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xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink" aria-hidden="true" role="img" data-testid="keyboard_arrow_down" style="" width="16" height="16" viewBox="0 0 16 16"></svg></span></button></div></div><p class="m-text text-sm text-color-error pt-sm">This version is not peer-reviewed</p></div></div><div class="py-md border-b border-color-default"><div class="m-select common-field bg-white"><div class="common-field__wrapper" data-headlessui-state><div class="relative w-full cursor-default"><div class="flex items-center"><button id="headlessui-listbox-button-mui-22098" type="button" aria-haspopup="listbox" aria-expanded="false" data-headlessui-state data-testid="select-activator" class="common-field__dropdown-input [&>span]:overflow-hidden h-10"><span class="flex w-full items-center gap-1 [&>span]:truncate"><span>Version 1</span></span><div class="flex"><svg xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink" aria-hidden="true" role="img" data-testid="keyboard_arrow_down" class="duration-400 transition-transform" style="" width="24" height="24" viewBox="0 0 16 16"></svg></div></button></div></div></div></div><div class="pt-sm"><div class="flex"><p class="m-text text-xs">Submitted:</p><p class="m-text text-xs pl-xs">11 May 2023</p></div><div class="flex pt-sm"><p class="m-text text-xs">Posted: </p><p class="m-text text-xs pl-xs">11 May 2023</p></div></div><div class="pt-sm"><div><p class="m-text text-xs">You are already at the latest version </p></div></div><div></div></div><div class="py-md border-b border-color-default flex items-center justify-between" data-v-d8c082ee><div class="flex-none flex items-center" data-v-d8c082ee><span class="m-text text-xs pr-xs font-semibold flex-none" data-v-d8c082ee>Alerts</span><span data-v-d8c082ee></span></div><div class="m-switch flex items-center gap-2" data-v-d8c082ee><button class="relative inline-flex h-[1.4rem] w-[2.75rem] shrink-0 cursor-pointer items-center rounded-full border-2 border-color-transparent transition-colors duration-150 ease-out ui-focus-visible:ring-2 ui-focus-visible:ring-brand-bold ui-focus-visible:ring-opacity-75 bg-content-default hover:bg-content-bold active:bg-content-bolder" aria-label="Switch on" id="headlessui-switch-mui-22099" role="switch" type="button" tabindex="0" aria-checked="false" data-headlessui-state><span aria-hidden="true" class="pointer-events-none inline-block h-[1rem] w-[1rem] transform rounded-full bg-white shadow-lg ring-0 transition duration-200 ease-in-out translate-x-0.5"></span></button></div></div><div data-v-d8c082ee data-v-03f1a75d></div><div><h6 class="m-heading text-inherit m-h6 font-semibold pt-lg">Abstract</h6><div class="">Bacterial transformation is the essential tool for introducing foreign DNA into bacterial cells in a host of recombinant technology applications useful for biomanufacturing and drug production. Thus far, chemically competent cells remain an important enabling technology for bacterial transformation given its facile transformation approach. However, many parameters affect transformation efficiency, and there is relatively little documentation of their cross-interacting effect in the literature. This work explores the roles played by incubation time on ice, competent cell concentration, and plasmid-to-cell ratio on transformation efficiency in <em>Escherichia coli</em> DH5α. Results revealed that increases in incubation time improve transformation efficiency, but 2 minutes incubation still yield appreciable efficiency. On the other hand, low concentration competent cells (10<sup>6</sup> CFU/mL) yielded significant improvement in transformation efficiency compared to high concentration competent cells (10<sup>8</sup> CFU/mL). In terms of plasmid-to-cell ratio, higher ratio increases transformation efficiency. Overall, bacterial transformation is incompletely understood particularly in the area of cross-interacting parameters. Experiments reported here revealed the possibility of short duration transformation where lower competent cell concentration and higher plasmid-to-cell ratio could improve transformation.</div></div><div data-v-3ce94018><div class="pt-lg" data-v-3ce94018><span class="m-text text-body font-semibold" data-v-3ce94018>Keywords: </span><span class="m-text text-body pt-sm" data-v-3ce94018></span></div></div><div class="flex-1 pt-lg mr-lg lg:pt-0" data-v-6afb3413><div class="pt-md" data-v-6afb3413><div class="flex flex-wrap lg:flex-nowrap items-center" data-v-6afb3413><span class="m-text text-body font-semibold" data-v-6afb3413>Subject: </span><span class="m-text text-body" data-v-6afb3413>Biology and Life Sciences</span><span class="" data-v-6afb3413> - </span><span class="m-text text-body" data-v-6afb3413>Biology and Biotechnology</span></div></div><div class="content-container" id="articleRef" data-v-6afb3413><script type="text/x-mathjax-config"> MathJax.Hub.Config({ menuSettings: { CHTMLpreview: false }, "CHTML-preview":{ disabled: true }, "HTML-CSS": { scale: 90, availableFonts: [], preferredFont: null, preferredFonts: null, webFont:"Gyre-Pagella", imageFont:'TeX', undefinedFamily:"'Arial Unicode MS',serif", linebreaks: { automatic: false } }, "TeX": { extensions: ["noErrors.js"], noErrors: { inlineDelimiters: ["",""], multiLine: true, style: { "font-size": "90%", "text-align": "left", "color": "black", "padding": "1px 3px", "border": "1px solid" } } } }); </script><script type="text/javascript" async="" src="https://www.mdpi.com/bundles/mathjax/MathJax.js?config=TeX-AMS-MML_HTMLorMML"></script> <section id="Introduction" type="intro"><h2 data-nested="1" id="preprints-h2-1"> Introduction</h2> <div class="html-p">Different parameters such as incubation time, competent cell concentration, and plasmid-to-cell ratio affects transformation efficiency [<a href="#B1-preprints-73387" class="html-bibr">1</a>,<a href="#B2-preprints-73387" class="html-bibr">2</a>]. And transformation is a physio-biological process that remains incompletely understood, particularly with respect to how different parameters cross-interact to affect the final outcome: transformation efficiency. This work sought to understand through conventional microtube-based experiments how different parameters affect transformation efficiency.</div> <div class="html-p">The parameters selected for study are incubation time on ice, competent cell concentration, and plasmid-to-cell ratio. Each parameter represents some aspect of the transformation system that can significantly affect transformation efficiency, and critically, where some trade-off in system design can help improve bacterial transformation. Firstly, conventional transformation in <span class="html-italic">Escherichia coli</span> would require a 30 min incubation on ice [<a href="#B3-preprints-73387" class="html-bibr">3</a>]. This is to a significant extent too time consuming for implementation in a microfluidic chip-based system. Hence, this work sought to examine the possibility of reducing incubation time on ice. Secondly, competent cells, especially chemically competent cells, are hard and time-consuming to prepare, but there is a popular notion that high concentration competent cells would result in higher transformation efficiency. Using microtube-based experiments, a series of experiments were performed to first understand the effect of competent cell concentration on transformation efficiency, and possibly to reduce the concentration of competent cells used. Thirdly, plasmid-to-cells ratio was examined to understand if there is a less understood biological effect governing how plasmid interact with cells, especially from the perspective of single cells taking up more than one plasmid that give rise to heightened metabolic burden, reduced cell growth, and reduced colony formation. The key question here is not solely the cost of plasmid or competent cells, but rather, how do we conduct transformation to ensure high transformation efficiency as perceived through colony counting on agar plates after transformation.