- Sonic (44th Place)
- Tails
- Dixie Kong
- Ness
Episode 1 - Dive of Doom
- Mario: Welcome to Total Game Island! Where 44 campers must compete to win $1,000,000! Today is the first Challenge but first things first! Breakfast!
(In The Kitchen)
- Diddy: What is this stuff?
- Zelda: I think it's paste mixed with yuck!
(In confessional Cam)
- Zelda: The food here is the most disgusting I've EVER tasted! and ive eaten poo! You hear that Ma! I was the one that ate poo that was on the table!
(Back in the Kitchen)
- Mario: Good Morning Campers! Are you ready for the first challenge?
- Everyone: No!
- Mario: Too Bad! Let's Go!
- Peach: What do you think the first challenge is gonna be?
- Shy Guy: Who Knows? It's our first challenge how hard is it gonna be?
(At the cliffside)
- Bowser: Aw! there's no food!
- Mario: Well theres no food! but there is water! the challenge is to jump into that itsy bitsy ring over there!
(Challenge ends)
- Mario: Well looks like I will be seeing the Rabid Spinies at the Campfire tonight as only 2 of them jumped that was Olimar and Fox.
(At the Campfire)
- Mario: On this plate I Have 10 marshmallows to give you, but there are 11 of you, the person who doesn't get a marshallow must leave immediately. Here we go.
List of Characters who got a marshmallow in order:
- Fox
- Olimar
- Krystal
- Tails
- Gliscor
- Dixie Kong
- Mr. Game and Watch
- Ness
- Slippy
- Mario: So Sonic, Au revoir!
- Gliscor: Adios!
- Ness: Sayanora!
- Tails: See Ya!
Episode 2 - Raving Racers
- Mario: Campers! Your challenge starts in ten! Meet me at the forest!
- Paratroopa: (to Diddy) What torture does he have for us this time?
- Diddy: I have no idea, but it might be something that makes us wish we never came here!
(at the forest)
- Mario: Welcome Campers! Yor challenge today is to race from here all the way to the mess hall! On your marks, get set, go!
Everyone starts running except Daisy who is painting her nails, when she realises that everyone is gone, she starts running.
- Slippy: Owch! I fell over!
Daisy runs past
- Slippy and Daisy are neck to neck and they both run past a lot of people and come in 2nd and 3rd place.
Race Places: Top 5:
- 1st: Diddy Kong
- 2nd: Daisy
- 3rd: Slippy
- 4th: Luigi
- 5th: Olimar
Race Places: Bottom 5:
- 39th: Peach
- 40th: Game and Watch
- 41st: Jigglypuff
- 42nd: Shy Guy
- Last: Kirby
- Mario: Looks like im seeing the Killer Goombas at the campfire tonight!
List of characters in order of obtaining marshmallow:
- Daisy
- Bowser
- King Boo
- Donkey Kong
- Falco
- Yoshi
- Birdo
- Peach
- Jigglypuff
- Kirby: Awww, that was too soon.
- Jigglypuff: Sorry, Kirby but you did lose it for us
- Kirby: I guess thats fair see ya guys!


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Surrounded by verdant rows of Swiss chard, eggplant, garlic, beets, amaranth and mesclun, the Wyckoff Farmhouse Museum seems worlds away from its urban perch on Utica Avenue in East Flatbush, Brooklyn. The farm's 1.5 acres, wedged between two busy roads, are the stuff that the dreams of idealistic urban planners are made of—a lush, green space readily accessible to an inner-city community. But Wyckoff is no idle fantasy. Its farmhouse, which dates to 1652, may be a monument to a bygone way of life, but the organic produce harvested on its grounds has been a vital resource for the neighborhood since 2003, when the farmers' market was established. Family-friendly events occur throughout the year, and usually focus on seasonal happenings, like an Apple Festival in September.
