Risk Assessment Referecen - Methods of Plant Genetic ...
July 2018Risk Assessment Reference:Methods of Plant Genetic ModificationIntroductionPlant genetic modification, also called plant transformation, is the process of introducing a new DNA fragment into the genome of a plant. Several methods are available to generate genetically modified (GM) plants. These have been described in a number of articles, e.g. PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ib295a2FhczwvQXV0aG9yPjxZZWFyPjIwMTA8L1llYXI+
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ADDIN EN.CITE.DATA Hooykaas, 2010; Barampuram and Zhang, 2011; Rivera et al., 2012; Peyret and Lomonossoff, 2015; Hwang et al., 2017; Zhao et al., 2017; Cunningham et al., 2018.Transformation methods can be divided into two major categories: indirect and direct DNA delivery. In the indirect method, the new DNA is introduced into the plant cell via bacteria, usually Agrobacterium tumefaciens or less commonly Agrobacterium rhizogenes. In the direct transformation methods, the new DNA is introduced without an intermediate host. The most commonly used direct transformation method is biolistic transformation (reviewed by ADDIN EN.CITE <EndNote><Cite AuthorYear="1"><Author>Rivera</Author><Year>2012</Year><RecNum>13</RecNum><DisplayText>Rivera et al. (2012)</DisplayText><record><rec-number>13</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1513206934">13</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rivera, A.L.</author><author>Gomez-Lim, M.</author><author>Fernandez, F.</author><author>Loske, A.M.</author></authors></contributors><titles><title>Physical methods for genetic plant transformation</title><secondary-title>Physics of Life Reviews</secondary-title></titles><periodical><full-title>Physics of Life Reviews</full-title></periodical><pages>308-345</pages><volume>9</volume><number>3</number><reprint-edition>In File</reprint-edition><keywords><keyword>Physical</keyword><keyword>methods</keyword><keyword>method</keyword><keyword>genetic</keyword><keyword>plant</keyword><keyword>plant transformation</keyword><keyword>transformation</keyword></keywords><dates><year>2012</year><pub-dates><date>2012</date></pub-dates></dates><label>19297</label><urls></urls></record></Cite></EndNote>Rivera et al. (2012)). These two transformation methods are outlined below.Agrobacterium-mediated transformation 277939515176500Agrobacterium tumefaciens is a common soil bacterium that naturally causes gall formation on a wide range of plant species, including most dicotyledonous and some monocotyledonous species ADDIN EN.CITE <EndNote><Cite><Author>Van Larebeke</Author><Year>1974</Year><RecNum>18</RecNum><DisplayText>(Van Larebeke et al., 1974)</DisplayText><record><rec-number>18</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1513206934">18</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Larebeke, N.</author><author>Engler, G.</author><author>Holsters, M.</author><author>Van den Elsacker, S.</author><author>Zaenen, I.</author><author>Schilperoort, R.A.</author><author>Schell, J.</author></authors></contributors><titles><title><style face="normal" font="default" size="100%">Large plasmid in </style><style face="italic" font="default" size="100%">Agrobacterium tumefaciens</style><style face="normal" font="default" size="100%"> essential for crown gall-inducing ability</style></title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>169-170</pages><volume>252</volume><number>5479</number><reprint-edition>In File</reprint-edition><keywords><keyword>Agrobacterium</keyword><keyword>Agrobacterium tumefaciens</keyword><keyword>AGROBACTERIUM-TUMEFACIENS</keyword><keyword>plasmid</keyword><keyword>TUMEFACIENS</keyword><keyword>Crown</keyword></keywords><dates><year>1974</year><pub-dates><date>11/8/1974 print</date></pub-dates></dates><label>9899</label><urls><related-urls><url><style face="underline" font="default" size="100%">;(Van Larebeke et al., 1974). The gall is induced by transfer of hormone-producing genes from the bacterial cell into the plant genome. The genes are carried on a circular DNA molecule found within the bacterial cell called