</div></section><section id="MaterialsandMethods" type><h2 data-nested="1" id="preprints-h2-2"> Materials and Methods</h2> <section id="CompetentCellPreparation" type><h4 class="html-italic" data-nested="2"> Competent Cell Preparation</h4> <div class="html-p"> <span class="html-italic">Escherichia coli</span> DH5α chemically competent cells were prepared as per instructions in the Zymo Research “Mix and go” competent cell preparation kit. Prepared cells were in aliquots of 200 µL per 1.5 mL microcentrifuge tube and stored at -80 <sup>o</sup>C prior to use.</div></section><section id="Plasmidusedinthisstudy" type><h4 class="html-italic" data-nested="2"> Plasmid used in this study</h4> <div class="html-p">pNAR plasmid of approximate size 10 kilobases and with ampicillin antibiotic resistance marker was used in this study. Purified plasmid was stored in elution buffer of Qiagen MiniPrep plasmid extraction kit and stored at -20 <sup>o</sup>C prior to use.</div></section><section id="Typicaltransformationprotocol" type><h4 class="html-italic" data-nested="2"> Typical transformation protocol</h4> <div class="html-p">A tube of <span class="html-italic">E. coli</span> DH5α competent cells was thawed on ice for 10 minutes prior to transformation. Typically, 100 µL of competent cells was pipetted into a sterile round bottom 14 mL Falcon two-stoppered tube which was first chilled on ice. 1 µL of pNAR plasmid was added to the competent cell mixture followed by a variable incubation period on ice. Except for comparison purpose experiments, no heat shock and outgrowth steps were used. Following incubation on ice for the cells to take up the plasmid, the transformation mixture was aliquot for spread plate culture on a LB agar plate with ampicillin antibiotic. After inoculation, agar plates were incubated at 37 <sup>o</sup>C overnight prior to counting of number of isolated colonies. </div></section></section><section id="Results" type="results"><h2 data-nested="1" id="preprints-h2-3"> Results</h2> <section id="ReducingtheTimeforTransformation" type><h4 class="html-italic" data-nested="2"> Reducing the Time for Transformation</h4> <div class="html-p">Bacterial transformation is a series of carefully choreographed steps that sought to generate cell transformed with plasmid for the next-step in biotechnology research. Given the number of plasmid variants that need to be tested in either biocatalysis or metabolic engineering research, protocols with reduced time requirement is highly sought after. Examining a typical transformation protocol such as the one depicted in <a href="#preprints-73387-f001" class="html-fig">Figure 1</a>, incubation on ice is perhaps the most time-consuming step that is suited for optimization in an effort to maximise potential trade-off between incubation time and transformation efficiency. The hypothesis here is that while incubation time should be positively correlated with transformation efficiency, a shorter incubation duration may generate sufficient colonies for single plasmid transformation.</div> <div class="html-fig-wrap" id="preprints-73387-f001"> <div class="html-fig_img"> <div class="html-figpopup html-figpopup-link" href="#fig_body_display_preprints-73387-f001"> <img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png" alt="Preprints 73387 g001" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png"> <a class="html-expand html-figpopup" href="#fig_body_display_preprints-73387-f001"></a> </div> </div> <div class="html-fig_description"> <b>Figure 1.</b> Schematic diagram showing the typical workflow for bacterial transformation that includes a lengthy incubation step of 30 minutes on ice.  </div> </div> <div class="html-fig_show mfp-hide" id="fig_body_display_preprints-73387-f001"> <div class="html-caption"> <b>Figure 1.</b> Schematic diagram showing the typical workflow for bacterial transformation that includes a lengthy incubation step of 30 minutes on ice.</div> <div class="html-img"><img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png" alt="Preprints 73387 g001" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png"></div> </div> <div class="html-fig-wrap" id="preprints-73387-f002"> <div class="html-fig_img"> <div class="html-figpopup html-figpopup-link" href="#fig_body_display_preprints-73387-f002"> <img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png" alt="Preprints 73387 g002" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png"> <a class="html-expand html-figpopup" href="#fig_body_display_preprints-73387-f002"></a> </div> </div> <div class="html-fig_description"> <b>Figure 2.</b> Effect of incubation duration on transformation efficiency showing that there is a positive correlation between incubation duration and transformation efficiency. Transformation was done without heat shock and outgrowth steps.  </div> </div> <div class="html-fig_show mfp-hide" id="fig_body_display_preprints-73387-f002"> <div class="html-caption"> <b>Figure 2.</b> Effect of incubation duration on transformation efficiency showing that there is a positive correlation between incubation duration and transformation efficiency. Transformation was done without heat shock and outgrowth steps.</div> <div class="html-img"><img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png" alt="Preprints 73387 g002" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png"></div> </div> <div class="html-p">Using chemically competent <span class="html-italic">E. coli</span> cells prepared by a commercial Zymo Research kit, experiments revealed an expected positive correlation between incubation time and transformation efficiency. Specifically, comparing 2 min and 30 min incubation on ice selects the latter as a better condition for conducting bacterial transformation in <span class="html-italic">E. coli</span>. But, do we need very high transformation efficiency? The answer is it depends. Given that 2 min incubation on ice gives an acceptable transformation efficiency of 9 x 10<sup>4</sup> CFU/µg DNA, 2 min may be sufficient for most applications in bacterial transformation.</div> <div class="html-p">With new understanding derived through experimentation, the project next compared a “Fast” transformation with a conventional transformation protocol. “Fast” transformation here refers to a 2 min incubation time on ice with no heat shock. <a href="#preprints-73387-f003" class="html-fig">Figure 3</a> illustrates that indeed, “Fast” transformation delivered less efficient transformation, but nevertheless, its performance remains sufficient for most applications.</div></section><section id="EffectofCompetentCellConcentrationonTransformationEfficiency" type><h4 class="html-italic" data-nested="2"> Effect of Competent Cell Concentration on Transformation Efficiency</h4> <div class="html-p"> <a href="#preprints-73387-f004" class="html-fig">Figure 4</a> describes the results of experiments conducted to examine the effect of competent cell concentration on transformation efficiency. Results revealed that low concentration competent cells delivered higher transformation efficiency compared to high concentration competent cells. This is a counter-intuitive result, but it can be understood in the following way. In high concentration competent cells transformation, the micro-environment surrounding a plasmid is highly negatively charged given the high negative charge on the cell membrane of many surrounding competent cells. This prevents the migration of negatively charged plasmid to the cell membrane of competent cells, and thus, reduces transformation efficiency.</div></section><section id="EffectofPlasmidtoCellRatioonTransformationEfficiency" type><h4 class="html-italic" data-nested="2"> Effect of Plasmid-to-Cell Ratio on Transformation Efficiency</h4> <div class="html-p">In conventional thinking, higher plasmid-to-cell ratio should lead to higher transformation efficiency. But, there may be cases where high plasmid-to-cell ratio lead to cells taking up more than one plasmid, which upon expression of plasmid genes, lead to high metabolic burden, reduced cell growth, and smaller and fewer colonies (<a href="#preprints-73387-f005" class="html-fig">Figure 5</a>).