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'''Transgenic plants''' are plants that possess [[gene]] or genes that have been transfered from a different species.  
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{{seealso|horizontal gene transfer in plants}}
'''Transgenic plants''' possess a [[gene]] or genes that have been transferred from a different species such as another plant, or a microorganism, or other source. They are created in nature during [[horizontal gene transfer]] 
They can also be created during [[plant breeding]], and especially in recent years through the use of the plant DNA [[transformation]] technique in [[biotechnology]]. (See [[Plant breeding]], [[Biotechnology and plant breeding]]).


The most efficient route for gene movement between species is cross-pollination, which generates hybrids.
==Introgression==
The most efficient natural route for gene movement between plant species is by [[cross-pollination]] to form fertile plant inter-species hybrids. Such hybrids are sometimes new plant species, but they can also form a gene transmission "bridge" in the hybrid-zone between two distinct plant populations, and thus facilitate [[introgression]], which is movement of genes from one distinct species or population to another.


For gene movement to be successful the added genes must replicate themselves successfully in in some way, or be inserted into existing chromosomes which are able to replicate succesfully. The cells carrying the added genes must also produce the normal reproductive process for the plant, that is,  in most cases produce fertile seeds.
==More distant natural transfers==
The best documented route for natural formation of transgenic plants is gene transfer between a plant [[epiphyte]] (such as [[moss]es]), or a parasitic plant (like dodder) and the host plant it colonizes.  


These conditions are most likely to be met when cross-pollination is carried out between '''closely related plant species'''. '''Distantly related species''' generally fail to succesfully cross-pollinate at some point, for instance by producing infertile seeds, or by lacking two copies of every chromosome which are needed to participate in sucessful [[meiosis]].
Non-standard fertilization of with more than one pollen grain has also been suggested for transfer of a nuclear genome located  gene from ''Poa'' grass genus into the distantly related sheep's fescue, ''Festuca ovina''<ref>Ghatnekar L, Jaarola M, Bengtsson BO.(2006) The introgression of a functional nuclear gene from ''Poa'' to ''Festuca ovina''.Proc Biol Sci. 2006 Feb 22;273(1585):395-9.</ref>.


There are, however, natural process for gene transfer between different species in addition to carriage of genes in pollen and straight-forward [[sexual recombination]].
Another mechanism for [[horizontal gene transfer]] is ''[[Agrobacterium tumefaciens]]'' and similar bacteria that inject DNA into plant cells. Biotechnology laboratories exploit ''[[Agrobacterium tumefaciens]]'' bacteria to make artificial transgenic plants with small segments of added transgenic DNA inserted in the host cell chromosomes. Others mechanisms may include plant sucking insects, mites, and possibly viruses.


One example is ''[[Agrobacterium tumefaciens]]'', which is a bacterium that injects DNA into plant cells. Biotechnology laboratories exploit this bacterium to make artificial transgenic plants with small segment of added DNA inserted in the host cell chromosomes.
Recent comparative studies of gene content of different [[genomes]]  provides strong circumstantial evidence that natural horizontal gene transfer does occur in plants at a frequency that is significant over evolutionary time scales. Over the evolutionary time-scales plant [[mitochondria]] are a stopping point for genes that may enter the nuclear genome from other species, and can in some cases be very active in inter-species gene-traffic (See [[Horizontal gene transfer]]).


But there are yet other routes for gene movement. This is shown by the widespread movement of mobile DNA such [[transposons]] and MULE elements between different species. It is also known that mobile DNA can move from site to site within a genome, and can sometimes mobilize adjacent genes during this movement, so the possibilities for [[horizontal gene transfer]] could include a combination of different mechanisms, including movements catalysed by mobile DNA. Chromosomes lost from progeny after inter-species cross-pollination can still occasionally leave behind DNA remnants with the help of mobile DNA.
The  natural DNA transfer of [[mobile DNA]] between rice and millet is well documented <ref>[http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040035 Jumping Genes Cross Plant Species Boundaries.]'' ''PLoS Biol 4(1): e35 DOI:10.1371/journal.pbio.0040035''
''Published: December 20, 2005''</ref>, but genes involved in metabolic process and bacterial genes can also take part in natural genes transfers (See [[horizontal gene transfer in plants]]).


A study has been carried out on MULE gene movement beween rice and millet specifically illustrates the type of recent discoveries that are being made about natural transgenic events, and this study is described fully below under the heading '''Jumping Genes Cross Plant Species Boundaries'''.  
==Hybrid formation in flowering plants and its role in introgression of genes between species==
Cross-pollination between plant species generates interspecies hybrids occurs widely in nature and has been exploited in plant breding for more than 100 years to create artificial transgenic plants ( see [[Plant breeding]]).


==Hybrid formation in flowering plants==
Hybrids can occur in the intermediate geographical zone between two species and provide a "bridge" for genes to "introgress" (or move) from one species to another<ref> Rieseberg, L.H. and Wendel, J. (1993). Introgression and its consequences in plants. In Hybrid Zones and the Evolutionary Process. (ed. J. Harrison) p 70-109, Oxford University Press, New York.</ref> <ref>Rieseberg, L.H. and Ellstrand, N.C. (1993) What can molecular and morphological markers tell us about plant hybridization/ Critical Reviews of Plant Science. 12 p213-241. </ref>.


Hybrid formation between two species by pollination joins two sets  of chromosomes together, one from each parent, is a common event in flowering plant evolution, and the main way new plant species are formed. Interestingly, in many cases hybrids are formed by adding '''two copies''' of each chromosome from each parent, forming a '''tetaploid''' that is reproductively isolated from both parents - and a new species.
Hybrid formation between two species by pollination joins two sets  of chromosomes together, one from each parent, is a common event in flowering plant evolution, and the main way new plant species are formed. Interestingly, in many cases hybrids are formed by adding two copies of each chromosome from each parent, forming an allotetraploid that is reproductively isolated from both parents - and a new species<ref>Ramsey, J. and Schemske, D.W. (1998) Pathways, mechanisms, and rates of polyploid formation in flowering plants. Annual Review of Ecology and Systematics. 29, 467-501.</ref>.  


