Summary of cotyledonary-node transformation system.
1. Introduction
Soybean (
Over the last two decades, the transfer of DNA into plant cells has been achieved by using several methods. In soybeans, the most frequently employed plant genetic engineering methods are
2. Different approaches for soybean transformation
In soybean transformation, two major methods are now widely utilized:
2.1. Cotyledonary-node-based transformation
The routine regeneration system was first reported by using the mature cotyledonary-node [14]. The multiple adventitious buds and shoots from explant tissues were proliferated and regenerated on culture media containing cytokinin by organogenesis. The transgenic soybean plants have been successfully and reproducibly produced using mature or immature cotyledon explants via
Recently, an alternative cotyledonary explant derived from mature soybean seed for
In fact, several laboratories have contributed to enhanced soybean transformation using a cotyledonary-node explant. To overcome the low transfer of
Since the transformation process by use of kanamycin or hygromycin B as selection agent has been proven to be genotype-dependent, the most widely used selection system has been the combination of
2.2. Immature embryos-based transformation
The regeneration using immature embryos via somatic embryogenesis was first reported by Christianson et al. [23]. The immature embryos excised from soybean pods were suspended on semi-solid media or liquid media containing high concentration of auxin, 2,4-Dichlorophenoxyacetic acid (2,4-D), and the whole plantlets were recovered [24-25]. After immature embryos were developed as an alternative plant material, transgenic plants were first obtained from this explant tissue via particle bombardment [26]. This system has been exclusively used to produce transgenic soybean such as glyphosate tolerant, hygromycin resistance, and
The use of particle bombardment with immature embryos tends to be highly variable, and multiple copies of the introduced DNAs are commons. Moreover, this problem has compounded with aged embryogenic suspension cultures from which a high percentage of regenerated plants lost their fertility [29]. In spite of this limitation, the embryogenic cultures have several advantages, one of which is its relatively high transformation efficiency and less chimeric plants recovered.
2.3. Embryogenic shoot tips-based transformation
The embryonic shoot tip explant is another source of explant which has been used for soybean transformation. McCabe et al., [30] first reported the stable transformation using meristemic cell, shoot apex, by particle acceleration. The shoot derived from these meristems via organogenesis has been produced to form multiple shoots prior to mature plants. However, all of the primary transgenic plants were chimeric. Martinell et al., [31] described the successful method using meristemic shoot tip from germinated seedling by
2.4. Immature cotyledonary-nodes
The regeneration capacity of immature cotyledonary-node was found by Parrott et al [33]. Based on this regeneration system, first transgenic soybean plants have been developed by
2.5. Hypocotyl based transformation
Another type of explant tissue, hypocotyl, was also investigated with 13 different soybean genotypes. Most of the genotypes initiated shoots from this type of explant [36]. This method was reported to be genotype-independent regeneration protocol via organogenesis and utilized the acropetal end of a hypocotyl section from a 7-day old seedling. Despite inducing adventitious shoots from the explant, most recovered shoot did not matured in the soil. Wang et al [37] reported successful production of fertile transgenic plants using hypocotyl-based
2.6. Leaf tissue-based transformation
The reproducible regeneration methods for whole plants from primary leaf tissue or epicotyls were first reported by Wright et al [38]. The multiple shoots from those explants were continually initiated and proliferated with cytokinin BAP hormone. Rajasckaren et al [39] described regeneration of several varieties of soybean by embryogenesis from epicotyls and primary leaf tissues, thereby inducing fertile plants from those explants. Kan et al [40] first tested transformation efficiency using epicotyls and leaf tissues by
3. Agrobacterium- mediated transformation of soybean
3.1. Agrobacterium- mediated transformation mechanism
The T-complex is imported into the nucleus by the phosphorylation of VirE2 Interacting Protein 1 (VIP1), induced by mitogen-activated protein kinase (MAPK), such as MPK3 [55]. After T-complex is imported into the host nucleus, VirE2 and VIP1 need to be degraded before T-DNA integration by a subunit of the SCF (SKP-CUL1-F-box protein) ubiquitin E3 ligase complex. Not only
3.2. History of Agrobacterium- mediated soybean transformation research
Among various transformation technologies,
After Hinchee et al., [8] developed
Recently,
There has been a significant improvement in soybean transformation over the past two decades. However, the efficiency of soybean transformation is not great enough for practical needs and shows high variation. Thus, considering the potential application of soybean transformation, the importance of
A208 | Peking, Maple Prest |
|
kanamycin | (8) |
AGL1 | Bert |
|
phosphinothricin | (67) |
EHA101 | Williams 82 | bar | glufosinate | (12) |
EHA101 | Williams, Williams 79, Peking, Thorne |
|
glufosinate or bialaphos | (68) |
EHA101 | Thorne, Williams, Williams 79, Williams 82 |
|
glufosinate | (19) |
EHA105 | AC Colibri |
|
kanamycin | (69) |
EHA105 | Hefeng 25, Dongnong 42, Heinong 37, Jilin 39, Jiyu 58 |
|
hygromycin | (70) |
EHA105 | A3237 |
|
glufosinate | (10) |
LBA4404 | Jungery |
|
phosphinothricin | (71) |
LBA4404, EHA105 | Bert |
|
hygromycin | (11) |
4. New directions of soybean genetic engineering, skills and vectors
To date, the
Among the various methods, co-transformation system is one of the most commonly used methods to produce marker free transgenic plants. In co-transformation systems, a marker gene and genes of interest are placed on separate DNA molecules and introduced into plant genomes. Then, the non-selectable genes segregate from the marker gene in the progeny generations. Most strains of
To improve plant genetic traits, many soybean research labs have developed tools for soybean functional genomics, such as several libraries containing large inserts of bacterial artificial chromosome (BAC) and plant transformation competent binary plasmids clone (BIBAC) (81). In functional genomic research, bacterial artificial chromosome (BAC) is a single copy artificial chromosome vector and is based on the
5. Zinc finger nucleases (ZFNs) and transcription activator-like effectors (TALEs)
Although many methods have been developed, soybean is considered a recalcitrant plant to transform compared to
Zinc finger nucleases (ZFNs) and meganucleases cut specific DNA target sequences
To overcome the ZFNs’s weakness, in late 2009, a novel DNA binding domain was identified which was a member of the large transcription activator-like (TAL) effector family [92-93]. Transcription activator-like effectors (TALEs) are produced by plant pathogens in the genus
6. Abbreviation
Acetosyringone: 4'-Hydroxy-3',5'-dimethoxyacetophenone
BAC: bacterial artificial chromosome
BAP: 6-Benzylaminopurine
2,4-D: 2,4-Dichlorophenoxyacetic acid
BIBAC: binary bacterial artificial chromosome
BT:
DSBs: double-strand breaks
HPT II: Hygromycin phosphotransferase
HR: homologous recombination
NAA: Naphthaleneacetic acid
NHEJ: non-homologous end-joining
NPT II: neomycin phosphotransferase II
RVD: repeat variable di-residue
Ti plasmid: tumour-inducing (Ti) plasmid
T4SS: type IV secretion system
TALEs: transcription activator-like effectors
VBF: VIP1-binding F-box
VIP1: VirE2 Interacting Protein 1
Vir protein: virulence protein
ZFNs: Zinc finger nucleases
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