</div> <div class="html-p">Results from experiments conducted with pNAR in <span class="html-italic">E. coli </span>cells showed that 2x plasmid-tocell ratio delivered higher transformation efficiency compared to 1x plasmid-to-cell ratio. This indicates very likely that only one copy of pNAR plasmid was uptake by E. coli cells, thereby, allowing the rest of the plasmid molecules to be distributed to other cells, which improves transformation efficiency.</div> <div class="html-fig-wrap" id="preprints-73387-f005"> <div class="html-fig_img"> <div class="html-figpopup html-figpopup-link" href="#fig_body_display_preprints-73387-f005"> <img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png" alt="Preprints 73387 g005" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png"> <a class="html-expand html-figpopup" href="#fig_body_display_preprints-73387-f005"></a> </div> </div> <div class="html-fig_description"> <b>Figure 5.</b> Schematic diagram illustrating the possible phenomenon of competent cells taking up more than 1 plasmid in a transformation mixture of high plasmid-to-cell ratio.  </div> </div> <div class="html-fig_show mfp-hide" id="fig_body_display_preprints-73387-f005"> <div class="html-caption"> <b>Figure 5.</b> Schematic diagram illustrating the possible phenomenon of competent cells taking up more than 1 plasmid in a transformation mixture of high plasmid-to-cell ratio.</div> <div class="html-img"><img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png" alt="Preprints 73387 g005" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png"></div> </div> <div class="html-fig-wrap" id="preprints-73387-f006"> <div class="html-fig_img"> <div class="html-figpopup html-figpopup-link" href="#fig_body_display_preprints-73387-f006"> <img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png" alt="Preprints 73387 g006" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png"> <a class="html-expand html-figpopup" href="#fig_body_display_preprints-73387-f006"></a> </div> </div> <div class="html-fig_description"> <b>Figure 6.</b> Effect of plasmid-to-cell ratio on transformation efficiency for pNAR plasmid in <span class="html-italic">E. coli</span> cells. 1x plasmid conc refers to 1 µl plasmid to 100 µl competent cell, 2x plasmid conc refers to 1 µl plasmid to 50 µl competent cell.  </div> </div> <div class="html-fig_show mfp-hide" id="fig_body_display_preprints-73387-f006"> <div class="html-caption"> <b>Figure 6.</b> Effect of plasmid-to-cell ratio on transformation efficiency for pNAR plasmid in <span class="html-italic">E. coli</span> cells. 1x plasmid conc refers to 1 µl plasmid to 100 µl competent cell, 2x plasmid conc refers to 1 µl plasmid to 50 µl competent cell.</div> <div class="html-img"><img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png" alt="Preprints 73387 g006" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png"></div> </div> </section></section><section id="Discussion" type="discussion"><h2 data-nested="1" id="preprints-h2-4"> Discussion</h2> <div class="html-p">From the theoretical perspective, bacterial transformation is the process whereby we manipulate experimental conditions to facilitate the movement of a plasmid close to a competent cell, and helping the cell to uptake the plasmid to endow it with new functions and properties. Along the way, many factors need to be considered to help optimise the journey and improve the efficiency of bacterial transformation. In terms of expediency, incubation time on ice is perhaps one area in need for improvement.</div> <div class="html-p">Current protocols for <span class="html-italic">Escherichia coli</span> transformation require 30 minutes incubation on ice to ensure higher transformation efficiency [<a href="#B4-preprints-73387" class="html-bibr">4</a>]. But, could we achieve similar transformation efficiency with a shorter time? This work demonstrates that incubation time positively correlates with transformation efficiency since more time is available for the plasmid to get close to the cells and be taken up by it. However, a short 2 minute incubation still yield good transformation efficiency suitable for most applications in biotechnology where the desire is to transformation the cell with one plasmid.</div> <div class="html-p">Perhaps the most intriguing finding from this work is the realisation that high concentration competent cells (10<sup>8</sup> CFU/mL) delivered poorer transformation efficiency compared to low concentration competent cells (10<sup>6</sup> CFU/mL). This is indeed counter-intuitive since conventional notion is that higher concentration of competent cells preparation will have more cells to uptake a given amount of plasmid. However, experiment results indicated that this is not true. One reason could be high concentration competent cells presenting a highly negatively charged microenvironment where the plasmid is surrounded by many negatively charged competent cells. Charge-charge repulsion would then present a significant hindrance for the facile diffusion of the plasmid to the cell surface, and subsequent uptake by the cell. This aspect of transformation is perhaps the least suspecting facet of the process, and its elucidation may help optimise many transformation protocols used in academic and industrial research labs.</div> <div class="html-p">Finally, plasmid-to-cell ratio concerns the usage and consumption of precious plasmid, which is still produced by the cell-based approach coupled with plasmid extraction [<a href="#B5-preprints-73387" class="html-bibr">5</a>]. Higher plasmid-to-cell ratio is perhaps the natural choice for many researchers interested in ensuring success of the experiment. However, possibility exists of high plasmid-to-cell ratio resulting in too many plasmids taken up by the cells, which may exhibit retarded growth due to high metabolic burden of expressing genes on the plasmid [<a href="#B6-preprints-73387" class="html-bibr">6</a>]. Coupled with strong replication origins on the plasmid, a high plasmid-to-cell ratio may lead to smaller and fewer colonies on the transformation plate. This work observes that a one-fold increase in plasmid-to-cell ratio improves transformation efficiency. The results suggests that the pNAR plasmid may be too large (10 kB in size) for efficient uptake by the cells. Plasmid smaller in size, and thus, more efficiently uptake by cells may result in a reverse correlation. Hence, plasmid size likely determines the effect of plasmid-to-cell ratio on transformation efficiency.</div></section><section id="Conclusions" type="conclusions"><h2 data-nested="1" id="preprints-h2-5"> Conclusions</h2> <div class="html-p">This work has elucidated new understanding in three areas of bacterial transformation. Firstly, possibility exists in reducing the incubation time on ice while still retaining appreciable level of transformation efficiency for most applications. Secondly, low concentration competent cells may deliver better transformation efficiency compared to high concentration competent cells. Finally, experiments should be conducted to examine species-specific effect of plasmid-to-cell ratio. In the case of <span class="html-italic">E. coli</span>, only one plasmid is taken up by each cell in the case of large plasmid like pNAR, and hence, higher plasmid-to-cell ratio leads to higher transformation efficiency in these cases.</div></section> <section class="html-notes"><h2 id="preprints-h2-6">Funding</h2> <div class="html-p">The author thank the National University of Singapore for financial support.</div></section><section class="html-notes"><h2 id="preprints-h2-7">Conflicts of Interest</h2> <div class="html-p">The authors declare no conflicts of interest.</div></section><section id="html-references_list"><h2 id="preprints-h2-8">References</h2> <ol class="html-x"> <li id="B1-preprints-73387" class="html-x" data-content="1.">H. Tan, L. Fu, and M. Seno, “Optimization of Bacterial Plasmid Transformation Using Nanomaterials Based on the Yoshida Effect. <span class="html-italic">Int. J. Mol. Sci.