next generation sequencing quality control

WebStep 2 in NGS Workflow: Sequencing. Synthetic lethality occurs when a combination of deficiencies in two genes leads to cell death, whereas deficiency in only one gene results in a viable phenotype [50]. Next-generation sequencing is based on the ability to sequence, in parallel, millions of DNA fragments, and introduction of NGS technology has resulted in a dramatic increase in speed and content of sequencing at a fraction of the cost.44 Described briefly, first a DNA library is prepared from the patients sample by fragmentation, purification and amplification of the DNA sample. ScienceDirect is a registered trademark of Elsevier B.V. ScienceDirect is a registered trademark of Elsevier B.V. Memorial Sloan-Kettering Cancer Center, New York, United States, David Geffen School of Medicine at UCLA, Los Angeles, United States, Kazusa DNA Research Institutes, Kisarazu, Japan, Accurate Results in the Clinical Laboratory (Second Edition), Pathology, Biomarkers, and Molecular Diagnostics, Abeloff's Clinical Oncology (Sixth Edition), Genetics, molecular biomarkers, and artificial intelligence to improve diagnostic and prognostic efficacy, Principles of Translational Science in Medicine (Third Edition), Genetics of Cardiovascular Disease and Applications of Genetic Testing, High-Throughput Methods in Molecular Pathology, Molecular and genomic basis of bronchial asthma, Advances in Molecular Genetics Including Fetal Sequencing. Li Y, Xi S, Yong J, Wu X, Yang X, Wang F. Morphologic, Immunohistochemical, and genetic differences between high-grade and low-grade fetal adenocarcinomas of the lung. The cost of NGS based carrier screening, which in the past has been a major deterrent, is also gradually decreasing. Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. conducted a prospective nested casecontrol study to analyze the house dust, moisture, and mold present in a household and relate it to later life asthma risk. The histone methyltransferase EZH2 is a therapeutic target in small cell carcinoma of the ovary, hypercalcaemic type. 2019;9:550. /index.php [L] 2014;46:4279. 8 0 obj The quality of the NGS library is the factor with the most influence on the success of the sequencing run, affecting both the sequence validity and the number of reads. Methods Association for Clinical Genetic Science. Arch Pathol Lab Med United States. A few studies have been conducted utilizing high-throughput NGS for asthma risk assessment, diagnosis, and treatment guidance. RNA-seq, Nextera, or bisulfite-converted DNA. Google Scholar. Google Scholar. Cancer Discov. 2018;36:63341. Reportedly, ARID1A-deficient gastric cancer cells are vulnerable to the AKT inhibitor, GSK690693, and the addition of GSK690693 possibly potentiates the suppressive function of conventional chemotherapy [105]. Prevalence of recurrent oncogenic fusion in mismatch repair-deficient colorectal carcinoma with hypermethylated MLH1 and wild-type BRAF and KRAS. CAS Basic local alignment search tool. Gut England. It should be highlighted that NGS-based gene panel testing is usually complemented with array comparative genomic hybridization and other ancillary methods, thus yielding a comprehensive and feasible approach for heterogeneous disorders.43 Based on recent evidence and marketing data, single tests are frequently used since they are suitable for conditions with distinct clinical features and minimal site heterogeneity; however, these data are constantly changing due to the rapid development of new technologies in genetic testing.12, Andrea Padoan, Mario Plebani, in Principles of Translational Science in Medicine (Third Edition), 2021. The sequence data, then, are compared with public databases such as ExAc and 1000 Genomes Project to exclude high prevalent and normal variants. p<0.1. This varies by browser, if you do not see a box on your page with a red X try right clicking on the page, then select View Page Info, and goto the Media Tab. LOF variants of the SWI/SNF complex can influence the response to ICIs by increasing the infiltration of CD8+T cells, enhancing the cytotoxicity of T cells [70], or by creating an immune-responsive milieu [24]. Chabanon RM, Morel D, Eychenne T, Colmet-Daage L, Bajrami I, Dorvault N, et al. Cancer Res. 2021;22:5869. Spatial immunoprofiling of the intratumoral and peritumoral tissue of renal cell carcinoma patients. Available online at: Chikhi R., Medvedev P. (2013). 