a Tumour-inducing (Ti) plasmid. During the infection process, only a section of the Ti plasmid known as the Transfer DNA (T-DNA) is transferred to the plant. The infection and T-DNA transfer process of A.?tumefaciens has been extensively studied. This natural process has been used to facilitate genetic modification of plants. A.?tumefaciens Ti plasmids have been produced that lack the genes responsible for gall formation (disarmed plasmids; Figure 1). Genes to be inserted into the plant are put into the T-DNA section of these disarmed plasmids.A.?tumefaciens cells carrying such plasmids cannot produce a gall in an infected plant but will transfer the T-DNA sequence carrying the genes of interest into the plant cell where they stably integrate into the plant genome PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5CZXZhbjwvQXV0aG9yPjxZZWFyPjE5ODQ8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Bevan, 1984; Klee and Rogers, 1989). Although most monocotyledoneous plants are not natural hosts of A. tumefaciens ADDIN EN.CITE <EndNote><Cite><Author>De Cleene</Author><Year>1976</Year><RecNum>7</RecNum><DisplayText>(De Cleene and De Ley, 1976)</DisplayText><record><rec-number>7</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1513206934">7</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>De Cleene, M.</author><author>De Ley, J.</author></authors></contributors><titles><title>The host range of crown gall</title><secondary-title>Botanical Review</secondary-title></titles><periodical><full-title>Botanical Review</full-title></periodical><pages>389-466</pages><volume>42</volume><reprint-edition>Not in File</reprint-edition><keywords><keyword>of</keyword><keyword>Crown</keyword></keywords><dates><year>1976</year><pub-dates><date>1976</date></pub-dates></dates><label>19294</label><urls></urls></record></Cite></EndNote>(De Cleene and De Ley, 1976), recent technical developments have enlarged the range of Agrobacterium-susceptible hosts, so that they now include dicotyledonous and monocotyledonous plants ADDIN EN.CITE <EndNote><Cite><Author>Sood</Author><Year>2011</Year><RecNum>16</RecNum><DisplayText>(Sood et al., 2011; Koh et al., 2015)</DisplayText><record><rec-number>16</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1513206934">16</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Sood, P.</author><author>Bhattacharya, A.</author><author>Sood, A.</author></authors></contributors><titles><title>Problems and possibilities of monocot transformation</title><secondary-title>Biologia Plantarum</secondary-title></titles><periodical><full-title>Biologia Plantarum</full-title></periodical><pages>1-15</pages><volume>55</volume><number>1</number><reprint-edition>In File</reprint-edition><keywords><keyword>and</keyword><keyword>of</keyword><keyword>transformation</keyword></keywords><dates><year>2011</year><pub-dates><date>2011</date></pub-dates></dates><label>19293</label><urls><related-urls><url><style face="underline" font="default" size="100%">file://S:\CO\OGTR\EVAL\Eval%20Sections\Library\REFS\Agrobacterium\Sood%20et%20al%202011.pdf</style></url></related-urls></urls></record></Cite><Cite><Author>Koh</Author><Year>2015</Year><RecNum>30</RecNum><record><rec-number>30</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1525391430">30</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Koh, Hee-Jong</author><author>Kwon, Suk-Yoon</author><author>Thomson, Michael</author></authors></contributors><titles><title>Current Technologies in Plant Molecular Breeding</title></titles><dates><year>2015</year></dates><publisher>Springer</publisher><isbn>9401799962</isbn><urls></urls></record></Cite></EndNote>(Sood et al., 2011; Koh et al., 2015).Agrobacterium-mediated transformation usually results in one or a few T-DNA insertions into the plant genome. Small segments of the T-DNA flanking sequence or A. tumefaciens chromosomal sequence may also be transferred into the plant genome at a low frequency PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TbWl0aDwvQXV0aG9yPjxZZWFyPjE5OTg8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Smith, 1998; ?lker et al., 2008). The likelihood of