</span> <b>2010</b>, <span class="html-italic">11</span>, 12. 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On the other hand, conventional transformation refers to the typical workflow outlined in <a href="#preprints-73387-f001" class="html-fig">Figure 1</a> with 30 min incubation on ice and with heat shock and outgrowth.  </div> </div> <div class="html-fig_show mfp-hide" id="fig_body_display_preprints-73387-f003"> <div class="html-caption"> <b>Figure 3.</b> Comparison between “Fast” transformation and conventional transformation. “Fast” transformation refers to transformation on ice with 2 min incubation and no heat shock. On the other hand, conventional transformation refers to the typical workflow outlined in <a href="#preprints-73387-f001" class="html-fig">Figure 1</a> with 30 min incubation on ice and with heat shock and outgrowth.</div> <div class="html-img"><img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png" alt="Preprints 73387 g003" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png"></div> </div> <div class="html-fig-wrap" id="preprints-73387-f004"> <div class="html-fig_img"> <div class="html-figpopup html-figpopup-link" href="#fig_body_display_preprints-73387-f004"> <img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png" alt="Preprints 73387 g004" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png"> <a class="html-expand html-figpopup" href="#fig_body_display_preprints-73387-f004"></a> </div> </div> <div class="html-fig_description"> <b>Figure 4.</b> Effect of competent cell concentration on transformation efficiency. Experiments with pNAR plasmid in <span class="html-italic">Escherichia coli</span> DH5α indicated that low concentration competent cells (1.45 x 10<sup>6</sup> CFU/ml) delivered higher transformation efficiency compared to high concentration competent cells (8.5 x 10<sup>8</sup> CFU/ml).  </div> </div> <div class="html-fig_show mfp-hide" id="fig_body_display_preprints-73387-f004"> <div class="html-caption"> <b>Figure 4.</b> Effect of competent cell concentration on transformation efficiency. Experiments with pNAR plasmid in <span class="html-italic">Escherichia coli</span> DH5α indicated that low concentration competent cells (1.45 x 10<sup>6</sup> CFU/ml) delivered higher transformation efficiency compared to high concentration competent cells (8.5 x 10<sup>8</sup> CFU/ml).</div> <div class="html-img"><img data-large="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png" data-original="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png" alt="Preprints 73387 g004" src="https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png"></div> </div> </section><section class="html-fn_group"><table><tr id> <td></td> <td><div class="html-p"> <b>Disclaimer/Publisher’s Note:</b> The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). 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Stay informed on research that impacts our environment and sustainability.","environmental-and-earth-sciences",{"id":96,"name":97,"name_system":98,"parent_id":50,"image_banner":99,"description":100,"converted_name_system":101},130,"Medicine and Pharmacology","medicine_and_pharmacology","https://www.preprints.org/media/cache/resolve/webp/upload/subject/2024-11-07/98c86d1fdf40abfda51ca959f75f3499.jpg","Embark on a journey through the vast world of medicine and pharmacology with our preprints. Explore breakthroughs in healthcare, from cardiology to neurology, which improve human health outcomes. Delve into the frontiers of medical research and pharmaceutical development that enhance quality of life.","medicine-and-pharmacology",{"id":103,"name":104,"name_system":105,"parent_id":50,"image_banner":106,"description":107,"converted_name_system":108},166,"Physical Sciences","physical_sciences","https://www.preprints.org/media/cache/resolve/webp/upload/subject/2024-11-07/c2d6e400b105b8c3d32ec59f26ab969d.jpg","Explore the fundamental laws governing matter and energy in the universe with our preprints. From quantum mechanics to cosmology, stay abreast of research that advances our understanding of the natural world. Dive into a world of scientific discovery that pushes the boundaries of knowledge.","physical-sciences",{"id":110,"name":111,"name_system":112,"parent_id":50,"image_banner":113,"description":114,"converted_name_system":115},185,"Public Health and Healthcare","public_health_and_healthcare","https://www.preprints.org/media/cache/resolve/webp/upload/subject/2024-11-07/ef7c6de55fa117659c3b6090edc7c4ff.jpg","Examine critical issues in public health policy, epidemiology, and healthcare systems with our preprints. Stay updated on research into health disparities, infectious disease control, and healthcare management, which informs public health interventions. Explore the latest trends in healthcare that impact population well-being and equity.","public-health-and-healthcare",{"id":117,"name":118,"name_system":119,"parent_id":50,"image_banner":120,"description":121,"converted_name_system":122},193,"Social Sciences","social_sciences","https://www.preprints.org/media/cache/resolve/webp/upload/subject/2024-09-12/910c960e3a6808861523728f0f26b2df.png","Gain insights into human behavior, societies, and cultures with our preprints. Explore disciplines like psychology, sociology, and political science, which shape our world and inform public policy. Stay engaged with research that explores social issues, cultural dynamics, and economic systems, enriching our understanding of human interactions.","social-sciences",{"code":41,"msg":42,"data":124},{"temp_id":125,"version":7,"id":126,"hash_key":127,"doi":128,"article_abstract":129,"article_title":130,"keywords":131,"is_registering_doi":132,"mdpi_topic":50,"preprints_collections":133,"subject":134,"top_subject":137,"article_type":138,"submitted_at":141,"published_at":142,"last_edited_at":50,"authors":143,"ethical_approval":132,"ethical_approval_number":50,"ethical_approval_body":50,"ethical_approval_for_publication":132,"article_supplementary":50,"final_file":152,"graphic_abstract":156,"statistics":157,"is_peer_reviewed":132,"peer_reviewed_article_url":50,"almetric":50,"preserved_by_portico":6,"ms_xml":160,"versions":162,"citation":164,"peer_reviewed_citation":50,"version_changes":165,"updating_alert_registered":132,"preprints_process_url":50,"comments_count":41,"latest_version":7,"author_notes":166,"withdrawed_at":50,"is_withdrwan":132},73387,"202305.0849","869f7a4b8e128c6dfc43eaa2e5a2a0ac","10.20944/preprints202305.0849.v1","Bacterial transformation is the essential tool for introducing foreign DNA into bacterial cells in a host of recombinant technology applications useful for biomanufacturing and drug production. Thus far, chemically competent cells remain an important enabling technology for bacterial transformation given its facile transformation approach. However, many parameters affect transformation efficiency, and there is relatively little documentation of their cross-interacting effect in the literature. This work explores the roles played by incubation time on ice, competent cell concentration, and plasmid-to-cell ratio on transformation efficiency in \u003Cem>Escherichia coli\u003C/em> DH5α. Results revealed that increases in incubation time improve transformation efficiency, but 2 minutes incubation still yield appreciable efficiency. On the other hand, low concentration competent cells (10\u003Csup>6\u003C/sup> CFU/mL) yielded significant improvement in transformation efficiency compared to high concentration competent cells (10\u003Csup>8\u003C/sup> CFU/mL). In terms of plasmid-to-cell ratio, higher ratio increases transformation efficiency. Overall, bacterial transformation is incompletely understood particularly in the area of cross-interacting parameters. Experiments reported here revealed the possibility of short duration transformation where lower competent cell concentration and higher plasmid-to-cell ratio could improve transformation.","Understanding the Different Parameters That Affects Bacterial Transformation Efficiency","Escherichia coli; bacterial transformation; plasmid-to-cell