|Venue name:
|Wyckoff Farmhouse Museum
5816 Clarendon Rd
|Cross street:
|at Ralph Ave
|Opening hours:
|May–Oct Tue–Sat, Nov–Apr Tue–Sun 10am–4pm
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I work for the federal government and my salary is $70,000 annually, single and no dependents. I was out on medical leave for 14 months without pay or any form of income. I'm just returning to work and before I can get a first pay check, the creditors are lined up with court filings and threats of garnishments. Since I have not had a salary in over a year will I still be required to take the pre-filing credit counseling course? Will future income be considered or the income I earned for the past 6 months which is $0. Are there any good bankruptcy attorneys in the landover Maryland area? how long do creditors have to contest a filing after the petition for chapter 7 is filed. I have over 80,000 in debt which includes unpaid federal taxes from (2007-2010).
You will need to take the credit counseling course - with VERY few exceptions, EVERYONE does. The means test (a mechanical set of assumptions) looks at the past six months income, so you should not have a problem with that. Your reality is what will matter in your filing. As long as your actual reasonable and necessary expenses use up all of your income, you should qualify for a chapter 7. You should probably wait until after your 2009 taxes were due (April 15, 2010 - unless you filed an extension - then Oct 15, 2010), so you file 3 years after they were due, which would likely make all but your 2010 taxes dischargeable. Generally creditors can only contest your bankruptcy if they can show you fraudulently obtained credit from them, which is rare. You should use the attorney finder at www.nacba.org to find a good attorney in your area.Ask a similar question
Yes you will need to still take the credit class, you should file sooner than later as you are correct in stating that the court will look back 6 months for your means test to determine if you qualify to file a chapter 7.Ask a similar question
You do have to take the course. If you file now, you will pass the "means test" which looks at the last 6 months income. But your current, or forward looking income is also considered to determine if your filing under Chapter 7 is abusive under the totality of the circumstances. Given that your 6 month income is below the statewide median income, creditors are limited in their ability to move to dismiss your case, but in any event that window runs until 60 days after your meeting of creditors, which is about 4-6 weeks after filing. You should see an experienced bankruptcy lawyer as soon as possible. And yes, there are good lawyers in your area (we have an office in Greenbelt, for example...)Ask a similar question
Good explanation.
Very good explanation. Time is of the essence in this case, especially with pending litigation.
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Hollyscoop headed to the set of TNT show 'Franklin & Bash' to talk to the shows stars Mark Paul Gosselaar and Breckin Meyer! We got the dirt on Mark Paul Gosselaar's upcoming wedding to the love of his life.
|Mark Paul Gosselaar Reveals Summer Wedding Plans
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Start a Call Center or Bpo Company
Edited by dequan, Eng, jojo, PapaNader and 5 others
- 1 Questions and Answers
- 1.1 Need full steps to setup a tech support BPO?
- 1.2 I want to open my own BPO company in Bangalore, so with whom I need to contact and how?
- 1.3 I am new to call center field and I want to know how to develop customer base and call center?
- 1.4 For a BPO services business plan, what is the table of contents?
- 1.5 BPO operations, Sales, techniques?
- 1.6 Give me a detailed description on how can I start a BPO company in Bangalore India?
- 1.7 How to find customers who wants data entry outsource?
- 1.8 I want to start a BPO/call center and I am unaware of how to get customers for the contract?
- 1.9 Please share project plan for BPO start up business in rural area?
- 1.10 Which sites can you recommend for a start up BPO company like us to get clients and market our services?
- 1.11 I would like to purchase a BPO company of data processing,what should I need to ask the selling person ,before I purchase?
- 1.12 What is the minimum number of seats a call center must have to be able to provide services for customers in USA and Europe?
- 2 Comments
Questions and Answers
Need full steps to setup a tech support BPO?
Tech support BPO setup, costing, and managing in details
The details are outlined in the main VisiHow article about starting a call center or BPO. As for funding, that will depend on how many staff you will need but most start ups require 10x the salaries for employees in the first year. This will include a contingency plan should you not get enough clients to support the business initially. Also, if you can't afford a manager on salary it will be solely up to you to train and manage as well as try to attract clients to the business.
I want to open my own BPO company in Bangalore, so with whom I need to contact and how?
