Cell wall

Cell Walls - Structure & Function

I. Functions of the cell wall:   The cell wall serves a variety of purposes including:

1. maintaining/determining cell shape (analogous to an external skeleton for every cell).  Since protoplasts are invariably round, this is good evidence that the wall ultimately determines the shape of plant cells.
    
2. Support and mechanical strength (allows plants to get tall, hold out thin leaves to obtain light)
    
3. prevents the cell membrane from bursting in a hypotonic medium (i.e., resists water pressure)
    
4. controls the rate and direction of cell growth and regulates cell volume
    
5. ultimately responsible for the plant architectural design and controlling plant morphogenesis since the wall dictates that plants develop by cell addition (not cell migration)
    
6. has a metabolic role (i.e., some of the proteins in the wall are enzymes for transport, secretion)
    
7. physical barrier to: (a) pathogens; and (b) water in suberized cells.  However, remember that the wall is very porous and allows the free passage of small molecules, including proteins up to 60,000 MW. The pores are about 4 nm (Tepfert & Taylor 1987)
    
8. carbohydrate storage - the components of the wall can be reused in other metabolic processes (especially in seeds).  Thus, in one sense the wall serves as a storage repository for carbohydrates
    
9. signaling  - fragments of wall, called oligosaccharins, act as hormones. Oligosaccharins, which can result from normal development or pathogen attack, serve a variety of functions including:  (a) stimulate ethylene synthesis; (b) induce phytoalexin (defense chemicals produced in response to a fungal/bacterial infection) synthesis; (c) induce chitinase and other enzymes; (d) increase cytoplasmic calcium levels and (d) cause an "oxidative burst".  This burst produces hydrogen peroxide, superoxide and other active oxygen species that attack the pathogen directly or cause increased cross-links in the wall making the wall harder to penetrate.
   
       Let's look at how this system works.  Consider a pathogenic fungus like Phytophthora.  In contact with the host plant the fungus releases enzymes such as pectinase that break down plant wall components into oligosaccharins.  The oligosaccharins stimulate the oxidative burst and phytoalexin synthesis, both which will deter the advance of the fungus.  In addition, the oligosaccharins stimulate chitinase and glucanase production in the plant.  These are released and begin to digest the fungal wall.  Fragments of fungal wall also act as oligosaccharins in the plant to further induce phytoalexin synthesis.  Cool!
    
10. recognition responses - for example: (a) the wall of roots of legumes is important in the nitrogen-fixing bacteria colonizing the root to form nodules; and (b) pollen-style interactions are mediated by wall chemistry.
     
11. economic products - cell walls are important for products such as paper, wood, fiber, energy, shelter, and even roughage in our diet.
    

II. Wall Components - Chemistry
    The main ingredient in cell walls are polysaccharides (or complex carbohydrates or complex sugars) which are built from monosaccharides (or simple sugars).  Eleven different monosaccharides are common in these polysaccharides including glucose and galactose. Carbohydrates are good building blocks because they can produce a nearly infinite variety of structures.  There are a variety of other components in the wall including protein, and lignin. Let's look at these wall components in more detail:
A. Cellulose
   β1,4-glucan (structure provided in class). Made of as many as 25,000 individual glucose molecules. Every other molecule (called residues) is "upside down". Cellobiose (glucose-glucose disaccharide) is the basic building block. Cellulose readily forms hydrogen bonds with itself (intra-molecular H-bonds) and with other cellulose chains (inter-molecular H-bonds). A cellulose chain will form hydrogen bonds with about 36 other chains to yield a microfibril. This is somewhat analogous to the formation of a thick rope from thin fibers. Microfibrils are 5-12 nm wide and give the wall strength - they have a tensile strength equivalent to steel. Some regions of the microfibrils are highly crystalline while others are more "amorphous". 
B. Cross-linking glycans (=Hemicellulose)
    Diverse group of carbohydrates that used to be called hemicellulose.  Characterized by being soluble in strong alkali. They are linear (straight), flat, with a β-1,4 backbone and relatively short side chains. Two common types include xyloglucans and glucuronarabinoxylans. Other less common ones include glucomannans, galactoglucomannans, and galactomannans.  The main feature of this group is that they don’t aggregate with themselves - in other words, they don’t form microfibrils. However, they form hydrogen bonds with cellulose and hence the reason they are called "cross-linking glycans". There may be a fucose sugar at the end of the side chains which may help keep the molecules planar by interacting with other regions of the chain.
C. Pectic polysaccharides
    These are extracted from the wall with hot water or dilute acid or calcium chelators (like EDTA).  They are the easiest constituents to remove from the wall. They form gels (i.e., used in jelly making). They are also a diverse group of polysaccharides and are particularly rich in galacturonic acid (galacturonans = pectic acids).   They are polymers of primarily β 1,4 galacturonans (=polygalacturonans) are called homogalacturons (HGA) and are particularly common.  These are helical in shape. Divalent cations, like calcium, also form cross-linkages to join adjacent polymers creating a gel.  Pectic polysaccharides can also be cross-linked by dihydrocinnamic or diferulic acids.  The HGA's (galacturonans) are  initially secreted from the golgi as methylated polymers; the methyl groups are removed by pectin methylesterase to initiate calcium binding.

    Other pectic acids include Rhamnogalacturonan II (RGII) which features rhamnose and galacturonic acid in combination with a large diversity of other sugars in varying linkages.  Dimers of RGII can be cross-linked by boron atoms linked to apiose sugars in a side chain.

    Although most pectic polysaccharides are acidic, others are composed of neutral sugars including arabinans and galactans.  The pectic polysaccharides serve a variety of functions including determining wall porosity, providing a charged wall surface for cell-cell adhesion - or in other words gluing cells together (i.e,. middle lamella), cell-cell recognition, pathogen recognition and others.
D. Protein
    Wall proteins are typically glycoproteins (polypeptide backbone with carbohydrate side chains). The proteins are particularly rich in the amino acids hydroxyproline (hydroxyproline-rich glycoprotein, HPRG), proline (proline-rich protein, PRP), and glycine (glycine-rich protein, GRP).  These proteins form rods (HRGP, PRP) or beta-pleated sheets (GRP).  Extensin is a well-studied HRGP.   HRGP is induced by wounding and pathogen attack. The wall proteins also have a structural role since: (1) the amino acids are characteristic of other structural proteins such as collagen; and (2) to extract the protein from the wall requires destructive conditions. Protein appears to be cross-linked to pectic substances and may have sites for lignification. The proteins may serve as the scaffolding used to construct the other wall components.