ratio; competent cell concentration; incubation time",false,[],{"id":135,"name":136},24,"Biology and Biotechnology",{"id":55,"name":56},{"id":139,"name":140},15,"Article","2023-05-11 04:48:12","2023-05-11 11:49:13",[144],{"id":145,"name":146,"email":147,"is_corresponding":6,"orcid_link":148,"author_mark":149,"sp_link":150,"avatar":151},338295,"Wenfa Ng","ngwenfa771@hotmail.com","https://orcid.org/0000-0001-7629-2140","*","https://sciprofiles.com/profile/954036","/statics/img/design/default-user.png",{"filename":153,"url":154,"filesize":155},"final_file.pdf","/frontend/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/download_pub",614913,[],{"viewed":158,"downloaded":159},"174","626",{"html_content":161},"\u003Cscript type=\"text/x-mathjax-config\">\n MathJax.Hub.Config({\n menuSettings: {\n CHTMLpreview: false\n },\n \"CHTML-preview\":{\n disabled: true\n },\n \"HTML-CSS\": {\n scale: 90,\n availableFonts: [],\n preferredFont: null,\n preferredFonts: null,\n webFont:\"Gyre-Pagella\",\n imageFont:'TeX',\n undefinedFamily:\"'Arial Unicode MS',serif\",\n linebreaks: { automatic: false }\n },\n \"TeX\": {\n extensions: [\"noErrors.js\"],\n noErrors: {\n inlineDelimiters: [\"\",\"\"],\n multiLine: true,\n style: {\n \"font-size\": \"90%\",\n \"text-align\": \"left\",\n \"color\": \"black\",\n \"padding\": \"1px 3px\",\n \"border\": \"1px solid\"\n }\n }\n }\n });\n \u003C/script>\u003Cscript type=\"text/javascript\" async=\"\" src=\"https://www.mdpi.com/bundles/mathjax/MathJax.js?config=TeX-AMS-MML_HTMLorMML\">\u003C/script>\n \u003Csection id=\"Introduction\" type=\"intro\">\u003Ch2 data-nested=\"1\" id=\"preprints-h2-1\"> Introduction\u003C/h2>\n\u003Cdiv class=\"html-p\">Different parameters such as incubation time, competent cell concentration, and plasmid-to-cell ratio affects transformation efficiency [\u003Ca href=\"#B1-preprints-73387\" class=\"html-bibr\">1\u003C/a>,\u003Ca href=\"#B2-preprints-73387\" class=\"html-bibr\">2\u003C/a>]. And transformation is a physio-biological process that remains incompletely understood, particularly with respect to how different parameters cross-interact to affect the final outcome: transformation efficiency. This work sought to understand through conventional microtube-based experiments how different parameters affect transformation efficiency.\u003C/div>\n\u003Cdiv class=\"html-p\">The parameters selected for study are incubation time on ice, competent cell concentration, and plasmid-to-cell ratio. Each parameter represents some aspect of the transformation system that can significantly affect transformation efficiency, and critically, where some trade-off in system design can help improve bacterial transformation. Firstly, conventional transformation in \u003Cspan class=\"html-italic\">Escherichia coli\u003C/span> would require a 30 min incubation on ice [\u003Ca href=\"#B3-preprints-73387\" class=\"html-bibr\">3\u003C/a>]. This is to a significant extent too time consuming for implementation in a microfluidic chip-based system. Hence, this work sought to examine the possibility of reducing incubation time on ice. Secondly, competent cells, especially chemically competent cells, are hard and time-consuming to prepare, but there is a popular notion that high concentration competent cells would result in higher transformation efficiency. Using microtube-based experiments, a series of experiments were performed to first understand the effect of competent cell concentration on transformation efficiency, and possibly to reduce the concentration of competent cells used. Thirdly, plasmid-to-cells ratio was examined to understand if there is a less understood biological effect governing how plasmid interact with cells, especially from the perspective of single cells taking up more than one plasmid that give rise to heightened metabolic burden, reduced cell growth, and reduced colony formation. The key question here is not solely the cost of plasmid or competent cells, but rather, how do we conduct transformation to ensure high transformation efficiency as perceived through colony counting on agar plates after transformation.\u003C/div>\u003C/section>\u003Csection id=\"MaterialsandMethods\" type>\u003Ch2 data-nested=\"1\" id=\"preprints-h2-2\"> Materials and Methods\u003C/h2>\n\u003Csection id=\"CompetentCellPreparation\" type>\u003Ch4 class=\"html-italic\" data-nested=\"2\"> Competent Cell Preparation\u003C/h4>\n\u003Cdiv class=\"html-p\">\n\u003Cspan class=\"html-italic\">Escherichia coli\u003C/span> DH5α chemically competent cells were prepared as per instructions in the Zymo Research “Mix and go” competent cell preparation kit. Prepared cells were in aliquots of 200 µL per 1.5 mL microcentrifuge tube and stored at -80 \u003Csup>o\u003C/sup>C prior to use.\u003C/div>\u003C/section>\u003Csection id=\"Plasmidusedinthisstudy\" type>\u003Ch4 class=\"html-italic\" data-nested=\"2\"> Plasmid used in this study\u003C/h4>\n\u003Cdiv class=\"html-p\">pNAR plasmid of approximate size 10 kilobases and with ampicillin antibiotic resistance marker was used in this study. Purified plasmid was stored in elution buffer of Qiagen MiniPrep plasmid extraction kit and stored at -20 \u003Csup>o\u003C/sup>C prior to use.\u003C/div>\u003C/section>\u003Csection id=\"Typicaltransformationprotocol\" type>\u003Ch4 class=\"html-italic\" data-nested=\"2\"> Typical transformation protocol\u003C/h4>\n\u003Cdiv class=\"html-p\">A tube of \u003Cspan class=\"html-italic\">E. coli\u003C/span> DH5α competent cells was thawed on ice for 10 minutes prior to transformation. Typically, 100 µL of competent cells was pipetted into a sterile round bottom 14 mL Falcon two-stoppered tube which was first chilled on ice. 1 µL of pNAR plasmid was added to the competent cell mixture followed by a variable incubation period on ice. Except for comparison purpose experiments, no heat shock and outgrowth steps were used. Following incubation on ice for the cells to take up the plasmid, the transformation mixture was aliquot for spread plate culture on a LB agar plate with ampicillin antibiotic. After inoculation, agar plates were incubated at 37 \u003Csup>o\u003C/sup>C overnight prior to counting of number of isolated colonies. \u003C/div>\u003C/section>\u003C/section>\u003Csection id=\"Results\" type=\"results\">\u003Ch2 data-nested=\"1\" id=\"preprints-h2-3\"> Results\u003C/h2>\n\u003Csection id=\"ReducingtheTimeforTransformation\" type>\u003Ch4 class=\"html-italic\" data-nested=\"2\"> Reducing the Time for Transformation\u003C/h4>\n\u003Cdiv class=\"html-p\">Bacterial transformation is a series of carefully choreographed steps that sought to generate cell transformed with plasmid for the next-step in biotechnology research. Given the number of plasmid variants that need to be tested in either biocatalysis or metabolic engineering research, protocols with reduced time requirement is highly sought after. Examining a typical transformation protocol such as the one depicted in \u003Ca href=\"#preprints-73387-f001\" class=\"html-fig\">Figure 1\u003C/a>, incubation on ice is perhaps the most time-consuming step that is suited for optimization in an effort to maximise potential trade-off between incubation time and transformation efficiency. The hypothesis here is that while incubation time should be positively correlated with transformation efficiency, a shorter incubation duration may generate sufficient colonies for single plasmid transformation.\u003C/div>\n\u003Cdiv class=\"html-fig-wrap\" id=\"preprints-73387-f001\">\n \u003Cdiv class=\"html-fig_img\">\n \u003Cdiv class=\"html-figpopup html-figpopup-link\" href=\"#fig_body_display_preprints-73387-f001\">\n \u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png\" alt=\"Preprints 73387 g001\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png\">\n \u003Ca class=\"html-expand html-figpopup\" href=\"#fig_body_display_preprints-73387-f001\">\u003C/a>\n \u003C/div>\n\n \u003C/div>\n \u003Cdiv class=\"html-fig_description\">\n \u003Cb>Figure 1.\u003C/b>\n Schematic diagram showing the typical workflow for bacterial transformation that includes a lengthy incubation step of 30 minutes on ice.