2016;34:15563. 24, 25. 2021;7:14453. 2017;355:11528. Springer Nature. PubMed PD-1 blockade in tumors with mismatch-repair deficiency. and transmitted securely. Gounder MM, Zhu G, Roshal L, Lis E, Daigle SR, Blakemore SJ, et al. Cancer Immunol Res. Notably, PARP inhibitors targeting BRCA1/2-mutant tumors represent a notable example of such synthetic lethality [82]. Low-quality bases can be easily trimmed; lots of low quality bases may indicate a poor library, or problems with phasing. Given that most SWI/SNF mutations were dispersed along the full length of each gene, NGS showed potential as the most suitable strategy for detecting SWI/SNF alterations. Nat Genet. Still, there are some limitations not only on the performance of genetic tests but also on the complexity of choosing the appropriate genetic test. A major chromatin regulator determines resistance of tumor cells to T cellmediated killing. <> The lysine-specific methyltransferase KMT2C/MLL3 regulates DNA repair components in cancer. The details of each of the abovementioned platforms are beyond the scope of this chapter. Oncol Rep Greece. Background: In recent years, metagenomic Next-Generation Sequencing (mNGS) has increasingly been used for an accurate assumption-free virological diagnosis. It was observed that lower fungal diversity of different taxonomic groups leads to an increased risk of asthma development in later life probably due to immunological priming. Identifying recurrent mutations in cancer reveals widespread lineage diversity and mutational specificity. The present study is the Matthijs, G. et doi: 10.1371/journal.pone.0060234. Notice that the CaSe is important in this example. Targeted next-generation sequencing identifies novel variants in candidate genes for Parkinsons disease in Black South African and Nigerian patients. This site needs JavaScript to work properly. Therefore, SWI/SNF-deficient cancers are vulnerable to the inhibition of homologous recombination repair factor, PARP1 [20, 49], cell cycle regulator, cyclin-dependent kinase (CDK)4/CDK6 [28, 56], DNA replication checkpoint factor, ATR [57], chromosomal segregation factor, Aurora kinase A [58], and oxidative phosphorylation [59] and glutathione [60] pathways. stream Circulating tumor DNA analyses predict progressive disease and indicate trastuzumab-resistant mechanism in advanced gastric cancer. Charles River Laboratories International, Inc. (NYSE: CRL) today announced the launch of Accugenix Next Generation Sequencing for Bacterial Identification and Fungal Identification (Accugenix NGS). 2018;9:3212. However, quality control (QC) of sequence data Oncogene. MeSH It is possible to perform whole-genome, whole-exome, and targeted-region sequencing using the NGS technology. This was the first whole-exome sequencing study performed for variant identification in an asthmatic family [68]. Next-generation sequencing (NGS) has been extensively applied as a cost-effective diagnostic tool in clinical practice and trials . A CRISPR/Cas9-engineered ARID1A-deficient human gastric cancer organoid model reveals essential and nonessential modes of oncogenic transformation. Careers. 15.7. Okamura R, Kato S, Lee S, Jimenez RE, Sicklick JK, Kurzrock R. ARID1A alterations function as a biomarker for longer progression-free survival after anti-PD-1/PD-L1 immunotherapy. FOIA Among the 58 cases of lung cancer with SMARCA4 LOF mutations in our study, the mutation rates corresponding to the above hot genes were almost consistent with the previously reported rates of 74.1%, 31.0%, 24.1%, 20.7%, and 15.5% for TP53, KEAP1, STK11, KRAS, and EGFR, respectively. Nat Commun. Proc Natl Acad Sci. Loss of PTEN promotes resistance to T cell-mediated immunotherapy. WebFor next-generation sequencing. Mod Pathol. DAfonseca V, Arencibia AD, Echeverra-Vega A, Cerpa L, Cayn JP, Varela NM, et al. NGS has been used to detect novel causative variants of a single gene and common disorders of the cardiovascular system including familial hypercholesterolemia, different types of cardiomyopathies, long QT syndrome, CHD, and thoracic aortic aneurysms and dissections (TAAD). Comparison and integration of deleteriousness prediction methods for nonsynonymous SNVs in whole exome sequencing studies. endobj Most of these variants were located in three asthma candidate genes, that is, PDE4DIP, CBLB, and KALRN. Sequencing Quality Control Phase 2 (SEQC2) is the fourth project of MAQC that performs analytical validation of NGS applications. 