    Another group of wall proteins are heavily glycosylated with arabinose and galactose. These arabinogalactan proteins, or AGP's, seem to be tissue specific and may function in cell signaling. They may be important in embryogenesis and growth and guidance of the pollen tube.
E. Lignin
    Polymer of phenolics, especially phenylpropanoids. Lignin is primarily a strengthening agent in the wall. It also resists fungal/pathogen attack.
F. Suberin, wax, cutin
    A variety of lipids are associated with the wall for strength and waterproofing.
G. Water
    The wall is largely hydrated and comprised of between 75-80% water. This is responsible for some of the wall properties. For example, hydrated walls have greater flexibility and extensibility than non-hydrated walls.
III. Morphology of the Cell Wall - there are three major regions of the wall:

1. Middle lamella - outermost layer, glue that binds adjacent cells, composed primarily of pectic polysaccharides.
   
2. Primary wall - wall deposited by cells before and during active growth. The primary wall of cultured sycamore cells is comprised of pectic polysaccharides (ca. 30%), cross-linking glycans (hemicellulose; ca 25%), cellulose (15-30%) and protein (ca. 20%) (see Darvill et al, 1980). The actual content of the wall components varies with species and age.  All plant cells have a middle lamella and primary wall.
   
3. Secondary Wall - some cells deposit additional layers inside the primary wall. This occurs after growth stops or when the cells begins to differentiate (specialize). The secondary wall is mainly for support and is comprised primarily of cellulose and lignin. Often can distinguish distinct layers, S1, S2 and S3 - which differ in the orientation, or direction, of the cellulose microfibrils.

IV. Tire analogy for the cell wall
    The wall is similar to a tire that has a series of steel belts or cords embedded in an amorphous matrix of rubber. In the plant cell wall, the "cords" are analogous to the cellulose microfibrils and they provide the structural strength of the wall. The matrix of the wall is analogous to the rubber in the tire and is comprised of non-cellulosic wall components. How are the various wall polymers arranged?  It appears that:

1. cross-linking glycans (hemicellulosic polysaccharides) are hydrogen bonded to the cellulose microfibrils
2. cross-linking glycans may also be entrapped inside cellulose microfibrils as they form
3. the different types of pectic polysaccharides are covalently bonded to one another
4. calcium bridges link pectic acids
5. connections between the protein and other wall polymers are still not clear
6. pectic polysaccharides and cross-linking glycans interact 
7. cross-linking glycans are linked by ferulic acid bridges or boron

V. Wall Formation
    The cell wall is made during cell division when the cell plate is formed between daughter cell nuclei. The cell plate forms from a series of vesicles produced by the golgi apparatus. The vesicles migrate along the cytoskeleton and move to the cell equator. The vesicles coalesce and dump their contents. The membranes of the vesicle become the new cell membrane. The golgi synthesizes the non-cellulosic polysaccharides. At first, the golgi vesicles contain mostly pectic polysaccharides that are used to build the middle lamella. As the wall is deposited, other non-cellulosic polysaccharides are made in the golgi and transported to the growing wall.
    Cellulose is made at the cell surface. The process is catalyzed by the enzyme cellulose synthase that occurs in a rosette complex in the membrane. Cellulose synthase, which is initially made in by the ribosomes (rough ER) and move from the ER → vesicles →  golgi →  vesicle →  cell membrane.  The enzyme apparently has two catalytic sites that transfer two glucoses at a time (i.e., cellobiose) from UDP-glucose to the growing cellulose chain.  Sucrose may supply the glucose that binds to the UDP.   Wall protein is presumably incorporated into the wall in a similar fashion.
    Remember that the wall is made from the outside in. Thus, as the wall gets thicker the lumen (space within the wall) gets smaller.
    Exactly how the wall components join together to form the wall once they are in place is not completely understood. Two methods seem likely: 

1. self assembly. This means that the wall components spontaneously aggregate; and
    
2. enzymatic assembly – various enzymatic reactions (XET) are designed for wall assembly. For example, one group of enzymes "stitches" xylans together in the wall to form long chains.  Oxidases may catalyze additional cross-linking between wall components and pectin methyl esterase may play an important role (see below).

VI. Strong Wall/Cell Expansion Paradox (don’t you just love a good paradox?)
    How can the wall be strong (it must withstand pressures of 100 MPa!), yet still allow for expansion? Good question, eh? The answer requires that the wall:
A. Be capable of expansion
    In other words, only cells with primary walls are capable of growth since the formation of the secondary wall precludes further expansion of the cell. The sequence of microfibril orientation changes during development. Initially the microfibrils are laid down somewhat randomly (isotropically). Such a cell can expand in any direction. As the cell matures, most microfibrils are laid down laterally, like the hoops of a barrel, which restricts lateral growth but permits growth in length.  As the cell elongates the microfibrils take on an overlapping cross-hatched pattern, similar to fiberglass. This occurs because the cell expands like a slinky - the width of the cell doesn't change by the microfibrils become aligned in the direction of growth just like the spring. This overlapping of microfibrils, which is strong and lightweight, prohibits further expansion.  
    But, what determines the orientation of the microfibrils? They are correlated with the direction of the microtubules in the cell. Evidence: treating a cell with colchicine or oryzalin (which inhibit microtubule formation) destroys the orientation of the microfibrils. The microtubules apparently direct the cellulose synthesizing enzymes to the plasma membrane.
    In addition to cellulose microfibril orientation, mature walls apparently loose their ability to expand because the wall components become resistant to loosening-activities. This would occur if there were increased cross-linking between wall components during maturation. This would result from:

1. producing wall polysaccharides in a form that makes tighter complexes with cellulose or other materials
2. increasing the lignin in the wall would increase cross-links between polymers
3. de-esterifying the pectic acids would increase calcium bridges;

B. Loosening (or Stress relaxation) the wall at the appropriate time
    Even though the microfibrils may be in the proper position to permit loosening, the wall is still rather strong. Recall that our wall model proposed strong (covalent) and weak (hydrogen bonds) links between the wall components. When the wall is loosened, weak bonds are temporarily broken to allow the wall components to slide or creep past one another. So, how is the wall temporarily loosened?

1. Protons are the primary wall loosening factor (Acid Growth Hypothesis). This idea was first proposed by David Rayle and R. Cleland in 1970. Some evidence:

• acid buffers stimulate elongation and rapid responses 5-15 min even in non-living tissues (Evans,1974);
• acid secretion is associated with sites of cell elongation (see Evans & Mulkey, 1981)
• Fusicoccin, a diterpene glycoside extracted from a fungus, stimulates proton secretion (activates a H⁺/K⁺ pump) and stimulates elongation.

2. Mechanism of proton action: Protons stimulate wall loosening by:

• disrupting acid-labile bonds such as H-bonds and calcium bridges; and
  
• enhancing the activity of enzymes that break wall cross-links including H-bonds and calcium bridges. Evidence for the enzyme involvement includes: (1) when primary walls are heated or treated with protein denaturing agents they can't be "loosened" by acid; and (2) adding proteins extracted from growing walls to heat-treated walls restores the acid response.
  