\n\u003C!-- \u003Cp>\u003Ca class=\"html-figpopup\" href=\"#fig_body_display_preprints-73387-f001\">\n Click here to enlarge figure\n \u003C/a>\u003C/p> -->\n\n \u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig_show mfp-hide\" id=\"fig_body_display_preprints-73387-f001\">\n \u003Cdiv class=\"html-caption\"> \u003Cb>Figure 1.\u003C/b>\n Schematic diagram showing the typical workflow for bacterial transformation that includes a lengthy incubation step of 30 minutes on ice.\u003C/div>\n \u003Cdiv class=\"html-img\">\u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png\" alt=\"Preprints 73387 g001\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g001.png\">\u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig-wrap\" id=\"preprints-73387-f002\">\n \u003Cdiv class=\"html-fig_img\">\n \u003Cdiv class=\"html-figpopup html-figpopup-link\" href=\"#fig_body_display_preprints-73387-f002\">\n \u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png\" alt=\"Preprints 73387 g002\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png\">\n \u003Ca class=\"html-expand html-figpopup\" href=\"#fig_body_display_preprints-73387-f002\">\u003C/a>\n \u003C/div>\n\n \u003C/div>\n \u003Cdiv class=\"html-fig_description\">\n \u003Cb>Figure 2.\u003C/b>\n Effect of incubation duration on transformation efficiency showing that there is a positive correlation between incubation duration and transformation efficiency. Transformation was done without heat shock and outgrowth steps.\n\u003C!-- \u003Cp>\u003Ca class=\"html-figpopup\" href=\"#fig_body_display_preprints-73387-f002\">\n Click here to enlarge figure\n \u003C/a>\u003C/p> -->\n\n \u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig_show mfp-hide\" id=\"fig_body_display_preprints-73387-f002\">\n \u003Cdiv class=\"html-caption\"> \u003Cb>Figure 2.\u003C/b>\n Effect of incubation duration on transformation efficiency showing that there is a positive correlation between incubation duration and transformation efficiency. Transformation was done without heat shock and outgrowth steps.\u003C/div>\n \u003Cdiv class=\"html-img\">\u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png\" alt=\"Preprints 73387 g002\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g002.png\">\u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-p\">Using chemically competent \u003Cspan class=\"html-italic\">E. coli\u003C/span> cells prepared by a commercial Zymo Research kit, experiments revealed an expected positive correlation between incubation time and transformation efficiency. Specifically, comparing 2 min and 30 min incubation on ice selects the latter as a better condition for conducting bacterial transformation in \u003Cspan class=\"html-italic\">E. coli\u003C/span>. But, do we need very high transformation efficiency? The answer is it depends. Given that 2 min incubation on ice gives an acceptable transformation efficiency of 9 x 10\u003Csup>4\u003C/sup> CFU/µg DNA, 2 min may be sufficient for most applications in bacterial transformation.\u003C/div>\n\u003Cdiv class=\"html-p\">With new understanding derived through experimentation, the project next compared a “Fast” transformation with a conventional transformation protocol. “Fast” transformation here refers to a 2 min incubation time on ice with no heat shock. \u003Ca href=\"#preprints-73387-f003\" class=\"html-fig\">Figure 3\u003C/a> illustrates that indeed, “Fast” transformation delivered less efficient transformation, but nevertheless, its performance remains sufficient for most applications.\u003C/div>\u003C/section>\u003Csection id=\"EffectofCompetentCellConcentrationonTransformationEfficiency\" type>\u003Ch4 class=\"html-italic\" data-nested=\"2\"> Effect of Competent Cell Concentration on Transformation Efficiency\u003C/h4>\n\u003Cdiv class=\"html-p\">\n\u003Ca href=\"#preprints-73387-f004\" class=\"html-fig\">Figure 4\u003C/a> describes the results of experiments conducted to examine the effect of competent cell concentration on transformation efficiency. Results revealed that low concentration competent cells delivered higher transformation efficiency compared to high concentration competent cells. This is a counter-intuitive result, but it can be understood in the following way. In high concentration competent cells transformation, the micro-environment surrounding a plasmid is highly negatively charged given the high negative charge on the cell membrane of many surrounding competent cells. This prevents the migration of negatively charged plasmid to the cell membrane of competent cells, and thus, reduces transformation efficiency.\u003C/div>\u003C/section>\u003Csection id=\"EffectofPlasmidtoCellRatioonTransformationEfficiency\" type>\u003Ch4 class=\"html-italic\" data-nested=\"2\"> Effect of Plasmid-to-Cell Ratio on Transformation Efficiency\u003C/h4>\n\u003Cdiv class=\"html-p\">In conventional thinking, higher plasmid-to-cell ratio should lead to higher transformation efficiency. But, there may be cases where high plasmid-to-cell ratio lead to cells taking up more than one plasmid, which upon expression of plasmid genes, lead to high metabolic burden, reduced cell growth, and smaller and fewer colonies (\u003Ca href=\"#preprints-73387-f005\" class=\"html-fig\">Figure 5\u003C/a>).\u003C/div>\n\u003Cdiv class=\"html-p\">Results from experiments conducted with pNAR in \u003Cspan class=\"html-italic\">E. coli \u003C/span>cells showed that 2x plasmid-tocell ratio delivered higher transformation efficiency compared to 1x plasmid-to-cell ratio. This indicates very likely that only one copy of pNAR plasmid was uptake by E. coli cells, thereby, allowing the rest of the plasmid molecules to be distributed to other cells, which improves transformation efficiency.\u003C/div>\n\u003Cdiv class=\"html-fig-wrap\" id=\"preprints-73387-f005\">\n \u003Cdiv class=\"html-fig_img\">\n \u003Cdiv class=\"html-figpopup html-figpopup-link\" href=\"#fig_body_display_preprints-73387-f005\">\n \u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png\" alt=\"Preprints 73387 g005\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png\">\n \u003Ca class=\"html-expand html-figpopup\" href=\"#fig_body_display_preprints-73387-f005\">\u003C/a>\n \u003C/div>\n\n \u003C/div>\n \u003Cdiv class=\"html-fig_description\">\n \u003Cb>Figure 5.\u003C/b>\n Schematic diagram illustrating the possible phenomenon of competent cells taking up more than 1 plasmid in a transformation mixture of high plasmid-to-cell ratio.\n\u003C!-- \u003Cp>\u003Ca class=\"html-figpopup\" href=\"#fig_body_display_preprints-73387-f005\">\n Click here to enlarge figure\n \u003C/a>\u003C/p> -->\n\n \u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig_show mfp-hide\" id=\"fig_body_display_preprints-73387-f005\">\n \u003Cdiv class=\"html-caption\"> \u003Cb>Figure 5.\u003C/b>\n Schematic diagram illustrating the possible phenomenon of competent cells taking up more than 1 plasmid in a transformation mixture of high plasmid-to-cell ratio.\u003C/div>\n \u003Cdiv class=\"html-img\">\u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png\" alt=\"Preprints 73387 g005\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g005.png\">\u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig-wrap\" id=\"preprints-73387-f006\">\n \u003Cdiv class=\"html-fig_img\">\n \u003Cdiv class=\"html-figpopup html-figpopup-link\" href=\"#fig_body_display_preprints-73387-f006\">\n \u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png\" alt=\"Preprints 73387 g006\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png\">\n \u003Ca class=\"html-expand html-figpopup\" href=\"#fig_body_display_preprints-73387-f006\">\u003C/a>\n \u003C/div>\n\n \u003C/div>\n \u003Cdiv class=\"html-fig_description\">\n \u003Cb>Figure 6.\u003C/b>\n Effect of plasmid-to-cell ratio on transformation efficiency for pNAR plasmid in \u003Cspan class=\"html-italic\">E. coli\u003C/span> cells. 1x plasmid conc refers to 1 µl plasmid to 100 µl competent cell, 2x plasmid conc refers to 1 µl plasmid to 50 µl competent cell.\n\u003C!