Wild emmer [[wheat]] is an example of a species formed by hybridization between two diploid wild grasses, ''Triticum urartu'' and a wild goatgrass such as ''Aegilops searsii'' or ''Ae. speltoides'' four sets of chomosomes (is a tetrapoid. [[Triticale]] is crop cultivated today mostly for forage and animal feed which is an artificial hybrid  between [[rye]] and wheat, first bred during the late 19th century.
Wild emmer [[wheat]] is an example of a species formed by hybridization between two [[diploid]] wild grasses, ''Triticum urartu'' (AA) and a wild goatgrass ''Ae. speltoides'' (BB) to form hybrid new species with four sets of chomosomes (AABB) which is a [[tetraploid]]. ''[[Triticale]]'' ''(Triticosecale)'' is a crop cultivated today mostly for forage and animal feed which is an artificial hybrid  between [[rye]] and [[wheat]], first bred during the late 19th century.


A surprising number of plants show evidence of being formed by such chromosome set addition processes; [[wheat]] and [[cotton]] are two other examples.
A surprising number of plants show evidence of being formed by such processes by which chromosome sets are  added : bread [[wheat]] ( an allohexaploid having three component genomes) , and [[cotton]] are two other examples <ref>J. A. Udall and J. F. Wendel (2006) Polyploidy and Crop Improvement. Crop Sci. 46, S-3-S-14 </ref>.


Gene transfer could occur by wide cross-pollination even if foreign chromosomes are lost. It has been suggested that 'wide crosses' are  a possible mechanism  of horizontal transfer of [[mobile DNA]] in plants, and that these might transfer only [[mobile DNA]]s, due to one of the participating sets of chromosomes being lost <ref>Ananiev EV, Riera-Lizarazu O, Rines HW, Phillips RL (1997) Oat maize chromosome addition lines: a new system for mapping the maize genome. Proc Natl Acad Sci USA 94: 3524–3528.</ref><ref>Bennetzen, J. L., (2000) Transposable element contributions to plant gene and genome evolution. Plant Molecular Biology 42: 251–269, 2000.</ref>


==Natural movements of genes between species by other routes than pollen==


==Natural movements of genes between species.==
:''See [[Horizontal gene transfer in plants]], [[Horizontal gene transfer]]''
Natural movement of genes between species, often called [[horizontal gene transfer]] or lateral gene transfer, can also because of gene transfer mediated by natural agents such as microrganisms, viruses or mites. Such transfers occur at a frequency that is low compared with the hybridization that occurs during natural pollination, but can be frequent enough to be a significant factor in genetic change of a [[chromosome]] on evolutionary time scales <ref>Syvanen, M. and Kado, C. I. Horizontal Gene Transfer. Second Edition. Academic Press 2002.</ref> 


This natural gene movement between species has been widely detected during genetic investigation of various natural [[Mobile genetic elements]], such as [[Transposon|Transposons]], and [[Retrotransposon|Retrotransposons]] that naturally transfer to new locations in a [[Genome]], and often move to new species host over an evolutionary time scale. There are many types of natural mobile DNAs, and they have been detected abundantly in food crops such as rice <ref>[http://nar.oxfordjournals.org/cgi/content/full/33/7/2153 DNA-binding specificity of rice mariner-like transposases and interactions with Stowaway MITEs]</ref>.
==Transgenic plants and crop improvement==


These various mobile or jumping genes play a major role in dynamic changes to chromosomes during evolution <ref>[http://www.pnas.org/cgi/content/full/103/21/8101]</ref>, <ref>[http://www.nature.com/nrg/journal/v4/n11/abs/nrg1204_fs.html]</ref>, and have often been given whimsical nanes, such as Mariner, Hobo, Trans-Siberian Express (Transib), Osmar, Helitron, Sleeping Princess, MITE and MULE, to emphasise their mobile and transient behaviour.  
Production of transgenic plants in 'wide-crosses' by plant breeders has been a vital aspect of conventional [[plant breeding]] for a century or so. Without it, security of our food supply against losses caused by crop pests such as rusts and mildews would be severely compromised. The first historically recorded interpecies transgenic cereal hybrid was actually between wheat and rye (Wilson, 1876).


Such genetically mobile DNA contitutute a major fraction of the DNA of many plants, and the natural dynamic changes to crop plant chromosomes caused by this natural transgenic DNA mimics many of the features of plant genetic engineering currently pursued in the laboratory, such as using [[Transposons as a genetic tool]], and molecular cloning. ''See also'' [[Transposon]], [[Retrotransposon]], [[Integron]], [[Provirus]], [[Endogenous retrovirus]], [[Heterosis]], <ref>[http://www.nature.com/ng/journal/v37/n9/abs/ng1615.html;jsessionid=367F14297326E4C7BF28B89F461CDB46 Gene duplication and exon shuffling by helitron-like transposons generate intraspecies diversity in maize.]</ref>
Transgenic varieties are frequently created by classical breeders by deliberately and artificially force hybridisation between distinct plant species with the intention of developing disease resistant crop varieties. Classical plant breeders may use use of a number of ''in vitro'' techniques such as protoplast fusion, embryo rescue or mutagenisis to generate diversity and produce plants that would not exist in nature (''see also [[Plant breeding]], [[Heterosis]], [[New Rice for Africa]]''). Chromosomal rearrangements and translocations occurring in these crosses help limit the amount of new DNA appearing in the final cultivated variety to a fraction of a chromosome, but still comprise substantial numbers of novel genes introduced into food.


There is large and growing scientific literature about natural transgenic events in plants, such as the creation of shibra millet in Africa, and movement of natural mobile DNAs called MULEs between rice and millet <ref>[http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040035]</ref>.
These "classical" techniques (used extensively since about 1930 on) have never been controversial, or been given wide publicity except among professional biologists, and have allowed crop breeders to develop varieties of basic food crop, wheat in particular, which resist devastating plant diseases such as rusts. ''Hope'' is one such transgenic wheat variety bred by E. S. McFadden with a transgene from a wild grass. ''Hope'' saved American wheat growers from devastating stem rust outbreaks in the 1930s.
An article about natural MULE gene movement between rice and millet is worth describing fully:


===Jumping Genes Cross Plant Species Boundaries===
Introduction of alien germplasm into common foods has repeatedly achieved novel genetic rearrangements of plant chromosomes, such as insertion of large blocks of rye (''Secale'') genes into wheat chromosomes ('[[translocations]]')<ref>[http://www.pnas.org/cgi/content/abstract/96/11/5937]</ref>.