A. tumefaciens DNA having an impact on GM plants is small because Agrobacterium chromosomal genes do not contain regulatory elements required for expression in plants, and are therefore unlikely to be expressed.Biolistic transformation (particle bombardment)In this technique, DNA is delivered into plant cells on small tungsten or gold carrier particles, approximately 2 microns in diameter. The particles are coated with the gene(s) of interest and fired into plant cells or tissues, usually using pressurised helium (Figure 2). Some of the particles penetrate the cell nucleus, where the introduced genetic material is incorporated into nuclear DNA PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TYW5mb3JkPC9BdXRob3I+PFllYXI+MTk5MDwvWWVhcj48
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ADDIN EN.CITE.DATA (Sanford, 1990). Biolistic transformation often results in multiple DNA insertions at different sites within the genome ADDIN EN.CITE <EndNote><Cite><Author>Rivera</Author><Year>2012</Year><RecNum>13</RecNum><DisplayText>(Rivera et al., 2012)</DisplayText><record><rec-number>13</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1513206934">13</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rivera, A.L.</author><author>Gomez-Lim, M.</author><author>Fernandez, F.</author><author>Loske, A.M.</author></authors></contributors><titles><title>Physical methods for genetic plant transformation</title><secondary-title>Physics of Life Reviews</secondary-title></titles><periodical><full-title>Physics of Life Reviews</full-title></periodical><pages>308-345</pages><volume>9</volume><number>3</number><reprint-edition>In File</reprint-edition><keywords><keyword>Physical</keyword><keyword>methods</keyword><keyword>method</keyword><keyword>genetic</keyword><keyword>plant</keyword><keyword>plant transformation</keyword><keyword>transformation</keyword></keywords><dates><year>2012</year><pub-dates><date>2012</date></pub-dates></dates><label>19297</label><urls></urls></record></Cite></EndNote>(Rivera et al., 2012).In recent years, biolistic transformation has become a very common method to genetically modify plants, and has been shown to be applicable to virtually all species investigated. It can also be used to deliver DNA to specific parts of plant cells, e.g. chloroplasts (Barampuram and Zhang, 2011). The process of plant transformationTransformation can be achieved using a variety of plant tissues, e.g. leaf discs, embryos and protoplasts. After the DNA delivery step, a whole plant must be regenerated from the transformed plant tissues. In most cases, plant regeneration involves a tissue culture process (Figure 2). In this process, plant tissues are transferred to a synthetic medium containing a selective agent such as an antibiotic to eliminate untransformed cells as well as nutrients and hormones to promote the growth of plants from single transformed cells. Each of the regenerated plantlets represents a GM line or event. In each GM line the new DNA fragment would have integrated randomly at a different position in the plant genome.All regenerated plants undergo a selection process that involves a phenotypic and molecular evaluation ADDIN EN.CITE <EndNote><Cite><Author>Koh</Author><Year>2015</Year><RecNum>30</RecNum><DisplayText>(Koh et al., 2015)</DisplayText><record><rec-number>30</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1525391430">30</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Koh, Hee-Jong</author><author>Kwon, Suk-Yoon</author><author>Thomson, Michael</author></authors></contributors><titles><title>Current Technologies in Plant Molecular Breeding</title></titles><dates><year>2015</year></dates><publisher>Springer</publisher><isbn>9401799962</isbn><urls></urls></record></Cite></EndNote>(Koh et al., 2015). At a phenotypic level, GM lines showing the trait of interest and without any undesired effects are selected. These selected GM lines are characterised at a molecular level, which includes determining the number of inserts they carry and their position within the genome. Selected GM lines are also usually backcrossed to their non-GM parent to eliminate any unintended effects of the transformation process (see below). Unintended effects of plant transformationThe process of plant transformation may induce other changes to the plant DNA. Unintended effects of the process of plant transformation are insertional effects and somaclonal variation.Insertional effects. The insertion of the new DNA into the genome can lead to insertions, deletions and rearrangements PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5XaWxzb248L0F1dGhvcj48WWVhcj4yMDA2PC9ZZWFyPjxS