      Expansins – appear to be the primary wall-loosening enzymes. This class of proteins are activated by low pH and break the hydrogen bonds between cellulose and the cross-linking glycans. Other candidates for enzymes involved include: (1) pectin methyl esterase which would break the calcium bridges between pectins by esterifying the carboxyl groups; and (2) hydrolases – which would hydrolyze the cross-linking glycans (hemicelluloses).  For example, xyloglucan endotransglycosylase (XET) has been shown to cleave cross-linking glycans that could allow slippage of the wall components

 3. The acid effect is induced by indole-3-acetic acid (IAA, auxin), one of the major plant hormones. IAA stimulates proton excretion and cell growth/elongation.  Evidence:

• peeled coleoptiles + IAA → medium acidic; peeled coleoptiles + water → not acidic; and
• flooding auxin-treated tissue with neutral buffers prevents the growth response.

4. Mechanism of Auxin Action – How does auxin stimulate proton excretion and wall elongation?  There are two ideas:

Hypothesis 1:  Auxin activates pre-existing H⁺-ATPase pump proteins in the cell membrane. These proteins transport protons from the protoplast into the wall. Auxin probably first binds to a receptor molecule and this complex then actives the pump. This process is active - thus the pump requires ATP. Evidence: ATP stimulated acidification is observed soon after auxin treatment. Hypothesis 2: Auxin stimulates transcription and translation. Transcription/translation (protein synthesis) would be required to produce proton pump proteins (a wonderful alliteration), respiratory enzymes to provide ATP to power the process; and even enzymes for the synthesis of wall components and cell solutes (see C. & D. below). Evidence for the involvement of transcription/translation:

• Nooden (1968) found that artichoke disks increased in size when incubated with IAA but that the addition of antimycin (a protein synthesis inhibitor) prevented this response;
• soybean hypocotyls incubated with 2,4-D (an analog of IAA) produce at least 3 new polypeptides within three hours (Zurfluh & Guilfoyle, 1980);
• in vitro translation of mRNA occurs within 15 minutes of IAA treatment
• The proton effect is short-lived. Cell elongation stops 30-60 minutes after acidification. Continuous elongation requires longer term metabolic changes such as protein synthesis.

C. Wall synthesis occurs
    As the cell grows, wall synthesis needs to occur. Think about the color of a balloon as it is blown up - it gets lighter in color as the balloon gets larger because the thickness of the balloon decreases as it expands and stretches. Using this logic, we expect that plant cells should become thinner as they expand. Right? Wrong - cell walls remain a relatively uniform thickness throughout cell growth. Thus, we can conclude that new wall material must be made during cell elongation.
D. Enhanced solute synthesis
    The solute concentration of the cell remains constant during cell enlargement. This suggests that solutes are being synthesized since the volume of the cell is increasing. Maintaining a high solute concentration is necessary to allow for water uptake.
E.  Lock wall in place after expansion is complete
    Once wall elongation is completed, the cell needs to "lock it" in place.  This likely happens as the temporary bonds that were broken reform, and due to increased interactions (including enzymatic) between wall molecules.
F.  Water Uptake/Pressure

Web Sites:

• Carpita, N, McCann (2000) Cell Walls. Chapter 2. In Biochemistry & Molecular Biology of Plants.  Buchanan, BB, Gruissem W, Jones, RL. eds. American Society of Plant Biology, Beltsville, MD.  This is a great article!

References

• Albersheim, P. 1975. The Walls of Growing Plant Cells. Sci. Amer. 232: 81-95.
• Albersheim, P. 1985. Oligosaccharins. Sci. Amer. 253: 58.
• Brett, C. T. and J.R.Hillman. 1985. Biochemistry of Plant Cell Walls. Cambridge University Press, NY.
• Brett, C. & K. Waldron. 1996. Physiology and Biochemistry of Plant Cell Walls. 2nd edn. Chapman & Hall, NY.
• Cosgrove, D. 1986. Cell growth. ARPP 37: 377
• Delmer, D. 1987. Cellulose biosynthesis. ARPP 38: 259
• Fry, S.C. 1989. Dissecting the complexity of the plant cell wall. Plants Today 2: 126-132.
• Mulkey, T.J., K.M. Kuzmanoff and M.L. Evans. 1981. The agar-dye method for visualizing acid efflux paterns during tropistic curvatures. What's New in Plant Physiol. 12:9-12.
• Preston, R.D. 1979. Polysaccharide conformation and cell wall formation. ARPP 30:50.
• Taiz, L. 1984. Plant cell expansion. ARPP 35: 585-647
• Ruszkowski, Martha - check this link for the Ukrainian translation by M Ruszkowski.

From: http://employees.csbsju.edu/ssaupe/biol327/lecture/cell-wall.htm

Prokaryotic and eukaryotic cell architecture

During the 1950s, scientists developed the concept that all organisms may be classified as prokaryotes or eukaryotes. The cells of all prokaryotes and eukaryotes possess two basic features: a plasma membrane and cytoplasm. However, the cells of prokaryotes are simpler than those of eukaryotes. For example, prokaryotic cells lack a nucleus, while eukaryotic cells have a nucleus. Prokaryotic cells lack internal cellular bodies (organelles), while eukaryotic cells possess them. Examples of prokaryotes are bacteria and cyanobacteria (formerly known as blue‐green algae). Examples of eukaryotes are protozoa, fungi, plants, and animals.

Plasma membrane

All prokaryote and eukaryote cells have plasma membranes. The plasma membrane (also known as the cell membrane) is the outermost cell surface, which separates the cell from the external environment. The plasma membrane is composed primarily of proteins and lipids, especially phospholipids. The lipids occur in two layers (a bilayer). Proteins embedded in the bilayer appear to float within the lipid, so the membrane is constantly in flux. The membrane is therefore referred to as a fluid mosaic structure. Within the fluid mosaic structure, proteins carry out most of the membrane's functions.

Cytoplasm and organelles

All prokaryote and eukaryote cells also have cytoplasm (or cytosol), a semiliquid substance that composes the foundation of a cell. Essentially, cytoplasm is the gel-like material enclosed by the plasma membrane.

Within the cytoplasm of eukaryote cells are a number of membrane-bound bodies called organelles (“little organs”) that provide a specialized function within the cell.

One example of an organelle is the endoplasmic reticulum (ER). The endoplasmic reticulum is a series of membranes extending throughout the cytoplasm of eukaryotic cells. In some places, the ER is studded with submicroscopic bodies called ribosomes. This type of ER is referred to as rough ER. In other places, there are no ribosomes. This type of ER is called smooth ER. The ER is the site of protein synthesis in a cell. Within the ribosomes, amino acids are actually bound together to form proteins. Cisternae are spaces within the folds of the ER membranes.