In future I want to open my own company as I am having 4 years experience in airport and currently working with 1 BPO so planing to open my own so I need your help and guidance. I have tried: I am trying to open own but not yet started. I think it was caused by: Might not stable, thinking about one particular business
The first thing you will need to do is register with NASSCOM. Make a comprehensive business plan and because you have doubts about stability, provide steps in the plan to negate this. This will include having in hand a year's salary for each employee you hire.
I am new to call center field and I want to know how to develop customer base and call center?
I want to start my call center in India, how to start or is there any BPO or KPO franchise available?
VisiHow QnA. This section is not written yet. Want to join in? Click EDIT to write this answer.
For a BPO services business plan, what is the table of contents?
As I start to consider building a BPO service, I know I can go in a variety of directions of services. I imagine the on thing common about all approaches is the table of contents on the business plan. What are the probable items in this table of contents?. I have tried: I'm an experienced CEO in a small company. I think it was caused by: No problem
Since you are looking to purchase a franchise, FranchiseIndia currently shows 18 BPO franchises available in the Consultant section of the industry. There are also companies like BPO Project World advertising for franchise buyers. These do not come cheap though so you will need to have the capital and prove you can sustain the business because you will be using their name. It will be higher financial scrutiny than if you were just registering with NASSCOM but at the same time, they can provide you with name recognition and possible clients that would take years to achieve on your own. Look for a company that will provide you with training and potential/initial clients.
BPO operations, Sales, techniques?
How is the market for a BPO startup?. How to start one?. Who is the decision maker in an organization to outsource work related activities in BPO?. How to get clients?
Depending on what country you live in, the government is the initial decision maker on whether you will own a BPO. To start one you need to come up with a business plan and the necessary capital to support a year's worth of salary for all employees. You can hire an intermediary company to gain clients for you or go after them yourself as the business owner. These clients will not seek you out, you will have to hustle like you never have before to get yourself in the door of these companies.
Give me a detailed description on how can I start a BPO company in Bangalore India?
What I am mainly looking for is, how to incorporate my company, what licenses should I get to get the business up and running, how to get in touch with clients
To incorporate in India, you need to fill out the applications and submit to NASSCOM for initial approval. Then you will move forward with Telecommunications India and complete their approval process. For these applications, you will need to provide a business plan.
Getting in touch with clients is all about marketing to gain name recognition. You can send an email with your business profile to prospective clients or go door to door with your brochures to companies that you can provide service to. Keep in mind that you will be not the only one doing this on a daily basis so make your approach stand out above the rest.
How to find customers who wants data entry outsource?
I have my BPO, want customers who want his data entry outsource
Create a profile on Freelancer and bid on projects for data entry. You will also want to make a website with contact information and get some Google Ads directed to your target market. Catchy lines in the advertisement like just $2 a day help with click rates.
Freelancer can help you gain name recognition with reviews/ratings from past clients on Freelancer. You will also get some income coming in while you start to build your regular client list. If you can come familiar with US medical data entry, you may have better success by marketing to the healthcare industry in the US. They largely outsource data entry.
I want to start a BPO/call center and I am unaware of how to get customers for the contract?
How to get a contract for BPO company. I have tried: Nill. I think it was caused by: Insufficient knowledge
It is purely about your sales hustle in terms of getting clients. If you are still unaware of how to go about it, purchasing a franchise with established name recognition and client list might be a better fit for you.
BPO project marketing and getting clients
Only you know the details but you should include:
- 1Number of employees.
- 2Training outline.
- 3Capital Funding.
- 4Start up costs such as leasing office space or machinery.
- 5List of potential clients and their contact person.
- 6Vision of where you want to be in 5 years.
- 7Contingency plans for financial/natural disaster.
Which sites can you recommend for a start up BPO company like us to get clients and market our services?
We are trying to create a Virtual assistant services company similar to the BPO setting but we want to know which sites are we able to post our company description and services to attract possible clients.