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PgB=
ADDIN EN.CITE.DATA (Latham et al., 2006; Wilson et al., 2006; Schnell et al., 2015). Additional DNA can be inserted in the plant genome. This DNA may come from the plant genome, the inserted DNA or it can simply be random filler DNA. Small deletions in the area flanking the site of insertion are also common. These deletions are typically less than 100 bp, although larger deletions have been occasionally observed. Transformation can also result in rearrangements of both the introduced DNA and the host plant DNA. For instance, chromosomal translocations have been documented, where the flanking genomic DNA on either side of the introduced DNA mapped to two different chromosomes. The new DNA may also knock out genes when integrating into the genome or affect the expression of neighbouring genes PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXRoYW08L0F1dGhvcj48WWVhcj4yMDA2PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Latham et al., 2006; Schnell et al., 2015). These alterations to the plant DNA are collectively known as insertional effects. Somaclonal variation. The tissue culture stage can also cause DNA substitutions, insertions, deletions, rearrangements and changes in chromosome number PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXRoYW08L0F1dGhvcj48WWVhcj4yMDA2PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Latham et al., 2006; Schnell et al., 2015). This is known as somaclonal variation. These newly generated mutations arise as a result of the tissue culture process and are not related to the genetic transformation. Horticultural species that are propagated by tissue culture can also display somaclonal variation. The mutagenicity of tissue culture has been attributed to the stress the plant cells suffer during this process ADDIN EN.CITE <EndNote><Cite><Author>Krishna</Author><Year>2016</Year><RecNum>34</RecNum><DisplayText>(Krishna et al., 2016)</DisplayText><record><rec-number>34</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1526875549">34</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Krishna, Hare</author><author>Alizadeh, Mahdi</author><author>Singh, Dhurendra</author><author>Singh, Udayvir</author><author>Chauhan, Nitesh</author><author>Eftekhari, Maliheh</author><author>Sadh, Radha Kishan</author></authors></contributors><titles><title>Somaclonal variations and their applications in horticultural crops improvement</title><secondary-title>3 Biotech</secondary-title></titles><periodical><full-title>3 Biotech</full-title></periodical><pages>54</pages><volume>6</volume><number>1</number><dates><year>2016</year></dates><isbn>2190-572X</isbn><urls></urls></record></Cite></EndNote>(Krishna et al., 2016). Stressing conditions include wounding, exposure to chemicals that prevent growth of bacteria, and high concentration of plant growth regulators in the media. Different factors affect the frequency of somaclonal variation ADDIN EN.CITE <EndNote><Cite><Author>Krishna</Author><Year>2016</Year><RecNum>34</RecNum><DisplayText>(Krishna et al., 2016)</DisplayText><record><rec-number>34</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1526875549">34</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Krishna, Hare</author><author>Alizadeh, Mahdi</author><author>Singh, Dhurendra</author><author>Singh, Udayvir</author><author>Chauhan, Nitesh</author><author>Eftekhari, Maliheh</author><author>Sadh, Radha Kishan</author></authors></contributors><titles><title>Somaclonal variations and their applications in horticultural crops improvement</title><secondary-title>3 Biotech</secondary-title></titles><periodical><full-title>3 