Another organelle is the Golgi body (also called the Golgi apparatus). The Golgi body is a series of flattened sacs, usually curled at the edges. In the Golgi body, the cell's proteins and lipids are processed and packaged before being sent to their final destination. To accomplish this function, the outermost sac of the Golgi body often bulges and breaks away to form droplike vesicles known as secretory vesicles.

1 ) is derived from the Golgi body. It is a droplike sac of enzymes in the cytoplasm. These enzymes are used for digestion within the cell. They break down particles of food taken into the cell and make the products available for use. Enzymes are also contained in a cytoplasmic body called the peroxisome. 

Figure 1

The components of an idealized eukaryotic cell. A cell such as this probably does not exist, but the diagram shows the relative sizes and locations of the cell parts.

The organelle that releases quantities of energy to form ATP (adenosine triphosphate) is the mitochondrion (the plural form is mitochondria). Because mitochondria are involved in energy release and storage, they are called the “powerhouses of the cells.”

Green plant cells contain organelles known as chloroplasts, which function in the process of photosynthesis. Within chloroplasts, energy from the sun is absorbed and transformed into the energy of carbohydrate molecules. Plant cells specialized for photosynthesis contain large numbers of chloroplasts, which are green because the chlorophyll pigments within the chloroplasts are green. Leaves of a plant contain numerous chloroplasts. Plant cells not specializing in photosynthesis (for example, root cells) have few chloroplasts and are not green.

An organelle found in mature plant cells is a large, fluid-filled central vacuole. The vacuole may occupy more than 75 percent of the plant cell. In the vacuole, the plant stores nutrients as well as toxic wastes. Pressure within the growing vacuole may cause the cell to swell.

An organelle called the cytoskeleton is an interconnected system of fibers, threads, and interwoven molecules that give structure to the cell. The main components of the cytoskeleton are microtubules, microfilaments, and intermediate filaments. All are assembled from subunits of protein.

The centriole organelle is a cylinder-like structure that occurs in pairs. Centrioles function in cell division.

Many cells have specialized cytoskeletal structures called flagella and cilia. Flagellaare long, hairlike organelles that extend from the cell, permitting it to move. In prokaryotic cells, such as bacteria, the flagella rotate like the propeller of a motorboat. In eukaryotic cells, such as certain protozoa and sperm cells, the flagella whip about and propel the cell. Cilia are shorter and more numerous than flagella. In moving cells, the cilia wave in unison and move the cell forward. Paramecium is a well-known ciliated protozoan. Cilia are also found on the surface of several types of cells, such as those that line the human respiratory tract.

Nucleus

Prokaryotic cells lack a nucleus; the word prokaryotic means “primitive nucleus.” Eukaryotic cells, on the other hand, have a distinct nucleus.

The nucleus of eukaryotic cells is composed primarily of protein anddeoxyribonucleic acid, or DNA. The DNA is organized into linear units calledchromosomes, also known as chromatin when the linear units are not obvious. Functional segments of the chromosomes are referred to as genes. Approximately 100,000 genes are located in the nucleus of all human cells. Nuclear proteins belong to a class of proteins called histones. The chromosome is coiled around the histones.

The nuclear envelope, an outer membrane, surrounds the nucleus of a eukaryotic cell. The nuclear envelope is a double membrane, consisting of two lipid layers (similar to the plasma membrane). Pores in the nuclear envelope allow the internal nuclear environment to communicate with the cell cytoplasm.

Within the nucleus are two or more dense organelles referred to as nucleoli (the singular form is nucleolus). In nucleoli, submicroscopic particles known asribosomes are assembled before their passage out of the nucleus into the cytoplasm.

Although prokaryotic cells have no nucleus, they do have DNA. The DNA exists freely in the cytoplasm as a closed loop. It has no protein to support it and no membrane covering it. A bacterium typically has a single looped chromosome with about 4,000 genes.

Cell wall

Many kinds of prokaryotes and eukaryotes contain a structure outside the cell membrane called the cell wall. With only a few exceptions, all bacteria have thick, rigid cell walls that give them their shape. Among the eukaryotes, fungi and plants have cell walls. Cell walls are not identical in these organisms, however. In fungi, the cell wall contains a polysaccharide called chitin. Plant cells, in contrast, have no chitin; their cell walls are composed exclusively of the polysaccharide cellulose.

Cell walls provide support and help cells resist mechanical pressures, but they are not solid, so materials are able to pass through rather easily. Cell walls are not selective devices, as plasma membranes are.

Function of MgCl2 in PCR

During replication dntp's gets broken down to dnmp's to form phosphodiester bond between 3' OH of adjacent nucleotide and 5' Phosphate of the upcoming nucleotide. in this reaction Mg++ binds to the alpha phosphate group of dntp and helps in the removal of beta and gamma Phosphate from dntp.

Another school of thought has it that increasing Mg++ increases Taq (DNA-dependent DNA polymerase) activity but at the expense of specificity, whereas less Mg++ lowers Taq's activity but increases its specificity. So the trick is knowing how much (or how little) to use. Generally, 3 mM Mg++ works well for most Taq-based mastermixes (used for PCR/qPCR) although as much as 5.5 mM Mg++ has been used with success in other mastermix formulations. Perhaps just use pi mM (3.14159265359 mM Mg++) and hope for the best~! You want good activity (productivity) but also good specificity...

From Research Gate.. 