<blockquote> In the early 1950s, legendary plant geneticist [[Barbara McClintock]] found the first evidence that genetic material can jump from one place to another within the genome. The variegated kernels of her [[maize]] plants, she determined, resulted from mobile elements that had inserted themselves into pigment-coding genes, changing their expression. McClintock's mobile elements, or [[transposons]], moved over generations within a single species. More recently, another form of genetic mobility has been discovered—genetic information can sometimes be transferred between species, a process called [[horizontal gene transfer]]. While horizontal genetic transfer occurs most commonly in bacteria, it has been detected in animals as well. Most transfers between higher animals involve the movement of transposons. Horizontal transfer can also occur between the mitochondrial DNA of different plant species. Until now, however, no one had found evidence for horizontal transfer in the nuclear DNA of plants.
The advent of drug [[colchicine]] in the late 1930s helped overcome fertility barriers in inter-specific crosses by stimulating doubling of chromosome numbers per cell, and after 1930 perennial wild-grasses were being frequently hybridized with wheat and other cereals with the aim of transferring disease resistance and perenniality into annual crops. Large-scale practical use of hybrids became well established, leading on to development of numerous ''Triticosecale'' (''[[Triticale]]'') varieties and other new transgenic cereal crops.
:
In a new report, Xianmin Diao, Michael Freeling, and Damon Lisch studied the genomes of millet and rice, '''two distantly related grasses that diverged 30–60 million years ago'''. While the two grasses show significant genetic divergence from accumulating millions of years of mutations, they carry some transposon-related DNA segments '''that are surprisingly similar'''. The authors conclude that these sequences were transferred horizontally between the two plants long after they went their separate ways.
:
Transposons of the class identified by Diao et al. typically consist of a variable length of DNA that codes for one or more enzymes flanked by repeating sequences called '''terminal inverted repeats''' ('''TIRs'''). These repeats can bind to each other to form a “lollipop” that is easily excised from the DNA strand, carrying the rest of the transposon along with it. Plant genomes are rife with transposons, many of which are relatively passive. '''Transposons from the “Mutator” family in maize, however, are especially active,''' frequently causing mutations as they insert themselves into new positions in the genome. They perform this jump with assistance from the two proteins they code for, a transposase and a helper gene.
:
DNA from many species of plants contains several families of '''cousins of the Mutator transposons'''. These “'''Mutator-like elements,” or MULEs''', code for a protein similar to the transposase, as well as the TIR sequences. Diao et al. identified 19 distinct MULEs in the DNA of various species of millet (genus Setaria), and compared these with the rice genome sequence, which was published in 2002. They compared the sequence similarity of these MULEs to that of other proteins that are also conserved in the same species for which sequences are available. Strikingly, they observed much higher sequence similarity between the MULEs from millet and rice than is typical for transposons. The greater similarity of the MULE DNA is easily explained if it jumped somehow, horizontally, between the species, but there could be alternative explanations. The match could have arisen without horizontal transfer, for example, if the MULE DNA had been under positive selection, as typically happens for protein-coding genes that confer some survival or reproductive benefit. In such cases, natural selection tends to preserve the integrity of these sequences.
:
To test for signs of selection, the researchers looked at regions of the MULE DNA that don't appear to code for protein. The similarity between these noncoding regions in millet and rice MULEs was just as high as for the coding regions, even though selection probably doesn't influence them. Even within the coding sections, “synonymous” mutations—which don't change the protein sequence and so are not prone to selection—showed few differences between these elements.
:
Another explanation for the low divergence of the rice and millet MULE sequences could be that they occur within a genomic region that, for whatever reason, experienced lower than average mutation rates. If this were the case, sequences adjacent to the elements should also show reduced variation. '''The authors tested this alternative hypothesis''' with the help of maize, which has more genomic sequence available than millet, by comparing genes flanking MULE regions in rice with evolutionarily conserved sequences in maize. The sequences did not show the similar degree of reduced variation predicted for below-average mutation rates.
:
Since neither selection nor low mutation frequency can explain the similar DNA between the grasses, the authors conclude, '''a transposon must have carried it between millet and rice long after these species diverged.''' Interestingly, the authors also found '''similar sequences in bamboo''', raising the question of how common horizontal transfer may be between plant species. '''Given that plant mitochondrial genes appear “particularly prone to horizontal transfer,”''' the authors note, “it is remarkable that these results represent the first well-documented case of horizontal transfer of nuclear genes between plants.” But as researchers begin to explore the growing databases of plant genomic sequences, they can determine whether this finding constitutes an anomaly—or points to a significant force in plant genome evolution. —Don Monroe</blockquote>


''Citation: (2006) [http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040035 Jumping Genes Cross Plant Species Boundaries.]'' ''PLoS Biol 4(1): e35 DOI:10.1371/journal.pbio.0040035''
Important transgenic pathogen and parasite resistance traits carried in current bread wheat varieties are <ref>[http://www.pnas.org/cgi/content/abstract/96/11/5937 Plant genetic resources: What can they contribute toward increased crop productivity? Hoisington, D. and others (1999) Proc. Natl. Acad Sci USA. Vol. 96, Issue 11, 5937-5943, May 25, 1999. (This paper was presented at the National Academy of Sciences colloquium "Plants and Population: Is There Time?" held December 5-6, 1998, at the Arnold and Mabel Beckman Center in Irvine, CA).]</ref>:  
''Published: December 20, 2005''
<!--THIS TABLE NEEDS RE-DOING IN WORD PLUS WORD2WIKI-->
{| <hiddentext>generated with [[:de:Wikipedia:Helferlein/VBA-Macro for EXCEL tableconversion]] V14<\hiddentext>
|-  valign="bottom"
|style="font-weight:bold" width="153.75" Height="12.75" | Disease resistance to Leaf rust
| width="107.25" |
| width="48" |
|style="font-weight:bold" width="231.75" | Disease resistance to powdery mildew
| width="98.25" |


''Copyright: © 2005 Public Library of Science. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.''
|-   valign="bottom"
| Height="12.75" | Gene
| Source
|
| Gene
| Source


|-  valign="bottom"
| Height="12.75" | Lr9
|style="font-style:Italic" | Aegilops umbellulata
|
| Pm12
|style="font-style:Italic" | Aegilops speltoides


|-  valign="bottom"
| Height="12.75" | Lr18
|style="font-style:Italic" | Triticum timopheevi
|
| Pm21
|style="font-style:Italic" | Haynaldia villosa


It is thus becoming clear that natural rearrangments of DNA and generation of transgenes play a pervasive role in natural evolution. Importantly many, if not most, flowering plants evolved by transgenesis - that is, the creation of natural interspecies hybrids in which chromosome sets from different plant species were added together. There is also the long and rich history of transgenic varieties in traditional breeding.
|-  valign="bottom"
| Height="12.75" | Lr19
|style="font-style:Italic" | Thinopyrum
|
| Pm25
|style="font-style:Italic" | T. monococcum


