Riboflavin biosynthesis in Bacillus Subtilis

Riboflavin biosynthesis in Bacillus Subtilis Motivation: Many important problems in cell biology require the dense nonlinear interactions between functional modules to be considered. The importance of computer simulation in understanding cellular processes is now widely accepted, and a variety of simulations algorithms useful for studying certain subsystems have been designed. 1 introduction Human and animal needs vitamins and minerals to help in the growth of the body and prevent disease, obtained from food or supplements. For example, an important vitamin is riboflavin and Riboflavin is water-soluble vitamins also sources of riboflavin are milk, meat, fish, fruits and vegetables. Diets that do not contain riboflavin or poor riboflavin cause serious health problems. For example, a disorder of the digestive system, heart disease and blood vessels. Riboflavin deficiency may also cause cancer. The name riboflavin comes from ribose (the sugar whose reduced form, ribitol, forms part of its structure) and flavin, the ring-moiety which imparts the yellow color to the oxidized molecule (from Latin flavus, yellow). The reduced form, which occurs in metabolism along with the oxidized form, is colorless. Riboflavin has been used in several clinical and therapeutic situations. For over 30 years, riboflavin supplements have been used as part of the phototherapy treatment of neonatal jaundice. The light used to irradiate the infants breaks down not only bilirubin, the toxin causing the jaundice, but also the naturally occurring riboflavin within the infants blood, so extra supplementation is necessary. Riboflavin is a yellow substance which was isolated in 1879. Riboflavin name comes from ribose sugar substance and also characterized by yeast an excellent dietary source. In addition, it contains two factors which are aide Flavin adenine dinucleotide (FAD) and a Flavin mononucleotide (FMN) (Daniel, 2012). 2 Background It has been pointed out by (Bretzel, et al.1999) and recently been produced commercially by riboflavin chemical manufacturing or by fermentation, where the production of riboflavin with a purity of at least 97-98% of foodstuffs and medicines. In addition, scientists biological prefer to biological and chemical manufacturing and much more than manufacturing because of the many advantages. For example, low material cost, low waste and energy requirements due to the use of renewable energy. It has been producing a large amount of riboflavin from many organisms’ example of the use of fungi, yeast and bacteria Bacillus Subtilis (Li Wu, Q, 2007). Research indicated that since 30 years the world’s consumption of riboflavin equivalent of 1.25 Ãâ€" 106 kg per year. However, the amount of riboflavin are currently more than 3.000 tons per year (Han Lim, et al.2001). Bacillus Subtilis is a gram positive, aerobic bacteria that converts glucose directly to riboflavin. Cell metabolisms are highly dependent on environmental conditions, so the metabolic state often shifts during the cultivation period. Characterizing the transience of metabolic fluxes is important for understanding how cells responded to environmental changes. (Boyle NR, Morgan JA, 2009) Fig. 1.Riboflavin synthesis reactions As the provide chart (Riboflavin synthesis reactions) shows, riboflavin bio-product of the yeast start from Guanosine triphosphate (GTP) and there are two molecules of ribulose-5-phosphate, which leads to the production of riboflavin with enzymes encoded with RIB1, RIB2, RIB4, RIB5, RIB7 and FMN1 names which helps in the production of Flavin mononucleotide also Flavin adenine dinucleotide by enzyme FAD. Moreover, ribose involved in the production of riboflavin with 3 parts of 4-dihydroxy-2-butanone-4-P and 6 parts of 7-dimethyl-8-(1-D-ribityl) lumazine. 3 methods 3.1 In-silico (computational) model In silico models have prompted considerable interest and debate because of their potential value in predicting the properties of chemical substances for regulatory purposes. The European REACH legislation promotes innovation and encourages the use of alternative methods, but in practice the use of in silico models is still very limited. (Benfenati E. and others, 2011) 3.2 Conceptual modelling – ER diagram Conceptual modeling is a way of carrying out an analysis of a system in order to identify the types of things in the system, the actual instances of those things and the relationships between those things. Entity relationship (ER) modeling is a conceptual modeling approach commonly used in Computer Science to model a domain for the development of a database. However, ER modeling provides a useful approach to starting to think about models of biological systems. 