List of Softwares for Genomics

Informatics
Name	Website	Notes
Abiomics	http://www.abiomics.eu	Bioinformatics
Accelerys	http://accelrys.com	Bioinformatics
Active Motif	http://www.activemotif.com	Acquired TimeLogic
AgileBio	http://www.agilebio.com	Custom bioinformatics
Annai Systems	http://www.annaisystems.com	Genome data management (Developed UCSC Cancer Genomics Hub)
Appistry	http://www.appistry.com	Data management, analysis, interpretation & storage (NGS/Other)
Ariadne Genomics	http://www.ariadnegenomics.com	Expression, pathways, text mining
aScidea	http://www.ascidea.com	Bioinformatics solutions including NGS products
Aspera	http://www.asperasoft.com	Fast efficient method for high-speed data transfer
AssurX	http://www.assurx.com	Risk and compliance management software for clinical NGS
Atragene	http://ri.atragene.com	Custom software solutions (NGS interpretation?)
Avadis	http://www.avadis-ngs.com	NGS through pathway analysis
Baseclear	http://www.baseclear.com	NGS, comprehensive
Bina Technologies	http://www.binatechnologies.com	NGS interpretation, comparison
Biobase	http://www.biobase-international.com	Identify functionally significant genome variants
Biocomputing Platforms Ltd	http://www.bcplatforms.com	NGS and variation data analysis and management
BioCOS Life Sciences	http://www.biocosls.in	NGS and microarray interpretation
BioDiscovery	http://www.biodiscovery.com	Copy number, expression, NGS
Bioinformatics Solutions	http://www.bioinformaticssolutions.com	Mapping
Biomatters	http://www.biomatters.com	Producers of Geneious
Biomol-Informatics	http://www.biomol-ngs.com	including NGS analysis
BioNeos	http://bioneos.com	Custom bioinformatics
Bionivid	http://www.bionivid.com	Bioinformatics services and development
Biosof	http://www.bio-sof.com	Sequence Analysis, Structure and Function Prediction
Bioteam	http://www.bioteam.net	MiniLIMS for NGS platforms
BMR Genomics	http://www.bmr-genomics.com	Assembly through interpretation
Cartagenia	http://www.cartagenia.com	web-based software and database, for diagnostic genome interpretation
cBio	http://www.cbiocorp.com	bioinformatics solutions (NGS and microbial diagnostics)
ChunLab	www.chunlab.com	NGS data analysis solutions
CLC Bio	http://www.clcbio.com	Comprehensive, Including NGS
Conexio Genomics	http://www.conexio-genomics.com	Next gen software for HLA and other applications
CSBI	Link dead as of June 2013	Software and training in bioinformatics
Cypher Genomics	http://cyphergenomics.com	Interpretation of NGS data, clinical
Diagnomics	http://www.diagnomics.com	Software and service for NGS and personalized healthcare
DNA 2.0	https://www.dna20.com	Tools for gene design through sequence analysis
DNA Guide	http://dnaguide.blogspot.com	Bring NGS to clinic
DNAlytics	http://www.dnalytics.com	Data Mining & IT solutions for personalized medicine
DNAnexus	http://dnanexus.com	Cloud-based DNA data management, analysis, and collaboration platform
DNASoftware	http://www.dnasoftware.com	Multiple DNA analysis packages
DNAstar	http://www.dnastar.com	NGS, assembly, other
Douglas Scientific	http://www.douglasscientific.com	Microarray, qPCR etc
Eagle Genomics	http://www.eaglegenomics.com	Pipelines, consulting, scalable genomics analysis
ecSeq Bioinformatics	http://www.ecseq.com	NGS interpretation, custom pipelines
Enlis Genomics	http://www.enlis.com	Powerful and simple NGS interpretation software
Era7 Bioinformatics	http://www.era7bioinformatics.com	Bacterial genomics
Euformatics	http://www.euformatics.com	Software platforms for NGS (research and clinical)
Existance Genetics	Bankrupt.  Website offline	Tertiary analysis (clinical interpretation) and reporting of NGS data
Fluidigm	http://www.fluidigm.com	Expression, targeted resequencing, SNP
Gataca	http://www.gatacallc.com	database construction, genomic analysis, bioinformatics algorithm construction
GE Healthcare	http://ge.geglobalresearch.com	Bring NGS to clinic
Genalice BV	www.genalice.com	very accurate and fast (up to 100 x faster than BWA) short read aligner
Genbios	http://www.genbios.in/	databases of genes, pathways etc.
GenDx	http://www.gendx.com/index.php/company	HLA typing
Gene Codes	http://www.genecodes.com	Bioinformatics software for DNA sequence analysis
GeneExplain	http://www.genexplain.com	Expression, proteomics
GeneGo (Thomson Reuters)	http://www.genego.com	pathway analysis
Geneious	http://www.Geneious.com	NGS assembly through annotation
GeneSpring (Agilent)	http://www.genomics.agilent.com	Gene expression, NGS and proteomics analysis, pathways
Genevia Technologies	http://www.genevia.fi	systems biological data analysis
GenoLogics	http://genologics.com	LIMS systems for -omics labs
Genomatix	http://www.genomatix.de	NGS analysis software
Genome Enterprise Limited	http://www.genome-enterprise.com	NGS analysis associated with TGAC
GenomeQuest	http://www.genomequest.com	NGS: research and clinical applications
Genomic Expression	http://www.genomicexpression.com	Proprietary cost-effective genomics diagnostics
Genomic Solutions	http://bioinformatics.genomicsolutions.com	proteomics, DNA sample prep
GenoSpace	http://www.genospace.com	Cloud based 'omics secure data sharing targeting clinical applications
Genostar	http://www.genostar.com	Solutions for microbial genome, proteome and metabolome comparison
Geospiza	http://www.geospiza.com	NGS/Microarray analysis and LIMS software
GNS healthcare	http://www.gnshealthcare.com	Big Data management devoted to healthcare solutions
Golden Helix	http://www.goldenhelix.com	Multiple genomics packages
Heracle BioSoft S.R.L.	http://www.dnabaser.com	Heracle makes DNABaser Assembly software
Igenbio.Inc	www.igenbio.com	Advanced bioinformatics services
Ingenuity Systems	http://www.ingenuity.com	Genome, pathway analysis
InSilicoDB	http://insilico.ulb.ac.be	Genomics databases and in-silico genomics tools
Integromics	http://www.integromics.com	Data handling and analysis for NGS, microarray, qPCR
Interactive Biosoftware	http://www.interactive-biosoftware.com	Visualization, interpretation and annotation of NGS data
Intrepid Bioinformatics	http://www.intrepidbio.com	NGS, SNP, Databases
IntuitiveGenomics	http://www.intuitivegenomics.com	NGS assemply, annotation, interpretation
Invitae	http://www.invitae.com	Working with Genomic Health and Locus Dev. (variant database)
JMP Genetics	http://www.jmp.com	Solutions for research, clinical, and agricultural genomics
JSI Medical Systems	http://www.jsi-medisys.com	NGS analysis software