==Transgenic plants and crop improvement==
|-  valign="bottom"
| Height="12.75" | Lr23
|style="font-style:Italic" | T. turgidum
|
|
|
 
|-  valign="bottom"
| Height="12.75" | Lr24
|style="font-style:Italic" | Ag. elongatum
|
|style="font-weight:bold" | Disease resistance to wheat streak mosaic virus
|
 
|-  valign="bottom"
| Height="12.75" | Lr25
|style="font-style:Italic" | Secale cereale
|
| Wsm1
|style="font-style:Italic" | Ag. elongatum
 
|-  valign="bottom"
| Height="12.75" | Lr29
|style="font-style:Italic" | Ag. elongatum
|
|
|
 
|-  valign="bottom"
| Height="12.75" | Lr32
|style="font-style:Italic" | T. tauschii
|
|style="font-weight:bold" | Pest resistance
|
 
|-  valign="bottom"
| Height="12.75" |
|
|
|style="font-weight:bold" | Hessian fly
|
 
|-  valign="bottom"
|style="font-weight:bold" Height="12.75" | Disease resistance to stem rust
|
|
| H21
|style="font-style:Italic" | Secale cereale


Production of transgenic plants in '''wide-crosses''' by plant breeders has been a vital aspect of conventional [[Plant breeding]] for a century or so. Without it, security of our food supply against losses caused by crop pests such as rusts and mildews would be severely compromised. The first historically recorded interpecies transgenic cereal hybrid was actually between wheat and rye (Wilson, 1876).
|-   valign="bottom"
| Height="12.75" | Sr2
|style="font-style:Italic" |  T. turgidum ("Hope" <ref>McFadden, E. S. (1930) J. Am. Soc. Agron. 22, 1020-1031.</ref>)
|
| H23
|style="font-style:Italic" | Secale cereale


Transgenic varieties are frequently created by classical breeders by deliberately and artificially force hybridisation between distinct plant species with the intention of developing disease resistant crop varieties. Classical plant breeders may use use of a number of ''in vitro'' techniques such as protoplast fusion, embryo rescue or mutagenisis to generate diversity and produce plants that would not exist in nature (''see also [[Plant breeding]], [[Heterosis]], [[New Rice for Africa]]''). Chromosomal rearrangements and translocations occurring in these crosses help limit the amount of new DNA appearing in the final cultivated variety to a fraction of a chromosome, but still comprise substantial numbers of novel genes introduced into food.
|-  valign="bottom"
| Height="12.75" | Sr22
|style="font-style:Italic" | Triticum monococcum
|
| H24
|style="font-style:Italic" | T. tauschii


These "classical" techniques (used extensively since about 1930 on) have never been controversial, or been given wide publicity except among professional biologists, and have allowed crop breeders to develop varieties of basic food crop, wheat in particular, which resist devastating plant diseases such as rusts. ''Hope'' is one such transgenic wheat variety bred by E. S. McFadden with a transgene from a wild grass. ''Hope'' saved American wheat growers from devastating stem rust outbreaks in the 1930s.
|-  valign="bottom"
| Height="12.75" | Sr36
|style="font-style:Italic" | Triticum timopheevii
|
| H27
|style="font-style:Italic" | Aegilops ventricosa


Introduction of alien germplasm into '''common foods has repeatedly achieved novel genetic rearrangements of plant chromosomes''', such as insertion of large blocks of rye (Secale) genes into wheat chromosomes ('[[translocations]]')<ref>[http://www.pnas.org/cgi/content/abstract/96/11/5937]</ref>.  
|-  valign="bottom"
| Height="12.75" |
|
|
|
|


The  advent of drug [[colchicine]] in the late 1930s helped overcome fertility barriers in inter-specific crosses by stimulating doubling of chromosome numbers per cell, and after 1930 perennial wild-grasses were being frequently hybridized with wheat and other cereals with the aim of transferring disease resistance and perenniality into annual crops. Large-scale practical use of hybrids became well established, leading on to development of numerous Triticosecale ([[Triticale]]) varieties and other new transgenic cereal crops.
|-   valign="bottom"
|style="font-weight:bold" Height="12.75" | Disease resistance to stripe rust
|
|
|style="font-weight:bold" | Cereal cyst nematode
|


Important transgenic pathogen and parasite resistance traits carried in current bread wheat varieties (gene, eg "Lr9" followed by the source species) are:  
|-  valign="bottom"
| Height="12.75" | Yr15
|style="font-style:Italic" | Triticum dicoccoides
|
| Cre3
|style="font-style:Italic" | T. tauschii


'''Disease resistance to Leaf rust'''
|}
*Lr9 (from ''Aegilops umbellulata'')
*Lr18 ''Triticum timopheevi''
*Lr19 ''Thinopyrum''
*Lr23 ''T. turgidum''
*Lr24 ''Ag. elongatum''
*Lr25 ''Secale cereale''
*Lr29 ''Ag. elongatum''
*Lr32 ''T. tauschii''
'''Disease resistance to Stem rust'''
*Sr2 ''T. turgidum'' ("Hope" ) <ref>McFadden, E. S. (1930) J. Am. Soc. Agron. 22, 1020-1031.</ref>
*Sr22 ''Triticum monococcum''
*Sr36 ''Triticum timopheevii''
'''Stripe rust'''
*Yr15 ''Triticum dicoccoides''
'''Powdery mildew'''
*Pm12 ''Aegilops speltoides''
*Pm21 ''Haynaldia villosa''
*Pm25 ''T. monococcum''
'''Wheat streak mosaic virus'''
*Wsm1 ''Ag. elongatum''
'''Pest resistance'''
*'''Hessian fly'''
**H21 ''S. cereale'' H23,
**H24 ''T. tauschii''
**H27 ''Aegilops ventricosa''
*'''Cereal cyst nematode'''
**Cre3 (Ccn-D1) ''T. tauschii''


The intentional creation of transgenic plants by laboratory based [[recombinant DNA]] methods is more recent (from the mid-1980s on) and has been a controversial development opposed vigourously by many NGOs, and several governments, particularly within the European Community. These transgenic recombinant plants (= biotech crops, modern transgenics) are transforming agricultural productivity in those regions that have allowed farmers to adopt them, and the area sown to these crops has continued to grow globally in each of the ten years since their first introduction in 1996.
The intentional creation of transgenic plants by laboratory based [[recombinant DNA]] methods is more recent (from the mid-1980s on) and has been a controversial development opposed vigourously by many NGOs, and several governments, particularly within the European Community. In those regions that have allowed farmers to adopt them these transgenic recombinant plants (= biotech crops, modern transgenics) are transforming agricultural productivity, and the area sown to these crops has continued to grow globally in each of the ten years since their first introduction in 1996.