3.3 Biological network analysis and construction using Cytoscape Cytoscape platform providing a full feature set for biological network analysis. It is an open source software platform for visualizing molecular interaction networks and biological pathways and integrating these networks with annotations, gene expression profiles and other state data. Although Cytoscape was originally designed for biological research, now it is a general platform for complex network analysis and visualization. Cytoscape core distribution provides a basic set of features for data integration, analysis, and visualization. 3.4 Exploring integrated networks using String and Ondex The Ondex is a data integration platform which enables data from diverse biological data sets to be linked, integrated and visualised through graph analysis techniques. Ondex uses a rich and flexible core data structure, which has the ability to bring together information from structured databases and unstructured sources such as biological sequence data and free text. Ondex also allows users to visualize and analyze the integrated data. Ondex is free and open-source software. STRING is a database of known and predicted protein interactions. The interactions include direct (physical) and indirect (functional) associations. Together String and Ondex are very useful in case of exploring integrated networks and simulation. 3.5 Dynamic modeling using SBML SBML is neutral with respect to programming languages and software encoding; however, its oriented towards allowing models to be encoded using XML. By supporting SBML as a format for reading and writing models, different software tools (including programs for building and editing models, simulation programs, databases, and other systems) can directly communicate and store the same computable representation of those models. This removes an impediment to sharing results and permits other researchers to start with an unambiguous representation of the model, examine it carefully, propose precise corrections and extensions, and apply new techniques and approaches — in short, to do better science (Rhoads DG, Pring M, 1998) 3.6 RuleBender and BioNetGen and a rule-based language for modelling biochemical reactions Modern biological experiments generate large amounts of data describing intracellular dynamics. Rule-based languages for describing intracellular biochemistry allow for the construction and simulation of detailed models with unprecedented scope and precision. Rule-based Models can be used to suggest new hypotheses and new ideas for future experimentation. Rule-based models can be difficult to understand and time consuming to debug. RuleBender is a free tool for constructing, debugging, simulating and analyzing rule-based biological models in the BioNetGen language. 3.7 Flux Balance Analysis using OptFlux Flux balance analysis is a mathematical approach for analyzing the flow of metabolites through a metabolic network. (Jeffrey D. Orth at el, 2010). OptFlux is an open-source and modular software to support in silico metabolic engineering tasks aimed at being the reference computational application in the field. 3.8 Stochastic simulation algorithm using BioNetGen Stochastic Simulation Algorithm (GSSA) and its variants are cornerstone techniques to simulate reaction kinetics in situations where the concentration of the reactant is too low to allow deterministic techniques such as differential equations. To generate a network in advance of a simulation, which may subsequently be carried out either by numerically solving ODEs or by implementing a stochastic simulation algorithm (SSA) (Gillespie, 2007) 4 Results 4.1 Ribose and Guanosine triphosphate (GTP) The non-specific precursors of riboflavin biosynthesis which are commonly used are guanosine triphosphate and ribulose-5-phosphate. Ribose and Guanosine triphosphate (GTP) is biologically responsible for the production of riboflavin with the help of enzymes (RIB1, RIB2, RIB4, RIB5, RIB7) that catalyze reactions to produced riboflavin, with help of (FMN) to produced Flavin mononucleotide. After that, produced Flavin adenine dinucleotide (FAD). It also has the role of a source of energy or an activator of substrates in metabolic reactions, like that of ATP, but more specific. It is used as a source of energy for protein synthesis and gluconeogenesis. GTP is essential to signal transduction, in particular with G-proteins, in second-messenger mechanisms where it is converted to guanosine diphosphate (GDP) through the action of GTPases. Rib A helps stimulate the production of riboflavin and direct link between the other proteins. In addition, is responsible for the outputs of the biochemical processes. Fig. 2.Rib A Simulation Fig. 3. Metal Phosphate Simulation Metal Phosphate, which is linked to a protein. Motivating force proton transfer of phosphate is associated. Thus, the transfer of the phosphate group of phosphate, ATP to amino acids. In conclusion, it is easy to extract the pure biological industries inexpensive and riboflavin without toxic substances. Therefore, it is expected in the future production of large amounts of riboflavin and faster by yeasts and fungi away from harmful chemicals. 