2002;62:3238. Loss of ARID1A activates mTOR signaling and SOX9 in gastric adenocarcinomarationale for targeting ARID1A deficiency. PRC2-mediated repression of SMARCA2 predicts EZH2 inhibitor activity in SWI/SNF mutant tumors. Interseasonal differences were also observed. M$Oo2m{1R@EUJ#Xy8>ci3H wa#`_m>+27& jofvWfr;+j\#,tB=`\Bcy9.RL. FCW:u5=&F$G$'F&j#"K&f,x,to`JrnywHWpZ2gP &4 Front Immunol. Mammalian SWI/SNF complex genomic alterations and immune checkpoint blockade in solid tumors. Cold Spring Harb Perspect Med. Alterations of DNA damage response pathway: biomarker and therapeutic strategy for cancer immunotherapy. Mol Med. (1990). 2014;20:217. Next generation sequencing (NGS) refers to large-scale DNA sequencing technology that allows for querying the entire genome (whole genome), the Please enable it to take advantage of the complete set of features! Part of 2016;34:220611. Traditional sequencing methods are limitedwhich is why we developed Accugenix NGS to provide more efficient and effective results, reducing downtime and uncertainty. Greg Marshall, Corporate Vice President & General Manager, Microbial Solutions, Charles River 2021;45:146475. 13 0 obj 2014 May 3;9:8. doi: 10.1186/1751-0473-9-8. The TMB-H and MSI-H ratios corresponding to SWI/SNF-mutant tumors were also significantly higher than those corresponding to the SWI/SNF-non-mutant tumors (TMB-H ratio: 44.3% vs. 10.3%, p <0.0001; MSI-H ratio: 16.0% vs. 0.9%, p <0.0001), even though the differences were not significant for certain malignancies, such as kidney cancer, pancreatic cancer, prostate cancer, and urothelial cancer. Huang DW, Sherman BT, Lempicki RA. Cancer Genet. Mutations in the SWI/SNF complex induce a targetable dependence on oxidative phosphorylation in lung cancer article. It is more important than ever that laboratories strive to develop pipelines with the highest possible performance standards and validate them as stringently as possible to reduce errors. Bis-seq. Nat Genet. Huang DW, Sherman BT, Lempicki RA. The resulting sequence data are computationally aligned against a normal reference genome.45 This enables the detection of many sequence alterations in a single reaction. endobj The results are sensitive to assumptions regarding mutation detection rates and carrier frequencies in multiethnic populations.49, Next-generation sequencingbased approaches have several shortcomings: some of the novel variants detected by NGS may have no clinical significance. Kerl K, Moreno N, Holsten T, Ahlfeld J, Mertins J, Hotfilder M, et al. SWI/SNF-deficiency defines highly aggressive undifferentiated endometrial carcinoma. c The PFS of patients treated by ICIs carrying mutations in two or more SWI/SNF genes was not significantly different from that of patients with mutations in single gene. Nat Genet. Elife. 2021;145:908. Additionally, the survival analysis for individual cancer types suggested that the PFS of the SWI/SNF-mutant group was significantly superior to that of the SWI/SNF-non-mutant group in colorectal cancer (NR vs. NR, HR=0.33 [0.190.59], p=0.0001; Additional file 2: Figure S2a) and gastric cancer (NR vs. 20.6months, HR=0.44 [0.190.97], p=0.0437; Additional file 2: Figure S2b); the same tendency was significant numerically but not statistically in non-small cell lung cancer (NR vs. 40.9months, HR=0.58 [0.331.02], p=0.0595; Additional file 2: Figure S2c). Ultimately the best QC of your NGS experiments is likely to come from the sequence data and, because of this, many labs will run a QC lane first. 