Knome	http://knome.com	NGS interpretation, clinical
Korilog	http://www.korilog.com	Products based on Klast high perfomance Seq. Comparison tool
Kovid Bioanalytics	www.kovidbioanalytics.com	Statisitical Analysis, Genomics Data Mining, Clinical Informatics
Lab 7 systems	www.lab7.io	Integrates multiple NGS software packages coherently
Leucine Rich Bio	http://www.leucinerichbio.com/	heuristic and statistical analysis of omics data and basic bioinf
Locus Development	http://locusdevelopmentinc.com	Bring NGS to clinic
Maverix Biomics	www.maverixbio.com	Cloud-based NGS data management, interpretation, and visualization
Microsoft	http://www.microsoft.com	Genomics
MolecularHealth	http://MolecularHealth.com	Molecular safety, molecular oncology
MultiD Analyses	www.multid.se	qPCR analysis software
NextBio	http://www.nextbio.com	Genomics, expression, literature, clinical
NuMedii	http://numedii.com	Genome-wide molecular profiles
Ocimum Biosolutions	http://www3.ocimumbio.com	Research Accelerator (for 'Omics comparison), extensive databases
Omicia	http://www.omicia.com	VAAST genome interpretation
Omixon	http://www.omixon.com	Targeted sequencing, HLA, variant analysis
OncoDNA	http://www.oncodna.com	Software packages for deep sequencing and cancer biomarker monitoring
Partek	http://www.partek.com	Software - NGS, expression etc.
Personalis	http://www.personalis.com	Interpretation of NGS data, clinical
Phenosystems	http://www.phenosystems.com	Software NGS and others
Premier Biosoft	http://www.premierbiosoft.com	Bioinformatics
Real Time Genomics	http://www.realtimegenomics.com	Sequence alignment through variant database comparison
Rimarcable	http://www.rimarcable.nl	Custom Software solutions (including healthcare applications)
Salsa Computing	http://www.salsacomputing.com	Big data, including bioinformatics
Samsunggenome	http://www.samsunggenome.com	NGS sequencing through interpretation
Sapio Sciences	http://www.sapiosciences.com	LIMS systems for genomics labs
Scale Genomics	http://www.scalegenomics.com	Cloud storage and analysis of NGS data (Domain seems to be parked)
ScaleMP	http://www.scalemp.com	Big data company, assembly through interpretation
Seven Bridges Genomics	https://igor.sbgenomics.com	NGS variant interpretation open source pipelines
SGI	Http://www.sgi.com	Big data company, assembly through interpretation
Silicon Valley Biosystems	http://www.svbio.com	NGS interpretation software
SimulConsult	http://www.SimulConsult.com	Genomic variant interpretation, phenomics
Simulconsult	http://www.simulconsult.com	Sofware packages to help clinicians interpret genetic variant tables
Soft Genetics Inc	http://www.softgenetics.com	NGS interpretation software packages
SolveBio	http://www.solvebio.com	Genome analysis
Spiral Genetics	http://www.spiralgenetics.com	Annotation
StationX	http://www.stationxinc.com	Genome interpretation
Strand Life Sciences	http://www.strandls.com	Genespring, Avadis-NGS
Syapse	http://www.syapse.com	Making omics data actionable in clinical research and development
TechTrendsIT	http://www.techtrendsit.com	Software builders.  Can assist with cloud computing and big data etc.
Translational Software	http://www.translationalsoftware.com	Bring NGS to clinic
Woodward Informatics Ltd	http://www.strychnine.co.uk	Custom bio- and chemo-informatics
xGen Cybernetics	www.xgencloud.com	Bioinformatics, clinical genetics and predictive medicine
Zymo Research	www.zymoresearch.com/services	Bioinformatics for all NGS applications/epigenetic analysis
Other resources for locating and evaluating NGS tools and services
Resource	Website	Notes
Allseq (formerly Blueseq)	http://www.allseq.com	Match sequencing capacity with sequencing need (advisors)
Omictools	http://omictools.com	Excellent set of lists of structural and functional genomics tools
Omicsmaps	http://omicsmaps.com	Interactive world map of NGS Instruments
Seqanswers	http://seqanswers.com	One of the oldest and most popular NGS forums
Genomics Service Providers
Company	Website	Notes
23 and me	http://www.23andme.com	Consumer genomics
5am Solutions	http://www.5amsolutions.com	Custom bioinformatics
Active Motif	http://www.activemotif.com	ChIP-Seq, (h)MeDIP-Seq and Bisulfite Sequencing among others
Adaptive Biotechnologies	http://www.adaptivebiotech.com	NGS on immune cells, moving to clinical
Allere Laboratory	http://www.AllereLabs.com	NGS - Based health and wellness
AllGenetics	http://www.allgenetics.eu	Molecular genetics support for academic and corporate labs
Almac group	http://www.almacgroup.com	CLIA lab, microarray including CNV and SNP
Ambry Genetics	http://www.ambrygen.com	Clinical Genetics and Genomics Services
Ancestry.com	http://www.ancestry.com	Integrating geneologies with DNA analysis
Applied Genomics Institute	http://www.appliedgenomics.org	NGS and related BioInformatics
Applied Microarrays	http://www.appliedmicroarrays.com	Custom microarray
Ariana Pharma	http://www.arianapharma.com	Multiple research/clinical genomics applications
AROS Applied Biotechnology	http://www.arosab.com	Extraction, PCR, microarray, and NGS services.
Asuragen	http://www.asuragen.com	Clinical NGS, ion torrent, FFPE capabilities
Atlas BioLabs	http://www.atlas-biolabs.de	NGS, Arrays, CGH, Bioinformatics
BairesDev	http://www.bairesdev.com/	BairesDev can develop software and web-apps for your algorithm
Baseclear	http://www.baseclear.com	NGS, comprehensive
Beckman Coulter Genomics	http://www.beckmangenomics.com	NGS, Sanger sequencing, data analysis, other
Beijing Genomics (BGI)	http://www.genomics.cn	NGS through assembly, interpretation
Bina Technologies	http://www.binatechnologies.com	NGS interpretation, comparison
Bio S & T	http://www.biost.com	Sanger sequencing and expression, WGS?
BioChain	http://www.biochain.com	NGS through genome interpretation
BioCOS Life Sciences	http://www.biocosls.in	NGS and microarray interpretation
BioDigital valley	http://www.biodigitalvalley.com	Proteomics, Genmics, Bioinformatics
Biodiscovery	http://www.biodiscovery.com	Genomics analysis consulting
Bioo	http://www.biooscientific.com	Comprehensive including NGS
Bioserve	http://www.bioserve.com	human DNA, microarray
BMR Genomics	http://www.bmr-genomics.com	NGS through interpretation
Caliper (PE)	http://www.caliperls.com	Lab auto and NGS services
Cartagenia	http://www.cartagenia.com	Enterprise integration of clinical NGS