'''Transgenic recombinant plants''' are now generally produced in a laboratory by adding one or more [[gene]]s to a plant's [[genome]],and the techniques frequently called [[transformation (genetics)|transformation]].  Transformation is usually acheived using gold particle bombardment or a soil bacterium (''[[Agrobacterium tumefaciens]]'') carrying an engineered plasmid vector, or carrier of selected extra genes.   
Transgenic recombinant plants are now generally produced in a laboratory by adding one or more [[gene]]s to a plant's [[genome]],and the techniques frequently called [[transformation (genetics)|transformation]].  Transformation is usually achieved using gold particle bombardment or a soil bacterium (''[[Agrobacterium tumefaciens]]'') carrying an engineered plasmid vector, or carrier of selected extra genes.   


Transgenic recombinant plants are identified as a class of [[genetically modified organism]](GMO); usually only transgenic plants created by direct DNA manipulation are given much attention in public discussions.
Transgenic recombinant plants are identified as a class of [[genetically modified organism]](GMO); usually only transgenic plants created by direct DNA manipulation are given much attention in public discussions.
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The preliminary step to using ''Agrobacterium'' for plant transformation is to carry out [[genetic engineering]], using [[recombinant DNA]] techniques, to create T-DNA plasmid vectors that carrying the desired foreign DNA. The recombinant T-DNA plasmids are then used to replace the natural plasmids in living ''Agrobacterium'' cells which can then do the job of conjugating with plant callus tissue.
The preliminary step to using ''Agrobacterium'' for plant transformation is to carry out [[genetic engineering]], using [[recombinant DNA]] techniques, to create T-DNA plasmid vectors that carrying the desired foreign DNA. The recombinant T-DNA plasmids are then used to replace the natural plasmids in living ''Agrobacterium'' cells which can then do the job of conjugating with plant callus tissue.


An alternative route to getting foreign DNA into plant cells is called [[biolistics]]. In this methods genetically manipulated DNA is coated onto small (gold) particles and these are fired into plant cells by a small gun-like device.
An alternative route to getting foreign DNA into plant cells is called [[biolistics]]. In this methods genetically manipulated DNA is coated onto small (gold) particles and these are fired into plant cells by a small gun-like device.  In addition, transgenic plants have been created by adding DNA to [[protoplasts]], inducing them to take it up, and then selecting and regenerating plants from those cells.


==Current global picture of modern transgenic crops==
==Current global picture of modern transgenic crops==
{{stub}}
A good source of information is the ISAAA [http://www.isaaa.org/]


==Regulation of transgenic plants==
==Regulation of transgenic plants==
In the [[United States]] the [http://usbiotechreg.nbii.gov Coordinated Framework for Regulation of Biotechnology] governs the regulation of transgenic organisms, including plants.  The three agencies involved are:
In the [[United States of America|United States]] the [http://usbiotechreg.nbii.gov Coordinated Framework for Regulation of Biotechnology] governs the regulation of transgenic organisms, including plants.  The three agencies involved are:


*[[USDA]] [[Animal and Plant Health Inspection Service]] - who state that
*[[USDA]] [[Animal and Plant Health Inspection Service]] - who state that
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==Ecological risks==
==Ecological risks==
The potential impact on nearby ecosystems is one of the greatest concerns associated with transgenic plants but most domesticated plants mate with wild relative a some location where they are grown, and gene flow from domesticated crops (irrespective of whether they transgenic or non-transgenic) can the have potentially harmful consequences of 1. evolution of increased weediness; 2. increased likihood of extinction of wild-relatives. Weediness of hybrids created with domesticated crops is quite common. For instance in California, cultivated rye hybridises with the wild ''Secale montanum'' to produce a weed, and this has led many Californian farmers to abandon rye as a crop. [http://links.jstor.org/sici?sici=0066-4162(1999)30%3C539%3AGFAIFD%3E2.0.CO%3B2-R]
The potential impact on nearby ecosystems is one of the greatest concerns expressed about transgenic plants. Most domesticated plants mate with wild relative a some location where they are grown, and gene flow from domesticated crops (irrespective of whether they transgenic or non-transgenic) can the have potentially harmful consequences <ref>Morris S.H. (2006) EU biotech crop regulations and environmental risk: a case of the emperor's new clothes? Trends Biotechnol. 2006 Nov 17; [Epub ahead of print]</ref>.
 
The main concerns are of 1. evolution of increased weediness; 2. increased likihood of extinction of wild-relatives. There are known instances of unwanted weediness of hybrids created by unintended gene flow from domesticated crops to wild-relatives. For instance in California, cultivated rye hybridises with the wild ''Secale montanum'' to produce a weed, and this has led many Californian farmers to abandon rye as a crop. <ref>[http://links.jstor.org/sici?sici=0066-4162(1999)30%3C539%3AGFAIFD%3E2.0.CO%3B2-R]</ref>


Transgenes (and traits present in domesticated crop created by conventional breeding) have the potential for significant ecological impact if the plants can increase in frequency and persist in natural populations.  This can occur:
Transgenes and other new traits such as mutation to herbicide tolerance present in domesticated crop created by conventional breeding have the potential for significant ecological impact if the plants receiving the trait ccan increase in frequency and persist in natural populations.  This can occur:
* if transgenic plants "escape" from cultivated to uncultivated areas.
* if transgenic plants "escape" from cultivated to uncultivated areas.
* &nbsp;if transgenic plants mate with similar wild plants, the transgene could be incorporated into the offspring.&nbsp;  
* &nbsp;if transgenic plants mate with similar wild plants, the transgene could be incorporated into the offspring.&nbsp;  
Line 176: Line 253:


==References==
==References==
<references/>
<div class="references-small" style="-moz-column-count:2; column-count:2;">
 