Odesk and Freelancer are the giants of the virtual assistant job market. Establish a profile for your company on both websites and bid on projects even if they are small so you can start establishing a rating. You can also look at websites like Kijiji (Canada), Gumtree (UK) and Craigslist (US) to post ads of availability on and reply to job postings.
You can also check out websites like Indeed and Monster to see if there are any postings that you can offer your services on.
I would like to purchase a BPO company of data processing,what should I need to ask the selling person ,before I purchase?
Purchasing a BPO data processing company
You want to see their financial books as well as a statement of revenue projections from clients. You should also ask for permission to speak with their largest clients directly or that they provide you with a contract from the client stating that the client will stay for at least 6 months after the transition of ownership.
What is the minimum number of seats a call center must have to be able to provide services for customers in USA and Europe?
We want to start a call center. Our target market is USA and Europe. What is the minimum number of seats we must have to be able to provide services for customers in USA and Europe? Can we get customers that will sign for at least 6 months contracts and buy call center service per seat?. I have tried: We are working on a Business Plan. I think it was caused by: None
Call Centers in North America hire around an average of 150 people. These are usually for one company such as a credit card or banking institution. Instead of determining what the US market will need, make your list based on qualified applicants that you know you can hire. So if the number is only 10 then go after smaller clients such as a medical office call center that covers doctor pages and messages when the office is closed.
A six-month contract might be attainable if you can prove that you have qualified applicants. Only small clients will agree to a per seat basis usually. Then again small clients can turn into larger clients over time. If you establish a good relationship with them it is likely that as they grow so will you by providing for their needs.
If you have problems with any of the steps in this article, please post in the comments section below.
Article Info
Categories : Communications & Education
Recent edits by: akeko.agba, Lola Todmann, sannidhan


text:
Wessex FM News
Hundreds Of Thousands Join Pension Strikes
1:05am 30th June 2011
(Updated 10:06am 1st July 2011)
Thousands of schools were closed as huge numbers of public sector workers went on strike over Government reforms to their pensions.
There were warnings of fresh industrial action to come after hundreds of thousands walked out in protest, disrupting courts, driving centres and jobcentres.
A row developed between unions and Downing Street over the impact of the strike after the Government said it had "minimal" effect.
As many as 600,000 joined the industrial action, and unions said as many as 20,000 marched in London. Thousands more protested in other cities.
They are against planned reforms which unions say would mean more work and higher contributions for a lower pension.
The Government has questioned the numbers taking part, but one of the four unions involved - the Public and Commercial Services Union (PCS) - said the strike was the best-supported ever.
The Cabinet Office said figures gathered from every Government department showed more than 75% of civil servants were not on strike.
It added fewer than 100,000 civil service workers were on strike - around a fifth of the total workforce.
But the PCS union, which includes police support and border staff, claimed its numbers suggested 84% of its members have gone on strike.
The Metropolitan police said nine out of ten of the force's 999 callers failed to turn up for work, with officers drafted in from other duties to make up the numbers.
Schools bore the brunt of the 24-hour walkout as members of three teaching unions staged walkouts.
The Department for Education said more than 11,000 out of 21,500 state schools in England were affected.
In total there were 5,679 schools closed, with nearly 5,000 partially closed. In Wales more than half the 1,800 schools were closed or partially shut.
The National Union of Teachers says 80% of schools in England and Wales have been affected.
Marches have been largely good-natured, although Sky News witnessed police remove about 10 youths dressed in black hoodies from a rally in central London.
Police in the capital said 30 people had been arrested on suspicion of criminal damage, breach of the peace and drugs offences.
One police officer and six members of the public were injured. Four arrests in Brighton were linked to a protest there.
The Government has criticised the strikes, with Prime Minister David Cameron describing the action as "wrong".
Cabinet Office minister Francis Maude said people needed to wake up to the stark economic realities.
"Those who say pensions are affordable should also say which hospitals to shut down to subsidise their pensions," he said.
"People are living longer, so should work longer."
But general secretary of the TUC, Brendan Barber, said the Government was not playing fair with the cuts.
"I'm a firm believer of negotiations and as I've said, I hope we're going to be able to resolve this through negotiations - without the need for further industrial action," he told Sky News.