-- \u003Cp>\u003Ca class=\"html-figpopup\" href=\"#fig_body_display_preprints-73387-f006\">\n Click here to enlarge figure\n \u003C/a>\u003C/p> -->\n\n \u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig_show mfp-hide\" id=\"fig_body_display_preprints-73387-f006\">\n \u003Cdiv class=\"html-caption\"> \u003Cb>Figure 6.\u003C/b>\n Effect of plasmid-to-cell ratio on transformation efficiency for pNAR plasmid in \u003Cspan class=\"html-italic\">E. coli\u003C/span> cells. 1x plasmid conc refers to 1 µl plasmid to 100 µl competent cell, 2x plasmid conc refers to 1 µl plasmid to 50 µl competent cell.\u003C/div>\n \u003Cdiv class=\"html-img\">\u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png\" alt=\"Preprints 73387 g006\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g006.png\">\u003C/div>\n\u003C/div>\n\u003C/section>\u003C/section>\u003Csection id=\"Discussion\" type=\"discussion\">\u003Ch2 data-nested=\"1\" id=\"preprints-h2-4\"> Discussion\u003C/h2>\n\u003Cdiv class=\"html-p\">From the theoretical perspective, bacterial transformation is the process whereby we manipulate experimental conditions to facilitate the movement of a plasmid close to a competent cell, and helping the cell to uptake the plasmid to endow it with new functions and properties. Along the way, many factors need to be considered to help optimise the journey and improve the efficiency of bacterial transformation. In terms of expediency, incubation time on ice is perhaps one area in need for improvement.\u003C/div>\n\u003Cdiv class=\"html-p\">Current protocols for \u003Cspan class=\"html-italic\">Escherichia coli\u003C/span> transformation require 30 minutes incubation on ice to ensure higher transformation efficiency [\u003Ca href=\"#B4-preprints-73387\" class=\"html-bibr\">4\u003C/a>]. But, could we achieve similar transformation efficiency with a shorter time? This work demonstrates that incubation time positively correlates with transformation efficiency since more time is available for the plasmid to get close to the cells and be taken up by it. However, a short 2 minute incubation still yield good transformation efficiency suitable for most applications in biotechnology where the desire is to transformation the cell with one plasmid.\u003C/div>\n\u003Cdiv class=\"html-p\">Perhaps the most intriguing finding from this work is the realisation that high concentration competent cells (10\u003Csup>8\u003C/sup> CFU/mL) delivered poorer transformation efficiency compared to low concentration competent cells (10\u003Csup>6\u003C/sup> CFU/mL). This is indeed counter-intuitive since conventional notion is that higher concentration of competent cells preparation will have more cells to uptake a given amount of plasmid. However, experiment results indicated that this is not true. One reason could be high concentration competent cells presenting a highly negatively charged microenvironment where the plasmid is surrounded by many negatively charged competent cells. Charge-charge repulsion would then present a significant hindrance for the facile diffusion of the plasmid to the cell surface, and subsequent uptake by the cell. This aspect of transformation is perhaps the least suspecting facet of the process, and its elucidation may help optimise many transformation protocols used in academic and industrial research labs.\u003C/div>\n\u003Cdiv class=\"html-p\">Finally, plasmid-to-cell ratio concerns the usage and consumption of precious plasmid, which is still produced by the cell-based approach coupled with plasmid extraction [\u003Ca href=\"#B5-preprints-73387\" class=\"html-bibr\">5\u003C/a>]. Higher plasmid-to-cell ratio is perhaps the natural choice for many researchers interested in ensuring success of the experiment. However, possibility exists of high plasmid-to-cell ratio resulting in too many plasmids taken up by the cells, which may exhibit retarded growth due to high metabolic burden of expressing genes on the plasmid [\u003Ca href=\"#B6-preprints-73387\" class=\"html-bibr\">6\u003C/a>]. Coupled with strong replication origins on the plasmid, a high plasmid-to-cell ratio may lead to smaller and fewer colonies on the transformation plate. This work observes that a one-fold increase in plasmid-to-cell ratio improves transformation efficiency. The results suggests that the pNAR plasmid may be too large (10 kB in size) for efficient uptake by the cells. Plasmid smaller in size, and thus, more efficiently uptake by cells may result in a reverse correlation. Hence, plasmid size likely determines the effect of plasmid-to-cell ratio on transformation efficiency.\u003C/div>\u003C/section>\u003Csection id=\"Conclusions\" type=\"conclusions\">\u003Ch2 data-nested=\"1\" id=\"preprints-h2-5\"> Conclusions\u003C/h2>\n\u003Cdiv class=\"html-p\">This work has elucidated new understanding in three areas of bacterial transformation. Firstly, possibility exists in reducing the incubation time on ice while still retaining appreciable level of transformation efficiency for most applications. Secondly, low concentration competent cells may deliver better transformation efficiency compared to high concentration competent cells. Finally, experiments should be conducted to examine species-specific effect of plasmid-to-cell ratio. In the case of \u003Cspan class=\"html-italic\">E. coli\u003C/span>, only one plasmid is taken up by each cell in the case of large plasmid like pNAR, and hence, higher plasmid-to-cell ratio leads to higher transformation efficiency in these cases.\u003C/div>\u003C/section>\n \n \u003Csection class=\"html-notes\">\u003Ch2 id=\"preprints-h2-6\">Funding\u003C/h2>\n\u003Cdiv class=\"html-p\">The author thank the National University of Singapore for financial support.\u003C/div>\u003C/section>\u003Csection class=\"html-notes\">\u003Ch2 id=\"preprints-h2-7\">Conflicts of Interest\u003C/h2>\n\u003Cdiv class=\"html-p\">The authors declare no conflicts of interest.\u003C/div>\u003C/section>\u003Csection id=\"html-references_list\">\u003Ch2 id=\"preprints-h2-8\">References\u003C/h2>\n\u003Col class=\"html-x\">\n\u003Cli id=\"B1-preprints-73387\" class=\"html-x\" data-content=\"1.\">H. Tan, L. Fu, and M. Seno, “Optimization of Bacterial Plasmid Transformation Using Nanomaterials Based on the Yoshida Effect. \u003Cspan class=\"html-italic\">Int. J. Mol. Sci.\u003C/span> \u003Cb>2010\u003C/b>, \u003Cspan class=\"html-italic\">11\u003C/span>, 12. [\u003Ca href=\"https://doi.org/10.3390/ijms11124962\" class=\"cross-ref\" target=\"_blank\" rel=\"noopener noreferrer\">CrossRef\u003C/a>]\u003C/li>\n\u003Cli id=\"B2-preprints-73387\" class=\"html-x\" data-content=\"2.\">W.-T. Chan, C. S. W.-T. Chan, C. S. Verma, D. P. Lane, and S. K.-E. Gan, A comparison and optimization of methods and factors affecting the transformation of Escherichia coli. \u003Cspan class=\"html-italic\">Biosci. Rep.\u003C/span> \u003Cb>2013\u003C/b>, \u003Cspan class=\"html-italic\">33\u003C/span>, e00086. [\u003Ca href=\"https://scholar.google.com/scholar_lookup?title=W.-T.+Chan,+C.+S.+Verma,+D.+P.+Lane,+and+S.+K.-E.+Gan,+A+comparison+and+optimization+of+methods+and+factors+affecting+the+transformation+of+Escherichia+coli&author=W.-T.+Chan,+C.+S.&publication_year=2013&journal=Biosci.+Rep.&volume=33&pages=e00086&doi=10.1042/BSR20130098&pmid=24229075\" class=\"google-scholar\" target=\"_blank\" rel=\"noopener noreferrer\">Google Scholar\u003C/a>] [\u003Ca href=\"https://doi.org/10.1042/BSR20130098\" class=\"cross-ref\" target=\"_blank\" rel=\"noopener noreferrer\">CrossRef\u003C/a>] [\u003Ca href=\"https://www.ncbi.nlm.nih.gov/pubmed/24229075\" class=\"cross-ref pub_med\" data-typ=\"pmid\" target=\"_blank\" rel=\"noopener noreferrer\">PubMed\u003C/a>]\u003C/li>\n\u003Cli id=\"B3-preprints-73387\" class=\"html-x\" data-content=\"3.\">A. Froger and J. E. Hall, “Transformation of Plasmid DNA into E. coli Using the Heat Shock Method. \u003Cspan class=\"html-italic\">JoVE J. Vis. Exp.\u003C/span> \u003Cb>2007\u003C/b>, e253. [\u003Ca href=\"https://doi.org/10.3791/253\" class=\"cross-ref\" target=\"_blank\" rel=\"noopener noreferrer\">CrossRef\u003C/a>]\u003C/li>\n\u003Cli id=\"B4-preprints-73387\" class=\"html-x\" data-content=\"4.