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. 2019 Sep 3;10:750. doi: 10.3389/fgene.2019.00750

Table 7.

The advantages and disadvantages of genetic manipulation tools applicable to ovine and caprine genomes

Tool Uses Advantages Disadvantages References
PNI Insertional transgenesis The first tool to be applied for the generation of transgenic animals Random integration, variable transgene copy number, low efficiency Hammer et al., 1985; Clark, 2002
SCNT Gene targeting, editing An alternative that facilitated the implementation of HR gene targeting in species that lack ESCs, a low-level mosaicism Development of a small proportion of reconstructed embryos that become live offspring, potential complications at birth of offspring as a result of developmental abnormalities Schnieke et al., 1997; Wilmut et al., 1997; Wilmut et al., 1999
SMGT Gene transfer, integration Simple, cost-effective, minimal embryo handling required Initial doubt with regard to its repeatability, variable results, low incorporation of the exogenous gene Lavitrano et al., 1989; Wall, 2002; Lavitrano et al., 2006
VMGT Gene transfer, integration Able to infect germline cells and dividing or non-dividing somatic cells, delivery of the system to the egg/zygote is less damaging compared to pronuclear injection, high integration Variability of transgenic expression, potential health risks, limited DNA capacity Whitelaw et al., 2008; Modric and Mergia, 2009
Recombinases Integration, selectable cassette excision Increased gene integration efficiency, offer different forms of modifications including the removal of unwanted DNA Conservative specificity, in specific cases, pre-introduction of specific target sites within the host genome is required which is an inefficient and time-consuming process, potential toxicity Xu et al., 2008; Gaj et al., 2014; Olorunniji et al., 2016
Transposons Integration Able to integrate transgenes and RNAi-expressing constructs for the mediation of knockdown expression, lower immunogenicity and larger DNA capacity compared to viral systems Classical transposons are less efficient for gene transfer compared to viral systems, potential cytotoxicity Muñoz-López and García-Pérez, 2010; Meir and Wu, 2011; Hudecek et al., 2017
RNAi Gene knock down Targeting gene expression at mRNA level, useful tool to elucidate gene functions Variability and incompleteness of knockdowns, potential off-target Boutros and Ahringer, 2008; Boettcher and McManus, 2015; Bradford et al., 2017
ZFNs Gene editing First “practical” endonuclease that has been applied to mediated gene-editing events Difficult to design, potential off-target, mosaicism in offspring generated from microinjected embryos Gaj et al., 2013; Gupta and Musunuru, 2014; Oliver et al., 2015; Zhang et al., 2019a
TALENs Gene editing A simplified alternative of the previously emerged ZFNs Moderate difficulty in design, potential off-target, mosaicism in offspring generated from microinjected embryos Bedell et al., 2012; Gaj et al., 2013; Gupta and Musunuru, 2014; Zhang et al., 2019a
CRISPR/Cas9 Gene editing Simple, cost-effective, customizable, precise compared to other endonucleases, able to mediate multiplex editing Potential off-target, mosaicism in offspring generated from microinjected embryos Gaj et al., 2013; and Gupta and Musunuru, 2014; Mehravar et al., 2019; Zhang et al., 2019a

PNI, pronuclear injection; SCNT, somatic cell nuclear transfer; SMGT, sperm-mediated gene transfer; VMGT, virus-mediated gene transfer; RNAi, RNA interference; ZFNs, zinc finger nucleases; TALENs, transcription activator-like effector nucleases; CRISPR/Cas9, clustered regularly interspaced short palindromic repeat/CRISPR-associated protein 9.