4.2 Dynamic modelling of Riboflavin biosynthesis in Bacillus subtilis using SBML Riboflavin synthase catalyzes the transformation of 6,7-dimethyl-8-ribityllumazine into riboflavin in the last step of the riboflavin biosynthetic pathway. Gram-negative bacteria and certain yeasts are unable to incorporate riboflavin from the environment and are therefore absolutely dependent on endogenous synthesis of the vitamin. (Markus F. et at, 204) Provided a strategy for developing and implementing genome-scale metabolic models. Simultaneous growth and riboflavin overproduction using Bacillus subtilis were investigated using Michaelis-Menten enzyme reaction model using different modern software packages for computer biochemical systems simulation. Fig. 4.Riboflavin Wild Type Simulation of Riboflavin biosynthesis in Bacillus subtilis After re-simulating Riboflavin level has been changed. The added reactions corresponds to a sharper transition of riboflavin level from the beginning of the simulated riboflavin function. Fig. 5.Riboflavin Wild Type Simulation of Riboflavin biosynthesis in Bacillus subtilis 4.3 RuleBender and BioNetGen Riboflavin simulation Rule-based modeling involves the representation of molecules as structured objects and molecular interactions as rules for transforming the attributes of these objects. Fig. 6.Simulation of Riboflavin biosynthesis in Bacillus subtilis using RuleBender The approach allows systematic incorporation of site-specific details about protein-protein interactions into a model for the dynamics of cell regulatory networks, including processes involving cell signaling, gene regulation, and metabolism. The consequences of protein-protein interactions are difficult to specify and track with a conventional modeling approach because of the large number of protein phosphoforms and protein complexes that these interactions potentially generate. Here, we demonstrate how a rule-based model is specified in the BioNetGen language (BNGL) using the RuleBender interface and demonstrate how models can be visualized, simulated, and analyze. 4.4 Riboflavin Flux Balance Analysis Provided a strategy for developing and implementing genome-scale metabolic models. Calculated Synthesis and Biomass Flux values. Simultaneous growth and riboflavin overproduction using Bacillus Subtilis were investigated using Michaelis-Menten enzyme reaction SBML model and OptFlux Software Platform for Metabolic Engineering. Fig. 7.Wild Type Simulation of Riboflavin biosynthesis in Bacillus subtilis using OptFlux Fig. 8.Riboflavin Synthesis and Biomass Flux Values 4.5 Using the SSA with BioNetGen Which Type of Model to use ODE or SSA? At low molecule numbers: use stochastic model. At high molecule numbers: both types of simulation can be used, but SSA might require more â€Å"computing power† than the ODE simulation. When the number of molecules is high SSA behaves like ODE. A chemical reaction model can be converted into a set of ordinary differential equations (ODEs) using the law of mass action. This law states that the rate of a reaction is its rate constant times the concentration of the reactants raised to the power of their stoichiometry. A better method for reasoning about genetic circuits is to utilize Gillespie’s stochastic simulation algorithm (SSA) (Gillespie, 1977). There are several variants of the SSA. This paper uses the direct method, which is shown in Algorithm 1. The SSA takes a chemical reaction network model, M, and computes a time series simulation ÃŽ ±. The SSA is essentially a Monte Carlo algorithm, which treats each reaction as a random event. For the SSA convert concentration to molecule number is using the formula: (1) Where is number of molecules, is concentration, is the Avogadro’s constant and is cell volume. Avogadro’s constant = 6.221023 Table 2. Riboflavin Enzymes convert concentration to molecule number |Enzyme| Value C Value n R5P 5e-07 5 RibA 2e-07 2 D2B4P 5e-07 5 RibH 5e-07 5 RibE 2e-07 2 Fig. 9. Simulating Riboflavin using the SSA with BioNetGen Fig. 10. Simulating Riboflavin using the SSA with BioNetGen CONCLUSION Provided a strategy for developing and implementing genome-scale metabolic models. Calculated synthesis and biomass flux values. Simultaneous growth and riboflavin overproduction using Bacillus Subtilis were investigated using Michaelis-Menten enzyme reaction SBML. An in-silico model of riboflavin biosynthesis in B. subtilis has been successfully developed. Designed the strategy for efficient producing of riboflavin