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(2015). Genome editing modifies a specific genome in precise and predictable manner. The products, which are obtained by editing through CRISPR-Cas9, have no exogenous DNA and furthermore editing can be done in such a way, which abides by all the rules and regulations that are complaisant to withstand against Genetically Modified issues and can get an easy approval by the Department of Agriculture (USDA). In addition, the complexity of wheat genome together with time-consuming tissue culture techniques has made it difficult for scientists to undergo ambitious genome editing projects via CRISPR/Cas9 in this important cereal. The immunity is acquired through the integration of short fragment of the foreign DNA as spacer between two adjacent repeats at the proximal end of a CRISPR locus [29]. Plant J. The type II adaptive immunity system by CRISPR/Cas9 involves three stages, which are acquisition, crRNA biogenesis and interference to cleavage the DNA target, resulting in the formation of DSBs. 7:475. doi: 10.3389/fpls.2016.00475. PubMed  Mol. After that, it will induce DSBs on the foreign DNA that carries the same protospacer sequence, which is accompanied by the protospacer adjacent motif (PAM) to disable the virus [7, 30]. As a result, the plant now displays red fluorescence. Rev. Sci. (2008). doi: 10.1007/s00239-004-0046-3, Moscou, M. J., and Bogdanove, A. J. A genome-wide association study reveals new loci for resistance to clubroot disease in Brassica napus. Targeted mutagenesis, precise gene editing, and site-specific gene insertion in maize using Cas9 and guide RNA. Cas9 nickase has also emerged as an alternative to reduce off target effects. Genome editing with engineered nucleases (GEEN) has evolved as a highly specific and efficient tool for crop improvement with the potential to rapidly generate useful novel phenotypes. (2001). He defined 45 CRISPR-associated (Cas) protein families that are categorized into core proteins (Cas1, Cas2, Cas3, Cas4, Cas5, Cas6), 8 CRISPR/Cas subtypes and RAMP (repair associated mysterious protein) module in prokaryotic genomes. (2015). CRISPR loci were discovered about two decades ago in Escherichia coli following viral infection as part of the immune system through which the bacteria keep a genetic memory of the viral genome fragments in order to defend against future infections if any other foreign DNA sequences match the unique spacer sequences in the memory (also known as protospacers) [2, 3]. Additionally, the DNA target specificity duplex is provided by what is called the ‘seed sequence,’ which is located at approximately 12 bases upstream of the PAM sequence that matches the RNA with the DNA target [29] (Figure 4). Genomes of other grain crops edited through this technique include maize, sorghum, barley, etc. Interestingly, four independent groups (Shan et al., 2013; Brooks et al., 2014; Zhang et al., 2014; Zhou et al., 2014) have demonstrated the introduction of biallelic or homozygous mutations in T1 generation of rice and tomato indicating the high efficiency of this system. ScienceDirect ® is a registered trademark of Elsevier B.V. ScienceDirect ® is a registered trademark of Elsevier B.V. Use of CRISPR/Cas9 for Crop Improvement in Maize and Soybean. Biotechnol. The transcript levels were reduced approximately by 60% as compared to control among the three independent transgenic lines. This was specifically aided by the discovery of nuclease enzyme, the utilization of different bacterial plasmids for cloning of target genes in bacteria as well as target plants and by the development of plasmid vectors carrying antibiotic resistance genes, a variety of constitutive, inducible as well as tissue-specific promoters, and different reporter genes that made the cloning and transformation easier. Proc Natl Acad Sci USA 108:20260–20264, Van de Wiel C, Schaart J, Lotz L, Smulders M (2017) New traits in crops produced by genome editing techniques based on deletions. Cas9 is a huge protein possessing multiple domains (RuvC domain at the amino terminus and the HNH nuclease domain positioned in middle) and two small RNAs namely crRNA and tracrRNA. Mol. doi: 10.1038/nrm.2015.2, Esvelt, K. M., Mali, P., Braff, J. L., Moosburner, M., Yaung, S. J., and Church, G. M. (2013). CRISPR/Cas9 applications in crop genetic improvement. These genome-editing technologies use programmable nucleases to increase the specificity of the target locus. doi: 10.1073/pnas.1420294112, Xie, K., and Yang, Y. Simultaneous editing of three homoeoalleles in hexaploid bread wheat confers heritable resistance to powdery mildew. doi: 10.1038/nbt.3117, Tsutsui, H., and Higashiyama, T. (2017). Then, the CRISPR will be transcribed into a long precursor crRNA (pre-crRNA) with the help of Cas proteins (Cas1, Cas2, Cas9 and Cas4/casn2) or simply known as biogenesis. Biotechnol. Plant Biotechnol J 15:1509–1519, Puchta H (2004) The repair of double-strand breaks in plants: mechanisms and consequences for genome evolution. Nat. Once the Cas9 protein is activated, it recognizes DNA target by binding with DNA sequence that matches its PAM. 33, 187–197. An example of CTLs is constructed through the CRISPR/Cas9 system by specifically inserting the transgenes into the desired region in the genome through HDR.

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