Biotech</full-title></periodical><pages>54</pages><volume>6</volume><number>1</number><dates><year>2016</year></dates><isbn>2190-572X</isbn><urls></urls></record></Cite></EndNote>(Krishna et al., 2016), including the plant species, the mode of regeneration, the length of culture period, and the culture environment. In general, the longer the period of tissue culture and the more disruptive the process, the more mutations are induced. For instance, the production of plants via axillary branching does not normally result in the production of somaclonal variants, while cultures that go through a callus phase promote a higher mutation rate ADDIN EN.CITE <EndNote><Cite><Author>Krishna</Author><Year>2016</Year><RecNum>34</RecNum><DisplayText>(Leva et al., 2012; Krishna et al., 2016)</DisplayText><record><rec-number>34</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1526875549">34</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Krishna, Hare</author><author>Alizadeh, Mahdi</author><author>Singh, Dhurendra</author><author>Singh, Udayvir</author><author>Chauhan, Nitesh</author><author>Eftekhari, Maliheh</author><author>Sadh, Radha Kishan</author></authors></contributors><titles><title>Somaclonal variations and their applications in horticultural crops improvement</title><secondary-title>3 Biotech</secondary-title></titles><periodical><full-title>3 Biotech</full-title></periodical><pages>54</pages><volume>6</volume><number>1</number><dates><year>2016</year></dates><isbn>2190-572X</isbn><urls></urls></record></Cite><Cite><Author>Leva</Author><Year>2012</Year><RecNum>35</RecNum><record><rec-number>35</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1528091184">35</key></foreign-keys><ref-type name="Book Section">5</ref-type><contributors><authors><author>Leva, AR</author><author>Petruccelli, R</author><author>Rinaldi, LMR</author></authors></contributors><titles><title>Somaclonal variation in tissue culture: a case study with olive</title><secondary-title>Recent advances in plant in vitro culture</secondary-title></titles><dates><year>2012</year></dates><publisher>InTech</publisher><urls></urls></record></Cite></EndNote>(Leva et al., 2012; Krishna et al., 2016). Somaclonal variation results in a similar spectrum of genetic variation as mutagenesis by chemical or physical agents ADDIN EN.CITE <EndNote><Cite><Author>Krishna</Author><Year>2016</Year><RecNum>34</RecNum><DisplayText>(Krishna et al., 2016)</DisplayText><record><rec-number>34</rec-number><foreign-keys><key app="EN" db-id="9eadptfx3rr2d3ew9t8vaxrkffs5eav2ppst" timestamp="1526875549">34</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Krishna, Hare</author><author>Alizadeh, Mahdi</author><author>Singh, Dhurendra</author><author>Singh, Udayvir</author><author>Chauhan, Nitesh</author><author>Eftekhari, Maliheh</author><author>Sadh, Radha Kishan</author></authors></contributors><titles><title>Somaclonal variations and their applications in horticultural crops improvement</title><secondary-title>3 Biotech</secondary-title></titles><periodical><full-title>3 Biotech</full-title></periodical><pages>54</pages><volume>6</volume><number>1</number><dates><year>2016</year></dates><isbn>2190-572X</isbn><urls></urls></record></Cite></EndNote>(Krishna et al., 2016). In fact, it is employed to develop novel cultivars in breeding programs. The extent of somaclonal variation showed by the GM plants will depend on the type of tissue culture technique used to regenerate them.Insertional effects and somaclonal variation may lead to unintended traits in the plant if they affect the expression of endogenous genes or create novel proteins ADDIN EN.CITE <EndNote><Cite><Author>Schnell</Author><Year>2015</Year><RecNum>19027</RecNum><DisplayText>(Schnell et al., 2015)</DisplayText><record><rec-number>19027</rec-number><foreign-keys><key app="EN" db-id="avrzt5sv7wwaa2epps1vzttcw5r5awswf02e" timestamp="1503881459">19027</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Schnell, J.</author><author>Steele, M.</author><author>Bean, J.</author><author>Neuspiel, M.</author><author>Girard, C.</author><author>Dormann, N.