in B. subtilis. The Riboflavin biosynthesis modeling has done using stochastic simulation algorithm. Presented methodology of Riboflavin biosynthesis modeling simulation using such software complexes as: RuleBender and BioNetGen, OptFlux, String and Ondex. Designed dynamic modelling of Riboflavin biosynthesis in Bacillus subtilis using SBML language. References Benfenati E., Oonella Diaza R., Cassano A., Pardoe S., Gini GM Mays C., KnaufR., Benighaus L. (2011) The acceptance of in silico models for REACH: Requirements, barriers, and perspectives, Chemistry Central Journal, Vol 5, pp 58. http://joumal.chemistry central com/content/5/1/58. Boyle NR, Morgan JA (2009) Flux balance analysis of primary metabolism in Chlamydomonas reinhardtii. BMC Syst Biol 3: 4 Bretzel, W Schurter, W Ludwig B, Kupfer, E Doswald, S Pfister, M van Loon, A (1999). Commercial riboflavin production by recombinant Bacillus Subtilis: down-stream processing and comparison of the composition of riboflavin produced by fermentation or chemical synthesis. Journal of Industrial Microbiology and Biotechnology. Issue 1, pp 19-26. Chem. 58, 36-57 Feng X, Xu Y, Chen Y, Tang YJ (2012) Integrating flux balance analysis into kinetic models to decipher the dynamic metabolism of Shewanella oneidensis MR-1. PLoS Comput Biol 8: e1002376 Gillespie, D. T. (1977). Exact stochastic simulation of coupled chemical reactions. J. Phys. Chem. 81, 2340–2361. Gillespie, D. T. (2007) Stochastic simulation of chemical kinetics. Annu. Rev. Phys. Han Lim, S Soo, J Enoch, Y. (Ed.) (2001). Microbial Production of Riboflavin Using Riboflavin Overproducers, Ashbya gossypii, Bacillus Subtilis, and Candida famate: An Overview. Jeffrey D Orth, Ines Thiele, and Bernhard ØPalsson. What is flux balance analysis? Nature biotechnology, 28(3):245– 8, March 2010. Li Wu, Q Chen, T Gan, Y Chen, X Ming Zhao, X (2007). Optimization of riboflavin production by recombinant Bacillus Subtilis RH44 using statistical designs. Applied Microbiology and Biotechnology. Issue 4, pp 783-794. Markus F., Werner R., Sabine S., Boris I., Gerald Richter, Felix R., Wolfgang E., Adelbert B. (2004) â€Å"Evolution of vitamin B2 biosynthesis: structural and functional similarity between pyrimidine deaminases of eubacterial and plant origin.†, J. Biol. Chem., vol. 279, no. 35 Rhoads DG, Pring M. The simulation and analysis by digital computer of biochemical systems in terms of kinetic models. IV. Automatic derivation of enzymic rate laws. J Theor Biol. 1998 Sep;20(3):297–313. Sauer U., Hatzimanikatis V., Bailey J.E., Hochuli M., Szyperski T., Wà ¼thrich K. (1997) Metabolic fluxes in riboflavin-producing Bacillus subtilis. Nat. Biotechnol. 15, 448–452. Stolz, J (2012). Riboflavin. http://www.nutrition.tum.de/index.php?id=115L=1 (29oct. 2014). 1 [*]To whom correspondence should be addressed.

Caribbean Literature 1900-1929 Essay

In the Caribbean, specifically Jamaica, during the year of 1900-1929 there were two poets whose work truly stood out and made a statement. Those poets were Thomas Redcam (1870-1933) and Claude McKay (1878-1972). Thomas Redcam was a Jamaican poet who came from Irish ancestry. Throughout his life he promoted Jamaican literature and was a notable poet. He was seen as helpful and encouragement to younger poets during that time period including Claude McKay. Both poets had such a love for their country and made it clear in their poems. They spoke about the beauty of Jamaica, how through thick and thin Jamaica would always be their heart and homeland. The themes of nature, faith, and love are very present within the six poems we read. In Thomas Redcam’s poem â€Å"My Beautiful Home† is where we first see the predominant themes of nature, faith, and love. In this poem he speaks of Jamaica in such vivid terms, making mention of the land only using words such as beautiful. He al so speaks of how the beauty of his homeland is the main reason he loves his land. He speaks of this love as being like none other, a love that is strong and rooted in heart and faith. He makes the idea clear that even when not there his love will always be there and living strong â€Å"Whenever I wander, for thee my love is abiding and strong† (Redcam, 45). This idea for love of country and nature continues in his next poem â€Å"O’little Green Island. Far Over the Sea† In this poem he speaks of the English rule throughout Jamaica and how it affects the people in one aspect but not at all in the other. Not once does he speak negatively about England but rather Redcam glorifies and praises their rule, â€Å"For England is England, brave, patient, and true.† (Redcam, 46) He speaks of how no matter who is ruling they are themselves and their love will always be for their own land Jamaica, â€Å"But my little Green Island, far over the sea, At eve-tide Jamaica, my heart turns to thee.