CCCB - Harvard	http://cccb.dfci.harvard.edu/cccb/analytical	Computational biology services for industry and academe @ Harvard
Centrillion Biosciences	https://www.centrillionbio.com	experimental design, data production and bioinformatic services
ChunLab	www.chunlab.com	Next gen sequencing and analysis services
Cofactor Genomics	http://www.cofactorgenomics.com	NGS, bioinformatics
Complete Genomics	http://www.completegenomics.com	NGS through assembly (Subsidiary of BGI)
Computomics	http://www.computomics.com	NGS interpretation for agricultural biotech
Contig Express	http://www.contigexpress.com	Integrated genomics data service provider
CorrZ	http://www.corrz.com	Biostatistics, analysis for NGS and microarray
Counsyl	www.counsyl.com	Carrier Screening via NGS or Targeted Genotyping
Courtagen	http://www.courtagen.com	NGS through interpretation, proteomics
Covance Genomics	http://www.covance.com	Full gervice genomics partner
CPGR	http://www.cpgr.org.za	Non profit genomics/bioinformatics center
Creative Bioinformatics	http://www.creativebioinformatics.com/	Consultancy that will tailor a bioinformatics team to your project.
Cypher Genomics	http://cyphergenomics.com	Interpretation of NGS data, clinical
Cytogenomics	http://www.cytognomix.com	High throughput interpretation of genomic variation
Data2Bio	http://www.data2bio.com	Design, Execute, analyse NGS projects
deCode Genetics	http://www.decode.com	Clinical genomics
Decode Nomics	Expired Website	Bioinformatics analysis services
DNA Link	www.dnalink.com	NGS, microarray, personal genomics, forensics etc.
DNAlytics	http://www.dnalytics.com	Data Mining & IT solutions for personalized medicine
DNASTAR	http://www.dnastar.com	Genome Assembly Services (and sequencing, downstream analysis etc.)
ecSeq Bioinformatics	http://www.ecseq.com	Sequencing- assembly and analysis
Edge Bio	http://www.edgebio.com	CLIA exome sequencing
Era7 Bioinformatics	http://www.era7bioinformatics.com	Bacterial genomics
Euformatics	http://www.euformatics.com	Consultation on NGS projects
Eureka Genomics	http://www.eurekagenomics.com	NGS through assembly, other
Eurofins, MWG, Operon	http://www.operon.com	Oligos through NGS analysis
Evogene	http://www.evogene.com	Identify agrinomically imortant plant genes
Expression Analysis	http://www.expressionanalysis.com	WGS, RNAseq, microarray
Eye.Q.Consulting	Website Pending	Biological interpretation of array data
Fasteris	http://www.fasteris.com	Including Illumina NGS
Fios Genomics	http://www.fiosgenomics.com	Exome, methylation for epigenetics, other
FitGenes	http://www.fitgenes.com	DNAprofiling, Nutrigenomics
Floragenex	http://www.floragenex.com	Plant and animal NGS
Foundation Medicine	http://www.foundationmedicine.com	Cancer diagnostics (including NGS)
Ganit labs	http://www.ganitlabs.in	Expression, NGS, genotyping
GATC-Biotech	http://www.gatc-biotech.com	NGS
GenAlysis	http://www.genalysislaboratory.com/	Veterinary genetic testing
Genbios	http://www.genbios.in/	Custom database creation service
Gene TLC	http://www.genetlc.com	Personalized genomics
GenebyGene	http://www.genebygene.com	Services include Ancestry, Health, Research, and Paternity
GeneDx	http://www.genedx.com	Clinical - including whole exome
Genergy Bio-tec	http://www.genenergy.cn	We prowide GWAS, NGS service
GeneWiz	http://www.genewiz.com	DNA services including NGS
Genialis	http://www.genialis.com/	Develops interactive visual analytics web applications and analyses NGS data
Genialis	http://www.genialis.com	Bioinformatics services as well as stand alone analysis products
Genohub	https://genohub.com	Helps researchers identify the best services for their NGS project
Genomatix	www.genomatix.de	NGS analysis services
Genome British Columbia	http://www.genomebc.ca	Investment and management of genomics projects
Genome Enterprise Limited	http://www.tgac.ac.uk/	Next Gen Sequencing associated with TGAC
Genome International Corporation	http://www.genome.com	Bioinformatics support for the life sciences
Genome Life Sciences	http://www.genomels.com	Bioinformatics support for the life sciences
Genome Project Solutions	http://genomeprojectsolutions.com	NGS sequencing though interpretation
GenomeDX	http://www.genomedx.com	Clinical cancer genomics
Genomic Express	http://www.GenomicExpress.com	CLIA laboratory, consumer genomics, medical genetics
Genomic Expression	http://www.genomicexpression.com	NGS libraries
Genomic Health	http://www.genomichealth.com	Clinical genomics
Genomic Health	http://www.oncotypedx.com	Clinical genomics
Genomnia	http://www.genomnia.it	NGS sequencing through interpretation
Genoptix Medical Laboratory	http://www.genoptix.com	personalized diagnostics some seq analysis
Genoscreen	http://http://www.genoscreen.com	NGS, interpretation, consultation
GenoSplice technology	http://www.genosplice.com	"Wet" lab and Bioinformatics services
Genostar	http://www.genostar.com	Solutions for microbial genome, proteome and metabolome analyses
Genotypic	http://www.nextgenseq.com	NGS, Microarrays, custom microarrays, Bioinformatics, PCR, and more
Genotypic	http://www.genotypic.co.in	NGS, Microarrays, custom microarrays, Bioinformatics, PCR, and more
Golden Helix	http://www.goldenhelix.com	Genomic analysis/genomics project consultation
Good Start Genetics	http://www.goodstartgenetics.com	Carrier screening
Health In Code	http://www.healthincode.com	Genetic diagnosis of inherited cardiovascular diseases: NGS/Sanger
High Throughput Sequencing	http://www.htseq.org	Service associated with UW
Holgentech	http://www.holgentech.com	Fractal defect mining, Fractal analytics (cancer)
Hybridstat	www.hybridstat.gr	Software design/bioinformatics consulting
Ibioa	http://www.ibioa.com/	CRO, with NGS analysis as one of many services
IDT	http://www.idtdna.com	In addition to Oligos, they do 454 sequencing as well
Igenbio.Inc	www.igenbio.com	Advanced bioinformatics services
Illumina	http://www.illumina.com	NGS
IMBcom	http://www.imbcom.com.au	Commercialization of academic genomics services
IMGM Laboratories	www.imgm.com	Comprehensive NGS/Bioinformatics provider
Inqaba Biotechnical Industries	www.inqababiotec.co.za	African NGS and Bioinformatics services
Interleukin Genetics	http://www.ilgenetics.com	Consumer genetic tests
iRepertoire	http://irepertoire.com	NGS on immune cells
Keygene	http://www.keygene.com	Including whole genome sequencing of plants
Knome	http://knome.com	NGS interpretation, clinical
LabCorp	https://www.labcorp.com	Including clinical genomics