<references />
==Further reading==
</div>
*Syvanen, M. and Kado, C. I. Horizontal Gene Transfer. Second Edition. Academic Press 2002.
*Chrispeels, M.J. and Sadova, D.E. Plants, Genes, and Crop Biotechnology. Second Edition. James and Bartlett 2003.
*[http://www.pnas.org/cgi/content/abstract/96/11/5937 Plant genetic resources: What can they contribute toward increased crop productivity? David Hoisington*, Mireille Khairallah, Timothy Reeves, Jean-Marcel Ribaut, Bent Skovmand, Suketoshi Taba, and Marilyn Warburton, Proc. Natl. Acad Sci USA. Vol. 96, Issue 11, 5937-5943, May 25, 1999. (This paper was presented at the National Academy of Sciences colloquium "Plants and Population: Is There Time?" held December 5-6, 1998, at the Arnold and Mabel Beckman Center in Irvine, CA).]
*[http://www.aphis.usda.gov/publications/biotechnology/index.shtml U.S. Department of Agriculture Animal and Plant Health Inspection Service (USDA-APHIS)  Publications Biotechnology.]
*[http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_FS_FedReg_02-06.pdf Biotechnology, Federal Regulation, and the U.S. Department of Agriculture, February 2006, USDA-APHIS Fact Sheet ]
*[http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_CoordFrameBro.pdf Biotechnology Regulatory Services, Coordinated Framework for the Regulation of Biotechnology, USDA-APHIS Outreach Material]
*[http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_QA_biotechandusda.pdf Questions and Answers About Biotechnology and the USDA, August 2006, USDA-APHIS Fact Sheet]
*[http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_FS_pharmaceutical_02-06.pdf Permitting Genetically Engineered Plants That Produce Pharmaceutical Compounds, February 2006, USDA-APHIS Fact Sheet]
 
==See Also==
*[[Plant breeding]]
*[[Food security]]
*[[Transposon]]
*[[Mobile genetic elements]]
*[[Transposons as a genetic tool]]
*[[Genome]]
*[[Arabidopsis thaliana]]
*[[Food security]]
*[[United States Department of Agriculture]]
*[[Foreign Agricultural Service]]
*[[Food and Agriculture Organization]]
*[[Developing country]]
*[[International development]]
*[[International Fund for Agricultural Development]]
*[[Hunger]]
 
==External links==
*[http://www.merid.org/fs-agbiotech/ Food Security and Ag-Biotech News] — balanced news on the debate over transgenic crops
*[http://www.pnas.org/cgi/content/abstract/96/11/5937 Plant genetic resources and transgenics contributions towards increased crop productivity] — Context of transgenics for food security
*[http://www.isb.vt.edu/ Information Systems for Biotechnology (ISB) based at Virginia Tech] — Authoritative information in the form of readable up-to-date articles to support the environmentally responsible use of agricultural biotechnology products, including transgenic plants.
*[http://www.fbae.org/index.htm  Foundation for Biotechnology Awareness and Education]
*[http://www.pgeconomics.co.uk/who.htm  PG Economics] - Reports on agricultural economic benefits.
*[http://www.agbioforum.org/  AgBioForum Journal]- Professional economics papers on crop biotechnology
*[http://www.ifpri.org/ Internation Food Policy Research Institute] The food security context.
*[http://www.truthabouttrade.org/  Truth About Trade]- Forthright support of agricultural technology for farmers and the value of free trade.
*[http://www.agbioworld.org/  AgbioWorld]- Comprehensive scientifically sound resource that supports the use on transgenic crops and provides news from around the world on agriculture.
 
[[Category:genetically modified organisms]]
[[Category:biotechnology]]
[[Category:Agriculture]]
[[Category:CZ Live]]
[[Category:Biology Workgroup]]


[[fa:گیاهان تراریخته]]
[[Category:Suggestion Bot Tag]]
[[it:Piante transgeniche]]
[[fi:Geenimuunneltu elintarvike]]

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See also: horizontal gene transfer in plants

Transgenic plants possess a gene or genes that have been transferred from a different species such as another plant, or a microorganism, or other source. They are created in nature during horizontal gene transfer They can also be created during plant breeding, and especially in recent years through the use of the plant DNA transformation technique in biotechnology. (See Plant breeding, Biotechnology and plant breeding).

Introgression

The most efficient natural route for gene movement between plant species is by cross-pollination to form fertile plant inter-species hybrids. Such hybrids are sometimes new plant species, but they can also form a gene transmission "bridge" in the hybrid-zone between two distinct plant populations, and thus facilitate introgression, which is movement of genes from one distinct species or population to another.

More distant natural transfers

The best documented route for natural formation of transgenic plants is gene transfer between a plant epiphyte (such as [[moss]es]), or a parasitic plant (like dodder) and the host plant it colonizes.

Non-standard fertilization of with more than one pollen grain has also been suggested for transfer of a nuclear genome located gene from Poa grass genus into the distantly related sheep's fescue, Festuca ovina[1].

Another mechanism for horizontal gene transfer is Agrobacterium tumefaciens and similar bacteria that inject DNA into plant cells. Biotechnology laboratories exploit Agrobacterium tumefaciens bacteria to make artificial transgenic plants with small segments of added transgenic DNA inserted in the host cell chromosomes. Others mechanisms may include plant sucking insects, mites, and possibly viruses.

Recent comparative studies of gene content of different genomes provides strong circumstantial evidence that natural horizontal gene transfer does occur in plants at a frequency that is significant over evolutionary time scales. Over the evolutionary time-scales plant mitochondria are a stopping point for genes that may enter the nuclear genome from other species, and can in some cases be very active in inter-species gene-traffic (See Horizontal gene transfer).

The natural DNA transfer of mobile DNA between rice and millet is well documented [2], but genes involved in metabolic process and bacterial genes can also take part in natural genes transfers (See horizontal gene transfer in plants).

Hybrid formation in flowering plants and its role in introgression of genes between species

Cross-pollination between plant species generates interspecies hybrids occurs widely in nature and has been exploited in plant breding for more than 100 years to create artificial transgenic plants ( see Plant breeding).

Hybrids can occur in the intermediate geographical zone between two species and provide a "bridge" for genes to "introgress" (or move) from one species to another[3] [4].

Hybrid formation between two species by pollination joins two sets of chromosomes together, one from each parent, is a common event in flowering plant evolution, and the main way new plant species are formed. Interestingly, in many cases hybrids are formed by adding two copies of each chromosome from each parent, forming an allotetraploid that is reproductively isolated from both parents - and a new species[5].

Wild emmer wheat is an example of a species formed by hybridization between two diploid wild grasses, Triticum urartu (AA) and a wild goatgrass Ae. speltoides (BB) to form hybrid new species with four sets of chomosomes (AABB) which is a tetraploid. Triticale (Triticosecale) is a crop cultivated today mostly for forage and animal feed which is an artificial hybrid between rye and wheat, first bred during the late 19th century.

A surprising number of plants show evidence of being formed by such processes by which chromosome sets are added : bread wheat ( an allohexaploid having three component genomes) , and cotton are two other examples [6].