† (Redcam, 46). Redcam makes it clear through his statement that living under the English rule does not change them but rather strengthens their love for their own homeland, Jamaica. Lastly but not least once again the themes of love and nature continue in Redcam’s last poem, but the theme of faith also becomes present in â€Å"Jamaica’s Coronation Ode†. Here more than in his other poems we find the idea of love for the country and how our love along with the idea of a strong faith helps us stand true to our country. He speaks of the war and hard times the people of Jamaica went through during this time period and although he speaks of hardship not once does he mention the idea of negativity. He speaks of positive thoughts and how their love and faith in Jamaica will carry them through the war and as a matter of fact anything in life. He speaks positive thoughts of the moving forward and the future to come after the war â€Å"We are marching to conquer the Future, We are sons of Jamaica the free, We are true to our King and our Country, We are heirs of the ages to be† (Redcam, 47) . This quote is mentioned throughout the poem which really shows the true faith and how they truly believed they could conquer it all with their strong love for Jamaica. Although in this poem the idea of faith in love is more predominant Redcam still mentions the nature he always speaks so highly of. A few years after the poems of Redcam came those of Claude McKay. Claude McKay was a Jamaican- American poet who inspired and guided by the work of Redcam. He was the author of four novels during the time of the Harlem Renaissances. During that time he also wrote short stories and poems, which he is noted for all around the world. McKay being under the guidance of Redcam had the same predominant themes in his poems which were that of nature, love, and faith. In McKay’s first poem â€Å"My Native Home Land† the theme of love is instantly noticeable. He speaks of his love and how he feels Jamaica has made him the man he is. He speaks of how through thick and thin he will stand by his country and defend her. â€Å" Though you cas’ me from your breas’ An’ trample me to deaf; My heart will trus’ you none de less, My land I won’t forget† (McKay, 64). McKay makes it clear that he will never lose his faith in Jamaica as well. In this poem Jamaica McKay believes that his country Jamaica is one that can do no wrong. Once again the ideas of nature, faith and love continue in his next poem â€Å"Flame Heart† but this time the idea of nature becomes the more predominant one. In this work Mckay speaks how he returns back to Jamaica. Throughout the poem he mentions aspects of nature in the land that he can not recall but then mentions the one part of the land he remembers so vividly. McKay speaks of poinsettia plants he remembers so well. He speaks of their beauty and color, â€Å"I have forgotten much, but I still remember The poinsettia’s red, blood-red in warm December† (McKay, 70). He speaks so vividly of this one aspect of nature and not only does he mention the beauty of it, he dives even deeper when mention of climate comes in. This statement about the poinsettia is mentioned a few times in the poem giving off a strong vibe and conviction. Everything to the speaker is a blur but through his blurs he is able to remember that red-blood poinsettia plant in warm December month. In his last poem â€Å"I Shall Return† McKay touches upon all three themes in such strong points. This poem speaks of someone who has left Jamaica but has intentions of returning. It speaks of how their love for their country has encouraged them to return, but also their faithfulness. It speaks of how they truly miss the nature and beauty of their homeland. The speaker believes that when they return they will save themselves pain. â€Å"I shall return, I shall return again, to ease my mind of long, long years of pain† (McKay, 71). Jamaica is their home and they feel it is where they belong no matter where life takes them. Their faith and love will always bring them back home. These 6 poems are the perfect example of the three themes mentioned. Not one of them speaks of negative thoughts of Jamaica but instead glorifies all its beauty and the faith and love the Jamaican people truly have for their land. Both poets have such strong and impacting words when speaking of their land and truly believe that there is no land like theirs. They speak from the heart and truly give their readers vivid ideas of what things were like. These poets are have left a good impact on not only the people of their homeland but of people around the world through their poems. Works Cited Donnell, Alison, and Sarah Lawson Welsh. Routledge Reader in Caribbean Literature. London: Routledge, 1996. Print.

The Effects Of Free Speech On Hate Groups, And The...