LC Sciences	http://www.lcsciences.com	Services include targeted sequencing and custom microarrays
Lexogen	http://www.lexogen.com	RNA-Seq R&D
LGC Genomics	http://www.lgc.co.uk	Nucleic acid extraction through NGS analysis
Lucigen	http://www.lucigen.com	Library prep through NGS analysis
Macrogen	http://macrogen.com	NGS, Sanger sequencing, microarray
Malaysian Genomics Resource Ctr	http://www.mgrc.com.my	Sequencing through analysis
Malaysian Genomics... (MGRC)	http://www.mgrc.com.my	NGS through Interpretation
Medomics	http://www.medomics.com	Clinical mitochondrial genomics
Metamark Genetics	http://www.metamarkgenetics.com	Clinical genomics, cancer focus
MOgene	http://mogene.com	NGS, microarray
MolecularMD	http://www.molecularmd.com	Comprensive CRO with significant NGS and bioinformatics capabilities
Moleculo	http://www.moleculo.com	DNA sequencing method:  very long and accurate reads
Multiplicom	http://www.multiplicom.com	Dx kits for targeted resequencing.
Mycroarray	http://mycroarray.com	Custom microarrays and capture bait libraries
Myriad Genetics	https://www.myriad.com	Clinical Genetics and Genomics Services
Natera	http://www.natera.com	Clinical prenatal genetic testing
Navigenics	http://www.navigenics.com	Consumer genomics
Nextcode	https://www.nextcode.com/	Delivering the resources developed at deCODE Genetics to the clinical domain.
Novocraft	http://www.novocraft.com	NGS Sequence Analysis and Custom pipelines
Nucleome Informatics	www.nucleomeinfo.com	Next gen sequencing and data analysis
Ocimum Biosolutions	http://www3.ocimumbio.com	Expression, LIMS, other
Oftalmic	http://www.oftalmic.com	Sequencing and analysis services (inherited eye diseases)
Omega Bioservices	http://www.omegabioservices.com	Exome sequencing, interpretation
Omics Maps	http://omicsmaps.com	Crowd Sourced list of sequencing centers
Omics Systems	http://www.omicssystems.com	Comprehensive list of genomics services
OncoDNA	www.oncodna.com	Targeted sequencing to exome with oncologic theranostic interpretation
Opgen	http://www.opgen.com	Genome mapping, assembly
Otogenetics	http://www.otogenetics.com	Whole exome NGS, RNAseq
Oxford Gene Technology	http://www.ogt.co.uk	Whole Exome, Targeted resequencing, microarray, ramily/trio analysis
Paradigm Diagnostics	http://www.paradigmdx.org	Clinical interpretation of NGS data
Paradigm	http://www.paradigmdx.org	Nonprofit organization (cancer pathway analysis, genomics, etc.)
Pathgroup	http://www.pathgroup.com	Pathology services including clinical NGS
Pathogenica	http://www.pathogenica.com	Clinical Sequencing/NGS
Pathtwork Genomics	http://www.pathworkdx.com	CLIA microarray (RNAseq?)
Pathway Genomics	http://www.pathway.com	Personal genomics/nutrition
Perkin Elmer	http://www.perkinelmer.com	NGS sequencing, exome
Personal Genome Diagnostics	http://www.personalgenome.com	Clinical cancer exome sequencing through interpretation
Personalis	http://www.personalis.com	Interpretation of NGS data, clinical
Population Genetics	http://www.populationgenetics.com	Targeted resequencing of large populations
Prevention Genomics	http://www.preventiongenetics.com	Clinical Genetic Screening
Progenus	http://www.progenus.be	Sequencing and analysis (Sanger through NGS)
Prognosys	http://www.prognosysbio.com/sequensys	NGS Sequencing and analysis services
QFAB	http://www.qfab.org	Bioinformatics service specialists
qGenomics	www.qgenomics.com	Genomics for human health
Qtlomics Technology	http://www.qtlomics.com/	Merging NGS/Bioinformatics with traditional plant breeding
Quality Partner	http://www.quality-partner.be	16S metagenomic analysis applied to food microbiology
Quest	http://www.questdiagnostics.com	Including clinical genomics
QUEST Diagnostics	http://www.QuestDiagnostics.com	Clinical Diagnostics including clinical genomics (see Genomic Vision)
Raindance	http://www.raindancetech.com	Raindance offers design services in addition to instrumentation
Response Genetics	http://www.responsegenetics.com	Genetic and genomic approaches to cancer diagnostics
Rheumakit	http://www.rheumakit.com	NGS-based Rheumatology diagnostics
Samsunggenome	http://www.samsunggenome.com	NGS sequencing through interpretation
SCR Consulting	www.scr-consulting.co.uk	Biotech IT service - Builds custom informatics for life scientists
Second Genome	http://www.secondgenome.com	NGS with microbiome focus
SEQLAB	http://www.seqlab.de	Including NGS
Sequenom	http://www.sequenom.com	Microarray structural/functional genomics
Sequenta	http://sequentainc.com	Clinical NGS and immunology
Seqwright	http://www.seqwright.com	NGS through assembly
ServiceXS	http://www.servicexs.com	NGS, genotyping, methylation and expression analysis services
Shanghai Biotech	http://www.shbiotech.org	NGS panels and Bioinformatics services
Skuldtech	www.skuldtech.com	Biomarker Discovery, Diagnostic NGS - very well established
SmartGene	http://www.smartgene.com	application-specific solutions for clinical sequencing
Sophia Genetics	http://www.sophiagenetics.com	Integrated Clinical NGS Dry Lab Service
Sorensen Genomics	http://www.sorensongenomics.com	Clinical and forensic genomics
Spiral Genetics	http://www.spiralgenetics.com	Annotation
STATSARRAY	http://statsarray.com	NGS data analysis services (also maintain omictools database)
Synexa Life Sciences	http://www.synexagroup.com	Clinical human genomics
TATAA Biocenter	www.tataa.com	gene expression profiling services
Taueret Laboratories	http://www.taueret.com	Genetic laboratory services to researchers of medical genetics
UD-GenoMed	http://www.ud-genomed.hu	Full service clinical genomics lab
Wuxi Apptec	http://www.wuxiapptec.com	Comprehensive, including NGS and expression analysis
xcelris labs	http://www.xcelrislabs.com	Sequencing and analysis (Sanger through NGS)
XDx	http://www.xdx.com	Clinical, expression-based transplant monitoring
Zymo Research	www.zymoresearch.com/services	Comprehensive NGS services for epigenetic analysis
Providers of cells and protein solutions
Company	Website	Notes
AllCells, LLC	http://www.allcells.com	Largest West Coast hematopoietic stem cell provider
Alstem	http://www.alstembio.com	Make and distribute monoclonal antibodies
Bioneer	http://www.bioneer.com	ExiprogenTM Automated protein synthesis/purification instrument
Novoprotein	http://www.novoprotein.com	custom antibody services and recombinant proteins
Nvigen	http://www.nvigen.com	Nanoparticles (Fluorescent/Magnetic etc.)
ProMab	http://www.promab.com	Largest monoclonal antibody supplier