Gene transfer could occur by wide cross-pollination even if foreign chromosomes are lost. It has been suggested that 'wide crosses' are a possible mechanism of horizontal transfer of mobile DNA in plants, and that these might transfer only mobile DNAs, due to one of the participating sets of chromosomes being lost [7][8]

Natural movements of genes between species by other routes than pollen

See Horizontal gene transfer in plants, Horizontal gene transfer

Transgenic plants and crop improvement

Production of transgenic plants in 'wide-crosses' by plant breeders has been a vital aspect of conventional plant breeding for a century or so. Without it, security of our food supply against losses caused by crop pests such as rusts and mildews would be severely compromised. The first historically recorded interpecies transgenic cereal hybrid was actually between wheat and rye (Wilson, 1876).

Transgenic varieties are frequently created by classical breeders by deliberately and artificially force hybridisation between distinct plant species with the intention of developing disease resistant crop varieties. Classical plant breeders may use use of a number of in vitro techniques such as protoplast fusion, embryo rescue or mutagenisis to generate diversity and produce plants that would not exist in nature (see also Plant breeding, Heterosis, New Rice for Africa). Chromosomal rearrangements and translocations occurring in these crosses help limit the amount of new DNA appearing in the final cultivated variety to a fraction of a chromosome, but still comprise substantial numbers of novel genes introduced into food.

These "classical" techniques (used extensively since about 1930 on) have never been controversial, or been given wide publicity except among professional biologists, and have allowed crop breeders to develop varieties of basic food crop, wheat in particular, which resist devastating plant diseases such as rusts. Hope is one such transgenic wheat variety bred by E. S. McFadden with a transgene from a wild grass. Hope saved American wheat growers from devastating stem rust outbreaks in the 1930s.

Introduction of alien germplasm into common foods has repeatedly achieved novel genetic rearrangements of plant chromosomes, such as insertion of large blocks of rye (Secale) genes into wheat chromosomes ('translocations')[9].

The advent of drug colchicine in the late 1930s helped overcome fertility barriers in inter-specific crosses by stimulating doubling of chromosome numbers per cell, and after 1930 perennial wild-grasses were being frequently hybridized with wheat and other cereals with the aim of transferring disease resistance and perenniality into annual crops. Large-scale practical use of hybrids became well established, leading on to development of numerous Triticosecale (Triticale) varieties and other new transgenic cereal crops.

Important transgenic pathogen and parasite resistance traits carried in current bread wheat varieties are [10]:

Disease resistance to Leaf rust Disease resistance to powdery mildew
Gene Source Gene Source
Lr9 Aegilops umbellulata Pm12 Aegilops speltoides
Lr18 Triticum timopheevi Pm21 Haynaldia villosa
Lr19 Thinopyrum Pm25 T. monococcum
Lr23 T. turgidum
Lr24 Ag. elongatum Disease resistance to wheat streak mosaic virus
Lr25 Secale cereale Wsm1 Ag. elongatum
Lr29 Ag. elongatum
Lr32 T. tauschii Pest resistance
Hessian fly
Disease resistance to stem rust H21 Secale cereale
Sr2 T. turgidum ("Hope" [11]) H23 Secale cereale
Sr22 Triticum monococcum H24 T. tauschii
Sr36 Triticum timopheevii H27 Aegilops ventricosa
Disease resistance to stripe rust Cereal cyst nematode
Yr15 Triticum dicoccoides Cre3 T. tauschii

The intentional creation of transgenic plants by laboratory based recombinant DNA methods is more recent (from the mid-1980s on) and has been a controversial development opposed vigourously by many NGOs, and several governments, particularly within the European Community. In those regions that have allowed farmers to adopt them these transgenic recombinant plants (= biotech crops, modern transgenics) are transforming agricultural productivity, and the area sown to these crops has continued to grow globally in each of the ten years since their first introduction in 1996.

Transgenic recombinant plants are now generally produced in a laboratory by adding one or more genes to a plant's genome,and the techniques frequently called transformation. Transformation is usually achieved using gold particle bombardment or a soil bacterium (Agrobacterium tumefaciens) carrying an engineered plasmid vector, or carrier of selected extra genes.

Transgenic recombinant plants are identified as a class of genetically modified organism(GMO); usually only transgenic plants created by direct DNA manipulation are given much attention in public discussions.

Transgenic plants have been deliberately developed for a variety of reasons: longer shelf life, disease resistance, herbicide resistance, pest resistance, non-biological stress resistances, such as to drought or nitrogen starvation, and nutritional improvement (see Golden rice). The first modern transgenic crop approved for sale in the US, in 1994, was the FlavrSavr tomato, which was intended to have a longer shelf life. The first conventional transgenic cereal created by scientific breeders was actually a hybrid between wheat and rye in 1876 (Wilson, 1876). The first transgenic cereal may have been wheat itself, which is a natural transgenic plant derived from at least three different parenteral species.

Commercial factors, especially high regulatory and research costs, have so far restricted modern transgenic criop varieties to major traded commodity crops, but recently R&D projects to enhance crops that are locally important in developing counties are being pursued, such as insect protected cow-pea for Africa [12], and insect protected Brinjal eggplant for India [13].

Plant transformation with foreign DNA

Modern biology can now be used to manipulate plant genomes and introduce short regions of foreign DNA into a plant by the process of plant transformation. This is the most common way transgenic plants are created in the laboratory.

One way this can be done is by exploiting one of the natural mechanisms for the relatively rare movement of DNA between species. The bacterium Agrobacterium tumefaciens has a natural mechanism called conjugation to inject small segments of DNA (T-DNA) into a plant cell. The T-DNA integrates randomly into the plant chromosomes and once inserted can function as a new gene. In the laboratory this mechanism is exploited to insert desired genes into the cells of plant callus tissue culture, which can then be regenerated into a full plant.

The preliminary step to using Agrobacterium for plant transformation is to carry out genetic engineering, using recombinant DNA techniques, to create T-DNA plasmid vectors that carrying the desired foreign DNA. The recombinant T-DNA plasmids are then used to replace the natural plasmids in living Agrobacterium cells which can then do the job of conjugating with plant callus tissue.

An alternative route to getting foreign DNA into plant cells is called biolistics. In this methods genetically manipulated DNA is coated onto small (gold) particles and these are fired into plant cells by a small gun-like device. In addition, transgenic plants have been created by adding DNA to protoplasts, inducing them to take it up, and then selecting and regenerating plants from those cells.