The relationship of free speech to that of hate groups, and the protection of hate speech under the first Amendment is a much debated topic of ethics and civil liberties. Although affirmative action protects against discrimination of race, religion, gender, and disabilities, it only protects from discrimination in educational institutions and employment. So how can a society that claims to protect civil rights allow the production of speech that opposes those same fundamental rights?, because this form of speech, although hateful in nature, are ideas, and â€Å"hateful ideas are just as protected under the First Amendment as other ideas† (McGough, 2015). The First Amendment guarantees freedoms in regards to expression, religion, assembly, and the right to petition. Freedom of expression prohibits Congress from restricting the press or individuals the right to speak freely (Legal Information Institute, n.d.). A hate group, as stated by Walker Bragman, is â€Å"an organized group or movement that advocates and practices hatred, hostility, or violence towards members of a race, ethnicity, religion, gender, sexual orientation or other designated sector of society† (Bragman, 2012). Speech that attacks, demeans, threatens, or insults a person or group on the basis of origin, ethnicity, religion, gender, gender identity, sexual orientation or disability is identified as hate speech (Random House Dictionary, 2016). By design the First Amendment was established to protect unpopular speech,Show MoreRelatedHate Speech And Freedom Of Speech1393 Words   |  6 Pages In the name of free speech, hate speech should not be tolerated. Hate speech has devastating effects on the people and communities it is targeted at. Left unchecked hate speech can lead to harmful and violent effects. 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There has been a lot of controversy over whether hate speech should be regulated. In analyzing the concept of free speech, one cannot ignore that it does not occur in a vacuum. There have been all types of debasements ranging from ethnic, religious, racial and gendered stereotyping. Freedom of speech inherently includes all other fundamental human rights. Hence, as acknowledged through naturalRead MoreHate Speech1536 Words   |  7 PagesHate Speech Essay In the first amendment of the United States constitution, American citizens are guaranteed the right to free speech. This is a fundamental right of American law, and one of the foundations of the U.S. Constitution. It is also the breeding ground for one of the most widely debated issues in America: What, if any, measures should be put into place to regulate hateful language? Most people will agree under one definition or another that hate speech is a socially deviant activity

Understanding the Concatenation of Strings in Java

Concatenation in the Java programming language is the operation of joining two strings together. You can join strings using either the addition () operator or the String’s concat() method. Using the Operator Using the operator is the most common way to concatenate two strings in Java. You can provide either a variable, a number, or a String literal (which is always surrounded by double quotes). To combine the strings â€Å"I’m a† and â€Å"student†, for example, write: I’m a student Be sure to add a space so that when the combined string is printed, its words are separated properly. Note above that student starts with a space, for example. Combining Multiple Strings Any number of operands can be strung together, for instance: I’m a student ! And so are you. Using the Operator in a Print Statement Frequently, the operator is used in a print statement. You might write something like: System.out.println(pan handle); This would print: panhandle Combining Strings Across Multiple Lines Java disallows literal strings to span more than a line. Using the operator prevents this: String quote Nothing in all the world is more dangerous than sincere ignorance and conscientious stupidity.;   Combining a Mixture of Objects The  operator   normally acts as an  arithmetic operator  unless one of its  operands  is a String. If so, it converts the other operand to a String before joining the second operand to the end of the first operand. For example, in the example below, age is an integer, so the operator will first convert it to a String and then combine the two strings. (The operator does this behind the scenes by calling its toString() method; you won’t see this occur.) int age 12;System.out.println(My age is age); This would print: My age is 12 Using the Concat Method The String class has a method concat() that performs the same operation. This method acts on the first string and then takes the string to combine as a parameter: public String concat (String str)​   For example: String myString I have decided to stick with love.;myString myString.concat( Hate is too great a burden to bear.);System.out.println(myString); This would print: I have decided to stick with love. Hate is too great a burden to bear. Differences Between the Operator and the Concat Method You may be wondering when it makes sense to use the operator to concatenate, and when you should use the concat() method.  Here are some differences between the two: The concat() method can combine only String objects — it must be called on a String object, and its parameter must be a String object. This makes it more restrictive than the operator since the operator silently converts any non-string argument to a string.The concat() method throws a NullPointerException if the object has a null reference, while the operator deals with a null reference as a â€Å"null† string.The concat()) method is capable of combining only two strings – it cannot take multiple arguments. The operator can combine any number of strings. For these reasons, the operator is more often used to combine strings. If you are developing a large-scale application, however, performance can differ between the two because of the way that Java handles string conversion, so be aware of the context in which you are combining strings.