\">C. T. Chung, S. L. C. T. Chung, S. L. Niemela, and R. H. Miller, One-step preparation of competent Escherichia coli: transformation and storage of bacterial cells in the same solution. \u003Cspan class=\"html-italic\">Proc. Natl. Acad. Sci. USA\u003C/span> \u003Cb>1989\u003C/b>, \u003Cspan class=\"html-italic\">86\u003C/span>, 2172–2175. [\u003Ca href=\"https://scholar.google.com/scholar_lookup?title=C.+T.+Chung,+S.+L.+Niemela,+and+R.+H.+Miller,+One-step+preparation+of+competent+Escherichia+coli:+transformation+and+storage+of+bacterial+cells+in+the+same+solution&author=C.+T.+Chung,+S.+L.&publication_year=1989&journal=Proc.+Natl.+Acad.+Sci.+USA&volume=86&pages=2172%E2%80%932175&doi=10.1073/pnas.86.7.2172&pmid=2648393\" class=\"google-scholar\" target=\"_blank\" rel=\"noopener noreferrer\">Google Scholar\u003C/a>] [\u003Ca href=\"https://doi.org/10.1073/pnas.86.7.2172\" class=\"cross-ref\" target=\"_blank\" rel=\"noopener noreferrer\">CrossRef\u003C/a>] [\u003Ca href=\"https://www.ncbi.nlm.nih.gov/pubmed/2648393\" class=\"cross-ref pub_med\" data-typ=\"pmid\" target=\"_blank\" rel=\"noopener noreferrer\">PubMed\u003C/a>]\u003C/li>\n\u003Cli id=\"B5-preprints-73387\" class=\"html-x\" data-content=\"5.\">J. Ohlson, Plasmid manufacture is the bottleneck of the genetic medicine revolution. \u003Cspan class=\"html-italic\">Drug Discov. Today\u003C/span> \u003Cb>2020\u003C/b>, \u003Cspan class=\"html-italic\">25\u003C/span>, 1891–1893. [\u003Ca href=\"https://doi.org/10.1016/j.drudis.2020.09.040\" class=\"cross-ref\" target=\"_blank\" rel=\"noopener noreferrer\">CrossRef\u003C/a>] [\u003Ca href=\"https://www.ncbi.nlm.nih.gov/pubmed/33075470\" class=\"cross-ref pub_med\" data-typ=\"pmid\" target=\"_blank\" rel=\"noopener noreferrer\">PubMed\u003C/a>]\u003C/li>\n\u003Cli id=\"B6-preprints-73387\" class=\"html-x\" data-content=\"6.\">F. Silva, J. A. Queiroz, and F. C. Domingues, Evaluating metabolic stress and plasmid stability in plasmid DNA production by Escherichia coli. \u003Cspan class=\"html-italic\">Biotechnol. Adv.\u003C/span> \u003Cb>2012\u003C/b>, \u003Cspan class=\"html-italic\">30\u003C/span>, 691–708. [\u003Ca href=\"https://doi.org/10.1016/j.biotechadv.2011.12.005\" class=\"cross-ref\" target=\"_blank\" rel=\"noopener noreferrer\">CrossRef\u003C/a>] [\u003Ca href=\"https://www.ncbi.nlm.nih.gov/pubmed/22244816\" class=\"cross-ref pub_med\" data-typ=\"pmid\" target=\"_blank\" rel=\"noopener noreferrer\">PubMed\u003C/a>]\u003C/li>\n\u003C/ol>\u003C/section>\u003Csection id=\"Figures\" type=\"display-objects\">\u003Cdiv class=\"html-fig-wrap\" id=\"preprints-73387-f003\">\n \u003Cdiv class=\"html-fig_img\">\n \u003Cdiv class=\"html-figpopup html-figpopup-link\" href=\"#fig_body_display_preprints-73387-f003\">\n \u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png\" alt=\"Preprints 73387 g003\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png\">\n \u003Ca class=\"html-expand html-figpopup\" href=\"#fig_body_display_preprints-73387-f003\">\u003C/a>\n \u003C/div>\n\n \u003C/div>\n \u003Cdiv class=\"html-fig_description\">\n \u003Cb>Figure 3.\u003C/b>\n Comparison between “Fast” transformation and conventional transformation. “Fast” transformation refers to transformation on ice with 2 min incubation and no heat shock. On the other hand, conventional transformation refers to the typical workflow outlined in \u003Ca href=\"#preprints-73387-f001\" class=\"html-fig\">Figure 1\u003C/a> with 30 min incubation on ice and with heat shock and outgrowth.\n\u003C!-- \u003Cp>\u003Ca class=\"html-figpopup\" href=\"#fig_body_display_preprints-73387-f003\">\n Click here to enlarge figure\n \u003C/a>\u003C/p> -->\n\n \u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig_show mfp-hide\" id=\"fig_body_display_preprints-73387-f003\">\n \u003Cdiv class=\"html-caption\"> \u003Cb>Figure 3.\u003C/b>\n Comparison between “Fast” transformation and conventional transformation. “Fast” transformation refers to transformation on ice with 2 min incubation and no heat shock. On the other hand, conventional transformation refers to the typical workflow outlined in \u003Ca href=\"#preprints-73387-f001\" class=\"html-fig\">Figure 1\u003C/a> with 30 min incubation on ice and with heat shock and outgrowth.\u003C/div>\n \u003Cdiv class=\"html-img\">\u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png\" alt=\"Preprints 73387 g003\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g003.png\">\u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig-wrap\" id=\"preprints-73387-f004\">\n \u003Cdiv class=\"html-fig_img\">\n \u003Cdiv class=\"html-figpopup html-figpopup-link\" href=\"#fig_body_display_preprints-73387-f004\">\n \u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png\" alt=\"Preprints 73387 g004\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png\">\n \u003Ca class=\"html-expand html-figpopup\" href=\"#fig_body_display_preprints-73387-f004\">\u003C/a>\n \u003C/div>\n\n \u003C/div>\n \u003Cdiv class=\"html-fig_description\">\n \u003Cb>Figure 4.\u003C/b>\n Effect of competent cell concentration on transformation efficiency. Experiments with pNAR plasmid in \u003Cspan class=\"html-italic\">Escherichia coli\u003C/span> DH5α indicated that low concentration competent cells (1.45 x 10\u003Csup>6\u003C/sup> CFU/ml) delivered higher transformation efficiency compared to high concentration competent cells (8.5 x 10\u003Csup>8\u003C/sup> CFU/ml).\n\u003C!-- \u003Cp>\u003Ca class=\"html-figpopup\" href=\"#fig_body_display_preprints-73387-f004\">\n Click here to enlarge figure\n \u003C/a>\u003C/p> -->\n\n \u003C/div>\n\u003C/div>\n\u003Cdiv class=\"html-fig_show mfp-hide\" id=\"fig_body_display_preprints-73387-f004\">\n \u003Cdiv class=\"html-caption\"> \u003Cb>Figure 4.\u003C/b>\n Effect of competent cell concentration on transformation efficiency. Experiments with pNAR plasmid in \u003Cspan class=\"html-italic\">Escherichia coli\u003C/span> DH5α indicated that low concentration competent cells (1.45 x 10\u003Csup>6\u003C/sup> CFU/ml) delivered higher transformation efficiency compared to high concentration competent cells (8.5 x 10\u003Csup>8\u003C/sup> CFU/ml).\u003C/div>\n \u003Cdiv class=\"html-img\">\u003Cimg data-large=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png\" data-original=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png\" alt=\"Preprints 73387 g004\" src=\"https://www.preprints.org/frontend/picture/ms_xml/manuscript/869f7a4b8e128c6dfc43eaa2e5a2a0ac/preprints-73387-g004.png\">\u003C/div>\n\u003C/div>\n\u003C/section>\u003Csection class=\"html-fn_group\">\u003Ctable>\u003Ctr id>\n\u003Ctd>\u003C/td>\n\u003Ctd>\u003Cdiv class=\"html-p\">\n\u003Cb>Disclaimer/Publisher’s Note:\u003C/b> The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.\u003C/div>\u003C/td>\n\u003C/tr>\u003C/table>\u003C/section>\n \u003Csection id=\"html-copyright\">\u003Cbr>© 2023 by the author. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (\u003Ca href=\"https://creativecommons.org/licenses/by/4.0/\" target=\"_blank\">https://creativecommons.org/licenses/by/4.0/\u003C/a>).\u003C/section>\n ",[163],{"version":7,"hash_key":127,"id":126,"change":50},"Ng, W. Understanding the Different Parameters That Affects Bacterial Transformation Efficiency. \u003Cem>Preprints\u003C/em> \u003Cb>2023\u003C/b>, 2023050849. https://doi.org/10.20944/preprints202305.0849.v1",[],[],["Reactive",168],{"$scookieConsent":169,"$stoast-ui":170,"$ssite-config":173},{"necessary":6,"functional":132,"statistic":132,"marketing":132,"unclassified":132},{"showToast":132,"toastMessage":171,"toastVariant":172},"","neutral",{"env":174,"name":175,"url":176},"production","nuxt-app","http://frontend.preprints.org/",["Set"],["ShallowReactive",179],{"fE3cVXSjOY":50,"mgOlEjP40X":50,"Ya7hZzqdE7":50},"/manuscript/202305.0849/v1",["Reactive",182],{"auth":183,"share-modal":185,"useHeader":186},{"userInfo":184,"isLogin":132},{},{"isShareModalOpen":132,"item":50},{"height":41}]</script> 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