2019;25:206471. The more time and effort you spend on QC the better quality your results and conclusion will be. WebAs a quality feature, an internal control consisting of 11 human mRNA targets is included in every sample. It is possible that you may need to edit the .htaccess file at some point, for various reasons.This section covers how to edit the file in cPanel, but not what may need to be changed. A series of inhibitors, ranging from chemical probes to FDA-approved drugs, that target the synthetic lethal partners of SWI/SNF members have been shown to exhibit clear therapeutic effects in several cancers [20, 21, 25, 48,49,50,51,52,53,54,55,56,57,58,59,60, 81,82,83,84,85,86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105]. These agreements bring together Charles Rivers curated, proprietary microbial identification database and the high-quality extensive testing services of Roche GS-FLX 454 Genome Sequencer: pyrosequencing. A hybrid captured-based NGS assay covering approximately 1.1 megabases (Mb) of the genomic sequences of 1021 cancer-related genes (GenePlus-Beijing, China) was used for the sequencing, which was performed using a GenePlus 2000 sequencing system (Beijing, China) with 2 100 bp paired-end reads. 2010;18:31628. Tessier-Cloutier B, Coatham M, Carey M, Nelson GS, Hamilton S, Lum A, et al. 3). 2015. 2021;34:222941. This selection of articles and reviews provide resources for assay design, quality assurance, quality control, data management and results analysis and interpretation. 2017;16:85060. Specifically, ARID, which is a conserved helix-turn-helix motif-containing domain, plays a role in recruiting SWI/SNF to the target gene promoters, whereas the function of the BAF250_C domain, which contains motifs, such as NES and LXXLL-motif, that putatively mediate proteinprotein interactions, is still unknown [63]. Bitler BG, Wu S, Park PH, Hai Y, Aird KM, Wang Y, et al. This will reset the permalinks and fix the issue in many cases. 2021. https://doi.org/10.1038/s41467-021-21893-y. Google Scholar. endstream Nucleic Acids Res. Newer approaches to overcome some of these difficulties are under development. If you do not allow these cookies we will not know when you have visited our site, and will not be able to monitor its performance. WebQuality control procedures monitor whether each component Gargis et al., Assuring the quality of nextgeneration sequencing in clinical laboratory practice. Sensitive detection of somatic point mutations in impure and heterogeneous cancer samples. Regardless of the origin of the starting material, successful sequencing experiments require purification of high-quality nucleic acids. sharing sensitive information, make sure youre on a federal *mfq3+sj8M= Bitler BG, Aird KM, Garipov A, Li H, Amatangelo M, Kossenkov AV, et al. These synthetic lethal interactions can be classified under four main categories. The clinicopathological significance of SWI/ SNF alterations in gastric cancer is associated with the molecular subtypes. The continuing decrease in massively parallel or next-generation sequencing (NGS) costs has enabled large projects consisting of hundreds or thousands of samples. Article Here, NGS in combination with sequence enrichment steps via SISPA allowed the obtention of complete CDS of WNV directly from clinical samples. However, the quality of this data is not always guaranteed. Cell Rep. 2018;22:3393400. 3 0 obj WebNext-generation sequencing (NGS) technologies have dramatically expanded the breadth of genomics. The PFS of SWI/SNF-mutant and SWI/SNF-non-mutant groups were not markedly different (Additional file 2: Figure S2dh) or could not be analyzed owing to the small sample size in the other malignancies. CAS Some of these errors may mimic biological signals (mutation), thus leading to incorrect conclusions. J Clin Oncol. Unable to load your collection due to an error, Unable to load your delegates due to an error. The companies currently marketing next-generation automated techniques are as follows: ABI SOLiD platform: sequencing by ligation; DNA nanoball sequencing by BGI Retrovolocity: sequencing by hybridization and ligation; Illumina/Solexa Genome Analyzer: sequencing by synthesis; Ion Torrent: Proton/PGM sequencing: sequencing by synthesis; and. The Centers for Disease Control and Prevention (CDC) cannot attest to the accuracy of a non-federal website. 2018;24:104757. WebNext Generation Sequencing. WILMINGTON, Mass., May 01, 2023--Charles River launches Accugenix NGS for bacterial identification and fungal ID; announces partner lab initiative to expand global coverage. J Clin Invest. Cancer Cell. A powerful diagnostic and surveillance tool like next generation sequencing (NGS) demands an equally powerful quality management system (QMS) to assure consistent, reliable data at any scale. 