Current global picture of modern transgenic crops

A good source of information is the ISAAA [5]

Regulation of transgenic plants

In the United States the Coordinated Framework for Regulation of Biotechnology governs the regulation of transgenic organisms, including plants. The three agencies involved are:

The Biotechnology Regulatory Services (BRS) program of the U.S. Department of Agriculture’s (USDA) Animal and Plant Health Inspection Service (APHIS) is responsible

for regulating the introduction (importation, interstate movement, and field release) of genetically engineered (GE) organisms that may pose a plant pest risk. BRS exercises this authority through APHIS regulations in Title 7, Code of Federal Regulations, Part 340 under the Plant Protection Act of 2000.

APHIS protects agriculture and the environment by ensuring that biotechnology is developed and used in a safe manner. Through a strong regulatory framework, BRS ensures the safe and confined introduction of new GE plants with significant safeguards to prevent the accidental release of any GE material.

APHIS has regulated the biotechnology industry since 1987 and has authorized more than 10,000 field tests of GE organisms. In order to emphasize the importance of the program, APHIS established BRS in August 2002 by combining units within the agency that dealt with the regulation of biotechnology. Biotechnology, Federal Regulation, and the U.S. Department of Agriculture, February 2006, USDA-APHIS Fact Sheet

  • EPA - evaluates potential environmental impacts, especially for genes which produce pesticides
  • DHHS, Food and Drug Administration (FDA) - evaluates human health risk if the plant is intended for human consumption

Ecological risks

The potential impact on nearby ecosystems is one of the greatest concerns expressed about transgenic plants. Most domesticated plants mate with wild relative a some location where they are grown, and gene flow from domesticated crops (irrespective of whether they transgenic or non-transgenic) can the have potentially harmful consequences [14].

The main concerns are of 1. evolution of increased weediness; 2. increased likihood of extinction of wild-relatives. There are known instances of unwanted weediness of hybrids created by unintended gene flow from domesticated crops to wild-relatives. For instance in California, cultivated rye hybridises with the wild Secale montanum to produce a weed, and this has led many Californian farmers to abandon rye as a crop. [15]

Transgenes and other new traits such as mutation to herbicide tolerance present in domesticated crop created by conventional breeding have the potential for significant ecological impact if the plants receiving the trait ccan increase in frequency and persist in natural populations. This can occur:

  • if transgenic plants "escape" from cultivated to uncultivated areas.
  •  if transgenic plants mate with similar wild plants, the transgene could be incorporated into the offspring. 
  • if these new transgene plants become weedy or invasive, which could reduce
  • if the transgenic crop trait confers a selective advantage in natural environments

Gene flow may affect biodiversity and might affect entire ecosystems.

Pollen flow from conventional crop plants to native species also poses gene-flow derived ecological risks, as crop plants are not selected to have optimal selective advantages in natural environments, and farm fields are different to natural ecosystems. Conventional varieties also posses new traits such as pest resistance that have been deliberately transferred into the crop variety from other species.

There are at least three possible avenues of hybridization leading to escape of a transgene:

  1. Hybridization with non-transgenic crop plants of the same species and variety.
  2. Hybridization with wild plants of the same species.
  3. Hybridization with wild plants of closely related species, usually of the same genus.

However, there are a number of factors which must be present for hybrids to be created.

  • The transgenic plants must be close enough to the wild species for the pollen to reach the wild plants.
  • The wild and transgenic plants must flower at the same time.
  • The wild and transgenic plants must be genetically compatible.
  • The hybrid offspring must be viable, and fertile.
  • The hybrid offspring must carry the transgene.

Studies suggest that a possible escape route for transgenic plants will be through hybridization with wild plants of related species.

  1. It is known that some crop plants have been found to hybridize with wild counterparts.
  2. It is understood, as a basic part of population genetics, that the spread of a transgene in a wild population will be directly related to the fitness effects of the gene in addition to the rate of influx of the gene to the population.  Advantageous genes will spread rapidly, neutral genes will spread with genetic drift, and disadvantageous genes will only spread if there is a constant influx.
  3. The ecological effects of transgenes are not known, but it is generally accepted that only genes which improve fitness in relation to abiotic factors would give hybrid plants sufficient advantages to become weedy or invasive.  Abiotic factors are parts of the ecosystem which are not alive, such as climate, salt and mineral content, and temperature.

References

  1. ↑ Ghatnekar L, Jaarola M, Bengtsson BO.(2006) The introgression of a functional nuclear gene from Poa to Festuca ovina.Proc Biol Sci. 2006 Feb 22;273(1585):395-9.
  2. ↑ Jumping Genes Cross Plant Species Boundaries. PLoS Biol 4(1): e35 DOI:10.1371/journal.pbio.0040035 Published: December 20, 2005
  3. ↑ Rieseberg, L.H. and Wendel, J. (1993). Introgression and its consequences in plants. In Hybrid Zones and the Evolutionary Process. (ed. J. Harrison) p 70-109, Oxford University Press, New York.
  4. ↑ Rieseberg, L.H. and Ellstrand, N.C. (1993) What can molecular and morphological markers tell us about plant hybridization/ Critical Reviews of Plant Science. 12 p213-241.
  5. ↑ Ramsey, J. and Schemske, D.W. (1998) Pathways, mechanisms, and rates of polyploid formation in flowering plants. Annual Review of Ecology and Systematics. 29, 467-501.
  6. ↑ J. A. Udall and J. F. Wendel (2006) Polyploidy and Crop Improvement. Crop Sci. 46, S-3-S-14
  7. ↑ Ananiev EV, Riera-Lizarazu O, Rines HW, Phillips RL (1997) Oat maize chromosome addition lines: a new system for mapping the maize genome. Proc Natl Acad Sci USA 94: 3524–3528.
  8. ↑ Bennetzen, J. L., (2000) Transposable element contributions to plant gene and genome evolution. Plant Molecular Biology 42: 251–269, 2000.
  9. ↑ [1]
  10. ↑ Plant genetic resources: What can they contribute toward increased crop productivity? Hoisington, D. and others (1999) Proc. Natl. Acad Sci USA. Vol. 96, Issue 11, 5937-5943, May 25, 1999. (This paper was presented at the National Academy of Sciences colloquium "Plants and Population: Is There Time?" held December 5-6, 1998, at the Arnold and Mabel Beckman Center in Irvine, CA).
  11. ↑ McFadden, E. S. (1930) J. Am. Soc. Agron. 22, 1020-1031.
  12. ↑ [2]
  13. ↑ [3]
  14. ↑ Morris S.H. (2006) EU biotech crop regulations and environmental risk: a case of the emperor's new clothes? Trends Biotechnol. 2006 Nov 17; [Epub ahead of print]
  15. ↑ [4]
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