Deputy Prime Minister Nick Clegg said the Government was "not seeking to be at loggerheads with the unions".
"I think it's a real shame there are strikes because there are talks ongoing between the Government and the trade unions," he said.
Labour leader Ed Miliband also criticised the mass walkout, saying it was "wrong" - but again accused the Government of "reckless" behaviour.
One protester at a rally in London branded him a "disgrace" for not supporting the action, prompting loud cheers from the audience.
Teacher Angela White told Sky News that while she understands cuts are needed, they should be less severe.
"We're not saying we don't want to take a hit - we understand people are being squeezed," she said.
"What we are saying is, please negotiate and not do all these things at once to us."
Up to 6,000 people joined the pensions protest in Newcastle, a union leader said, and around 1,500 gathered in the centre of Sheffield. A major protest has also taken place in Manchester.
The UK Border Agency warned passengers travelling into Britain would face delays at ports and airports as passport officers join the strike.
Heathrow Airport and the PCS said 70% of border staff at the airport were striking.
Downing Street said there had been "minimal impact" on the public.
"Our border controls are in place and JobCentres and pension offices are open for business," said a spokeswoman.
Special needs teacher Mark Baker, from Rochdale, said a pension of £18,500 that he had expected to receive when he retired at 60 would be cut to £13,500 under Government proposals.
Mr Baker, a regional official for the Association of Teachers and Lecturers, insisted the organisation was moderate but added: "There must be something serious if a union that doesn't strike is taking strike action."
The British Chambers of Commerce said the stoppage would result in tens of thousands of parents having to take a day off work to look after their children.
:: Unions insists public sector workers are facing an unprecedented attack on pensions
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Running Time: 86 minutes
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Books: The Prince by Niccolo Machiavelli, A Clockwork Orange by Anthony Burgess, and anything with Greek Gods, or history
Music: the black dahlia murder, cradle of filth, cannibal corpse, as blood runs black, as I lay dying, at the gates, kataklysm, Lamb of God, behemoth, all that remains, silverstien, rage against the machines, dethklok, mastodon, children of bodom, gwar, amon amarth, the clash, the misfits, the ramones, the buzzcocks, bad religion, no fx, rancid, him, Judas priest, iron maiden, king diamond, arch enemy, meatloaf, arsis, atreyu, and much more. Also if you know starbomb or ninja sex party, I would love to know your favorite song.
Games: all Zelda's, Mario's, metroid, final fantasy's, contras, and just too many to name.
2. Friends
3. Internet
4. Phone
5. Car
6. Games


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go round (something or someone)
Definition of go round (something or someone)
chiefly British
1 : to go or pass from one person to another person at or in (a place) There's a rumor going round the office that the boss is about to get fired.
2 : to be long enough to pass all the way around (something or someone) This belt isn't long enough to go round my waist.
Word by Word Definitions
: to move on a course : proceed — compare stop
: to move out of or away from a place expressed or implied : leave, depart
: to take a certain course or follow a certain procedure
: the act or manner of going
: the height of fashion : rage
: an often unexpected turn of affairs : occurrence
: functioning properly : being in good and ready condition
: a game played between two players who alternately place black and white stones on a board checkered by 19 vertical lines and 19 horizontal lines in an attempt to enclose the larger area on the board
: having every part of the surface or circumference equidistant from the center
: cylindrical
: approximately round
: around
: something (such as a circle, globe, or ring) that is round
: a knot of people
: a circle of things
: to make round
: to make (the lips) round and protruded (as in the pronunciation of \ü\)
: to pronounce with lip rounding : labialize
: around
: all during : throughout
: whisper
: to speak to in a whisper
: some indeterminate or unspecified thing
: a person or thing of consequence
: one having more or less the character, qualities, or nature of something different
: in some degree : somewhat
: used as an intensive giving adverbial force to an adjective
: some person : somebody
Seen and Heard
What made you want to look up go round (something or someone)? Please tell us where you read or heard it (including the quote, if possible).