4d). J Thorac Oncol. This may put a strain on laboratory personnel, genetic counsellors and physicians alike. S1 The distributions of variant allele frequencies (VAFs) of ARID1A, ARID1B, ARID2, PBRM1, SMARCA4, and SMARCB1. Both methods use proprietary software and alignment to a reference sequence to produce the final template sequence. statement and In addition to QC tools, it consists of many subsidiary tools for handling and processing of data obtained from Illumina and Roche 454 sequencing platforms. Updated May 2014. Truncating mutations of hSNF5/INI1 in aggressive paediatric cancer. Chan TA, Yarchoan M, Jaffee E, Swanton C, Quezada SA, Stenzinger A, et al. Peng W, Chen JQ, Liu C, Malu S, Creasy C, Tetzlaff MT, et al. 2017;10:41539. All authors read and approved the final manuscript. WebAssay 100 and Assay 200 for 400 x 25 l reactions (400 l) each; QuantiMIZE Control gDNA (50 l); RNase- and DNase-free water (1 ml x 4); GeneRead qPCR SYBR Green Mastermix (1.35 ml x 4) $478.00 CONFIGURE AT GeneGlobe QIAseq DNA QuantiMIZE Kits are intended for molecular biology applications. These cookies perform functions like remembering presentation options or choices and, in some cases, delivery of web content that based on self-identified area of interests. Evidently, the current role of NGS in asthma pathophysiology is limited to the following: discovery of new variantsrare variants as a part of missing heritability; understanding the microbiological environment of the pulmonary system as well as the external environment and its interaction with the disease; and. Moreover, it is usually difficult to know if quality features are relevant in all experimental Next-generation sequencing Quality management Data quality Standardization Validation Guideline 1. Cancers. The PubMed wordmark and PubMed logo are registered trademarks of the U.S. Department of Health and Human Services (HHS). RewriteCond %{REQUEST_FILENAME} !-d <> 2019;30:4456. The NGS Quality Initiative promotes implementation of quality management based on the CLSI quality framework for laboratories. 8600 Rockville Pike Sample quality must be tracked and maintained along the workflow to ensure that only samples of suitable quality are processed into the resource-intensive sequencing runs, because the final result is not a good time to discover a problem with the sample. The PFS of SWI/SNF-mutant and SWI/SNF-non-mutant were not markedly different in melanoma (d), soft tissue sarcoma (e), urothelial cancer (f), endometrial cancer (g) and other cancers (h). Several consensus documents have been issued with respect to NGS including laboratory standards by the College of American Pathologists (CAP)68, joint standards and guidelines on validating oncology panels and bioinformatics pipelines by CAP and the Association for Molecular Pathology (AMP),69,70 as well as joint standards and guidelines for interpreting and reporting sequence variants in cancer by CAP, AMP, and the American Society of Clinical Oncology.71 Current clinical applications for NGS have been mainly limited to the sequencing of panels of genes associated with a particular disease. Comprehensive characterization of distinct genetic alterations in metastatic breast cancer across various metastatic sites. Evaluation of POLE and pold1 mutations as biomarkers for immunotherapy outcomes across multiple cancer types. Sandhya S, Maulik A, Giri M, Singh M. Domain architecture of BAF250a reveals the ARID and ARM-repeat domains with implication in function and assembly of the BAF remodeling complex. All patients provided signed informed consent, and the study was performed in accordance with the Declaration of Helsinki. Hong AL, Tseng Y-Y, Wala JA, Kim W-J, Kynnap BD, Doshi MB, et al. Hum Mol Genet. CCND1 amplification contributes to immunosuppression and is associated with a poor prognosis to immune checkpoint inhibitors in solid tumors. https://doi.org/10.1186/s12881-020-0953-1. Oike T, Ogiwara H, Tominaga Y, Ito K, Ando O, Tsuta K, et al. 2019;8: e44161. NGS technology has been rapidly adapted to clinical testing and it is radically changing the dogma of clinical diagnostics. U &6$Z}g@$Ns]73ofopr\.lq3;pyAG"L?9Q]_ ZLb4#'Z3? Xue Y, Meehan B, Macdonald E, Venneti S, Wang XQD, Witkowski L, et al. Epigenetic antagonism between polycomb and SWI/SNF complexes during oncogenic transformation. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. Next-generation sequencing is a relatively new technology based on massively parallel sequencing (MPS). Minoli M, Cantore T, Hanhart D, Kiener M, Fedrizzi T, La Manna F, Karkampouna S, Chouvardas P, Genitsch V, Rodriguez-Calero A, Comprat E, Klima I, Gasperini P, Kiss B, Seiler R, Demichelis F, Thalmann GN, Kruithof-de Julio M. Nat Commun. D1850Tfs*33 and D1850Gfs*4, which are frameshift truncating mutations, brought about the loss of more amino acids than did R1989*. 2016;37:23541. 2020;19:117693512092215. The properties will tell you the path and file name that cannot be found. 2018;17:2279. Lee D, Yu EJ, Ham I-H, Hur H, Kim Y-S. AKT inhibition is an effective treatment strategy in ARID1A-deficient gastric cancer cells. Over the past few years, pre-clinical and clinical evidence has implicated the SWI/SNF complex as a potential predictor of response to ICIs [6]. 2015;21:14916. Sun D, Tian L, Zhu Y, Wo Y, Liu Q, Liu S, et al. There are three main areas where QC can be applied to next-generation sequencing (NGS) On the starting nucleic acid samples After library preparation Post QC-Chain: fast and holistic quality control method for next-generation sequencing data. 2016;6:20216. You can share information with colleagues by sending the Web address to this site or by sharing this PDF. Thank you for taking the time to confirm your preferences. Biegel JA, Kalpana G, Knudsen ES, Packer RJ, Roberts CWM, Thiele CJ, et al. Previous studies have revealed the existence of a potential linkage between the SWI/SNF chromatin remodeling complex and DNA repair, TMB, and MSI [6]. Rampias T, Karagiannis D, Avgeris M, Polyzos A, Kokkalis A, Kanaki Z, et al. 2011;71:322535. Romero OA, Vilarrubi A, Alburquerque-Bejar JJ, Gomez A, Andrades A, Trastulli D, et al. Selective Killing of SMARCA2- and SMARCA4-deficient small cell carcinoma of the ovary, hypercalcemic type cells by inhibition of EZH2: in vitro and in vivo preclinical models. 2016. Optimized Illumina PCR-free library preparation for bacterial whole genome sequencing and analysis of factors influencing de novo assembly. Either the entire library of fragments or only a selected subset of fragments of interest is used as templates for the synthesis of millions of short and overlapping DNA fragments. Nat Commun. 2019;32:105364. Berns K, Caumanns JJ, Hijmans EM, Gennissen AMC, Severson TM, Evers B, et al. Clinical and public health laboratory (PHL) personnel can find free tools and resources to introduce or strengthen quality management in their workflows, available online from the Next Generation Sequencing (NGS) Quality Initiative. BMC Medical Genetics. Mod Pathol. Immunologic correlates of the abscopal effect in a SMARCB1/INI1-negative poorly differentiated chordoma after EZH2 inhibition and radiotherapy. Genome Sequencing and Assembly by Long Reads in Plants. 2021;16:117687. Versteege I, Svenet N, Lange J, Rousseau-Merck MF, Ambros P, Handgretinger R, et al. endobj ARID1A-mutated ovarian cancers depend on HDAC6 activity. Genes that belong to families of highly homologous genes or have a pseudogene are also difficult to sequence. The use of next-generation sequencing technologies has raised the need for quality control assessment as required in clinical laboratories. 2016;7:13837. JAMA Oncol. Individual fragments are then physically isolated by attachment to solid surfaces or small beads. http://www.bioinformatics.babraham.ac.uk/projects/fastqc/, http://www.bioinformatics.babraham.ac.uk/projects/fastq_screen/, MR/K001744/1/MRC_/Medical Research Council/United Kingdom, Altschul S. F., Gish W., Miller W., Myers E. W., Lipman D. J. Specifically, patients carrying mutations at two or more SWI/SNF genes did not show a superior PFS than single gene mutation carriers (NR vs. NR, HR=0.85 [0.511.42], p =0.5397; Fig. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. Cancer Res. 2020;8:107584. Laboratories usually limit their reports to include variants in classes 1 and 2 only, but for some variants, it may be difficult to make a call (Table 26.2). The GO analysis was performed on all the mutated genes in 1001 SWI/SNF-mutant samples. ARID1A deficiency impairs the DNA damage checkpoint and sensitizes cells to PARP inhibitors. <> Each nucleotide incorporated into these fragments is labelled with a different coloured fluorescent probe so that the sequence or genetic code of each fragment is identifiable.

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