</author><author>Pearson, C.</author><author>Savoie, A.</author><author>Bourbonniere, L.</author><author>Macdonald, P.</author></authors></contributors><titles><title>A comparative analysis of insertional effects in genetically engineered plants: considerations for pre-market assessments</title><secondary-title>Transgenic Research</secondary-title></titles><periodical><full-title>Transgenic Research</full-title></periodical><pages>1-17</pages><volume>24</volume><reprint-edition>Not in File</reprint-edition><keywords><keyword>comparative analysis</keyword><keyword>analysis</keyword><keyword>of</keyword><keyword>effects</keyword><keyword>genetically</keyword><keyword>genetically engineered</keyword><keyword>genetically engineered plants</keyword><keyword>PLANTS</keyword><keyword>plant</keyword><keyword>Assessment</keyword></keywords><dates><year>2015</year><pub-dates><date>2015</date></pub-dates></dates><label>20530</label><urls></urls></record></Cite></EndNote>(Schnell et al., 2015). If unintended traits arise from changes in gene expression, these may or may not be harmful.The range of DNA changes observed in GM plants is similar to other genetic changes that occur spontaneously in plants and during conventional breeding PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Db21taXR0ZWUgb24gSWRlbnRpZnlpbmcgYW5kIEFzc2Vz
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ADDIN EN.CITE.DATA (Committee on Identifying and Assessing Unintended Effects of Genetically Engineered Foods on Human Health, 2004; Bradford et al., 2005; Weber et al., 2012; Steiner et al., 2013; Ladics et al., 2015; Schnell et al., 2015). Differences in gene expression and metabolic composition are typically greater between conventionally bred cultivars than they are between a GM plant and its non-GM parent PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TY2huZWxsPC9BdXRob3I+PFllYXI+MjAxNTwvWWVhcj48
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ADDIN EN.CITE.DATA (Baudo et al., 2006; Batista et al., 2008; Clarke et al., 2013; Schnell et al., 2015).It is important to bear in mind that new plant varieties developed by both conventional breeding and genetic modification undergo a selection process in which any undesired effects are eliminated before reaching a commercial product PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TY2huZWxsPC9BdXRob3I+PFllYXI+MjAxNTwvWWVhcj48
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ADDIN EN.CITE.DATA (Ladics et al., 2015; Schnell et al., 2015).ConclusionDespite the widespread employment of these methods of plant transformation, there have been no reports of adverse effects on human health and safety or the environment as a result of their use. Nevertheless, irrespective of which method of transformation was used, the potential for adverse unintended effects as a result of gene technology is assessed on a case-by-case basis, in the context of the proposed dealings, in each risk assessment for release of a GM plantReferences ADDIN EN.REFLIST Barampuram, S., and Zhang, Z.J. (2011). Recent advances in plant transformation. Methods in Molecular Biology 701, 1-35.Batista, R., Saibo, N., Louren?o, T., and Oliveira, M.M. (2008). Microarray analyses reveal that plant mutagenesis may induce more transcriptomic changes than transgene insertion. Proceedings of the National Academy of Sciences 105, 3640-3645.Baudo, M.M., Lyons, R., Powers, S., Pastori, G.M., Edwards, K.J., Holdsworth, M.J., and Shewry, P.R. (2006). Transgenesis has less impact on the transcriptome of wheat grain than conventional breeding. Plant Biotechnology Journal 4, 369-380.Bevan, M. (1984). Binary Agrobacterium vectors for plant transformation. Nucleic Acids Research 12, 8711-8721.Bradford, K.J., van Deynze, A., Gutterson, N., Parrott, W., and Strauss, S.H. (2005). Regulating transgenic crops sensibly: lessons from plant breeding, biotechnology and genomics. Nature Biotechnology 23, 439-444.Clarke, J.D., Alexander, D.C., Ward, D.P., Ryals, J.A., Mitchell, M.W., Wulff, J.E., and Guo, L. (2013). Assessment of Genetically Modified Soybean in Relation to Natural Variation in the Soybean Seed Metabolome. Scientific Reports 