text:
Gúta (pronounced "Gueta") are a non-sapient extraterrestrial species that was indigenous to the planet Reach, where they were the largest terrestrial species found on-world. They were bipedal, standing roughly 20 or 30 feet tall and while aggression was rare, rutting males would attack if interrupted. They were presumably far too large and violent to serve any exportation purpose, and therefore they were very likely rendered extinct when Reach was glassed by the Covenant in 2552.
Appearances Edit
- Halo: Reach


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It's somewhat telling that the most memorable Georgia Bulldogs moment Justin Scott-Wesley has experienced came before he ever stepped foot on the Sanford Stadium turf.
Two years ago, the redshirt freshman flanker made public his commitment to Georgia from the winner's stand at the Georgia Olympics in Jefferson, just minutes after claiming the Class AA state title in the 200 meters.
"It was a great day for me and for my family," said the 5-foot-11, 218-pound Scott-Wesley, a graduate of Mitchell County High in Camilla. "I won the 100 and 200 and I (committed) after the 200. I committed a week earlier to Coach (Mark) Richt, but I told him I was going to come out with it on the track stand. I had planned it for a long time."
Since that day, there's not been a lot of attention cast Scott-Wesley's way.
But with the season-ending injury suffered by Marlon Brown last Saturday against Ole Miss (coupled with Michael Bennett's ACL tear earlier this fall), there's a good chance that could change in the near future.
Although he's not listed on the No. 5 Georgia's depth chart as the Bulldogs prepare to visit Auburn today with a trip to the SEC championship game in Atlanta on the line, Scott-Wesley could find himself in the middle of the action in a hurry.
The swift Scott-Wesley — who has appeared in four games and caught one pass, a 43-yarder against Florida Atlantic — has toiled in the shadows created by Georgia's cadre of pass-catchers, but he's not grumbling about missed playing time.
"Justin has a great opportunity," Richt said. "He was, of course, eligible to play this year; he wasn't redshirted or anything, but he hasn't had a lot of snaps. Sometimes guys will say, 'I'm not playing this year so I'm just going to go over here and feel sorry for myself and wait until the spring.' Justin has not done that. He's come to work every day to improve. We watch film as coaches, and his effort hasn't gone unnoticed.
"I don't know what's going to happen. I think he's going to play more than he's played, and if you are in the game you could get a ball. We'll call a play and we don't know who it's going to go to a lot of the time. It just depends on the coverage and the protection, so he may get his hands on the ball a couple of times and maybe something good will happen, and (it will) give him some confidence… and give us more confidence
in him."
Scott-Wesley, who was redshirted last year ("Last year, to be honest, I needed a redshirt. I wasn't ready," he said) has a host of teammates looking forward to seeing him make contributions on Saturdays.
"Losing a guy like Marlon is huge for our offense and our receiving corps and I told Justin that it's his time to stand up, and I feel he's embracing it," said flanker Tavarres King. "He's come a ways since fall camp and has gotten so much better. This is his time to shine and I think he can do it."
"I think he can make a huge impact, especially with his size," added quarterback Aaron Murray. "Justin's a very big, strong receiver and that's what we lost with injuries to Marlon and Bennett, two physical receivers. He's a guy who can definitely play that part and go against bigger (defensive backs) and linebackers and create space for us. And he's got speed. Once he gets going, there aren't many guys who can catch him."
Speed is perhaps what Scott-Wesley is best known for, but he wants it to be known that's just one aspect of his arsenal.
"Coming in, I had to show people I was a receiver," he said. "People really didn't acknowledge my skills as a receiver much. So I had to show people I was a football player. Day in and day out, I'm showing people I'm more than just a fast guy. I can catch the ball, I can take a hit, I can give a hit, I can block."
"Yes, he is a speed guy but he's also able to catch the ball," said flanker Rantavious Wooten. "He can take it to the house, he can catch the deep ball, he can go across the middle. I feel he can do everything. That's what we practice each and every day so when stuff like this happens, we have somebody ready. He doesn't have a lot of experience, but his time is now."