3, mittee on Identifying and Assessing Unintended Effects of Genetically Engineered Foods on Human Health, National Research Council (2004). Unintended Effects from Breeding. In Safety of Genetically Engineered Foods: Approaches to Assessing Unintended Health Effects (The National Academies Press), pp. 39-71.Cunningham, F.J., Goh, N.S., Demirer, G.S., Matos, J.L., and Landry, M.P. (2018). Nanoparticle-Mediated Delivery towards Advancing Plant Genetic Engineering. Trends in biotechnology.De Cleene, M., and De Ley, J. (1976). The host range of crown gall. Botanical Review 42, 389-466.Hooykaas, P. (2010). Plant transformation. In Encyclopedia of Life Sciences (ELS) (John Wiley and Sons, Ltd).Hwang, H.-H., Yu, M., and Lai, E.-M. (2017). Agrobacterium-Mediated Plant Transformation: Biology and Applications. The Arabidopsis Book, e0186.Klee, H.J., and Rogers, S.G. (1989). Plant gene vectors and genetic transformation: plant transformation systems based on the use of Agrobacterium tumefaciens. Cell Culture and Somatic Cell Genetics of Plants 6, 1-23.Koh, H.-J., Kwon, S.-Y., and Thomson, M. (2015). Current Technologies in Plant Molecular Breeding (Springer).Krishna, H., Alizadeh, M., Singh, D., Singh, U., Chauhan, N., Eftekhari, M., and Sadh, R.K. (2016). Somaclonal variations and their applications in horticultural crops improvement. 3 Biotech 6, 54.Ladics, G.S., Bartholomaeus, A., Bregitzer, P., Doerrer, N.G., Gray, A., Holzhauser, T., Jordan, M., et al. (2015). Genetic basis and detection of unintended effects in genetically modified crop plants. Transgenic Res 24, 587-603.Latham, J., Wilson, A.K., and Steinbrecher, R. (2006). The mutational consequences of plant transformation. Journal of Biomedicine and Biotechnology 2006, 1-7.Leva, A., Petruccelli, R., and Rinaldi, L. (2012). Somaclonal variation in tissue culture: a case study with olive. In Recent advances in plant in vitro culture (InTech).Peyret, H., and Lomonossoff, G.P. (2015). When plant virology met Agrobacterium: the rise of the deconstructed clones. Plant biotechnology journal 13, 1121-1135.Rivera, A.L., Gomez-Lim, M., Fernandez, F., and Loske, A.M. (2012). Physical methods for genetic plant transformation. Physics of Life Reviews 9, 308-345.Sanford, J.C. (1990). Biolistic plant transformation. Physiologia Plantarum 79, 206-209.Schnell, J., Steele, M., Bean, J., Neuspiel, M., Girard, C., Dormann, N., Pearson, C., et al. (2015). A comparative analysis of insertional effects in genetically engineered plants: considerations for pre-market assessments. Transgenic Research 24, 1-17.Smith, N. (1998). More T-DNA than meets the eye. Trends in Plant Science 3, 85-85.Sood, P., Bhattacharya, A., and Sood, A. (2011). Problems and possibilities of monocot transformation. Biologia Plantarum 55, 1-15.Steiner, H.Y., Halpin, C., Jez, J.M., Kough, J., Parrott, W., Underhill, L., Weber, N., et al. (2013). Evaluating the potential for adverse interactions within genetically engineered breeding stacks. Plant Physiology 161, 1587-1594.?lker, B., Li, Y., Rosso, M.G., Logemann, E., Somssich, I.E., and Weisshaar, B. (2008). T-DNA-mediated transfer of Agrobacterium tumefaciens chromosomal DNA into plants. Nature Biotechnology 26, 1015-1017.Van Larebeke, N., Engler, G., Holsters, M., Van den Elsacker, S., Zaenen, I., Schilperoort, R.A., and Schell, J. (1974). Large plasmid in Agrobacterium tumefaciens essential for crown gall-inducing ability. Nature 252, 169-170.Weber, N., Halpin, C., Hannah, L.C., Jez, J.M., Kough, J., and Parrott, W. (2012). Editor’s Choice: Crop Genome Plasticity and Its Relevance to Food and Feed Safety of Genetically Engineered Breeding Stacks. Plant Physiology 160, 1842-1853.Wilson, A.K., Latham, J.R., and Steinbrecher, R.A. (2006). Transformation-induced mutations in transgenic plants: analysis and biosafety implications. Biotechnology and Genetic Engineering Reviews 23, 209-234.Zhao, X., Meng, Z., Wang, Y., Chen, W., Sun, C., Cui, B., Cui, J., et al. (2017). Pollen magnetofection for genetic modification with magnetic nanoparticles as gene carriers. Nature Plants. ................
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