Can feral weeds evolve from cultivated radish (Raphanus sativus, Brassicaceae)?

Cultivated plants that cannot survive on their own often have maladaptive domestication traits. Unharvested crop seeds may generate feral populations, at times causing serious weed problems, but little is known about the evolution of ferality. We explored the potential for cultivated radish, Raphanus sativus, to become feral, given that closely related taxa (e.g., R. raphanistrum and crop-wild hybrids) are well-documented weeds. First, we measured the population growth of five experimental, cultivated, self-seeding radish populations in Michigan, USA, for three generations. Three late-flowering populations went extinct, and two others apparently hybridized with local R. raphanistrum. A common garden experiment showed that the two surviving populations had earlier flowering, smaller root diameters, and greater individual fecundity than did nonhybridized populations. We also used artificial selection to measure the evolutionary potential for earlier flowering. After two generations of strong selection, two of three lineages flowered earlier and produced more seeds than control lineages, but insufficient genetic variation prevented dramatic evolution of crop phenotypes. In summary, it seems unlikely that radishes could spontaneously become feral in our study area without gene flow from R. raphanistrum. Applying these approaches to other cultivated species may provide a better understanding of mechanisms promoting the evolution of feral weeds.     

The Biology and Ecology of Feral Alfalfa (Medicago sativa L.) and Its Implications for Novel Trait Confinement in North America

Alfalfa (Medicago sativa) is an important forage crop worldwide. Apart from cultivated fields, alfalfa is also found along roadsides and in natural and semi-natural habitats. However, little information is available on the establishment capabilities of alfalfa in noncultivated areas and the potential of these founding populations to become feral. Some crop species have not lost all their wild characteristics during the domestication process and with several inherent traits favoring weediness, alfalfa could be one among those that can become feral. There is great interest in the feral potential of alfalfa, particularly due to the concerns that feral plants could act as genetic bridges and facilitate novel trait movement at the landscape level. Alfalfa is the first perennial, insect-pollinated crop to be genetically engineered and approved for unconfined release into the environment. This review investigates and compiles information in the literature that reveals the life history components that can influence ferality in alfalfa. Characteristics that can contribute to ferality in alfalfa include high genetic diversity, perenniality, quick regrowth potential, persistence, symbiotic nitrogen fixation, deep tap root system, drought and cold tolerance, and seed dormancy. With these traits, alfalfa is equipped to invade and dominate unmanaged habitats. Feral alfalfa populations can and will act as bridges for long-distance gene flow and facilitate the adventitious presence of novel traits in the environment. As such, feral populations will become a potential barrier for achieving coexistence of transgenic and nontransgenic alfalfa fields. Implications of ferality, including gene flow and hybridization with compatible wild relatives are also discussed in detail. This review serves as a resource for environmental risk assessment for the release of alfalfa containing novel traits.     

Crop Ferality and Volunteerism

Volunteers and weedy crop (feral) plants have been with us eversince the domestication of crops. Whilst we know a great dealabout their biology and management as weeds, little scientificattention has been paid to their evolution, and understandingtheir role in the domestication and de-domestication of crops.With the commercialization of transgenic crops, there is a greaterurgency to understand these processes and establish whethervolunteer (crop plants that germinate in subsequent years aftera crop has been harvested) and feral (plants derived in fullor in part     

Introgression from cultivated rice influences genetic differentiation of weedy rice populations at a local spatial scale

Hybridization and introgression can play an important role in genetic differentiation and adaptive evolution of plant species. For example, a conspecific feral species may frequently acquire new alleles from its coexisting crops via introgression. However, little is known about this process. We analyzed 24 weedy rice (Oryza sativa f. spontanea) populations and their coexisting rice cultivars from northern Italy to study their genetic differentiation, outcrossing, and introgression based on microsatellite polymorphisms. A total of 576 maternal plants representing 24 weedy populations were used to estimate their genetic differentiation, and 5,395 progeny (seedlings) derived from 299 families of 15 selected populations were included to measure outcrossing rates. Considerable genetic differentiation (F _st = 0.26) was detected among weedy rice populations, although the differentiation was not associated with the spatial pattern of the populations. Private alleles (28%) were identified in most populations that exhibited a multiple cluster assignments, indicating stronger genetic affinities of some weedy populations. Outcrossing rates were greatly variable and positively correlated (R ² = 0.34, P = 0.02) with the private alleles of the corresponding populations. Paternity analysis suggested that ~15% of paternal specific alleles, a considerable portion of which was found to be crop-specific, were acquired from the introgression of the coexisting rice cultivars. Frequent allelic introgression into weedy populations resulting from outcrossing with nearby cultivars determines the private alleles of local feral populations, possibly leading to their genetic differentiation. Introgression from a crop may play an important role in the adaptive evolution of feral populations.     

Community response to transgenic plant release: predictions from British experience of invasive plants and feral crop plants

The ten-ten rule can be used to predict the community consequences of releasing GMOs. This, combined with data on the feral state of British crop plants, predicts that almost all GMOs will become at least casual, that more than 10% will establish, and about 10% of those will become pests. Which constructs will fall into which category cannot be predicted from their characters. In particular, Baker characters have no predictive value for weediness. The fourteen most troublesome invasive plant pests show a remarkably diverse set of characters. GMOs, being mainly derived from crop plants, and in some cases with genes that are likely to enhance survival, can be expected to have an appreciable effect on nonagricultural ecosystems, once a range of different constructs have been released. Familiarity is unlikely to be an effective defence against new ecological effects.     

Transcriptional profile and differential fitness in a specialist milkweed insect across host plants varying in toxicity

Interactions between plants and herbivorous insects have been models for theories of specialization and co‐evolution for over a century. Phytochemicals govern many aspects of these interactions and have fostered the evolution of adaptations by insects to tolerate or even specialize on plant defensive chemistry. While genomic approaches are providing new insights into the genes and mechanisms insect specialists employ to tolerate plant secondary metabolites, open questions remain about the evolution and conservation of insect counterdefences, how insects respond to the diversity defences mounted by their host plants, and the costs and benefits of resistance and tolerance to plant defences in natural ecological communities. Using a milkweed‐specialist aphid (Aphis nerii) model, we test the effects of host plant species with increased toxicity, likely driven primarily by increased secondary metabolites, on aphid life history traits and whole‐body gene expression. We show that more toxic plant species have a negative effect on aphid development and lifetime fecundity. When feeding on more toxic host plants with higher levels of secondary metabolites, aphids regulate a narrow, targeted set of genes, including those involved in canonical detoxification processes (e.g., cytochrome P450s, hydrolases, UDP‐glucuronosyltransferases and ABC transporters). These results indicate that A. nerii marshal a variety of metabolic detoxification mechanisms to circumvent milkweed toxicity and facilitate host plant specialization, yet, despite these detoxification mechanisms, aphids experience reduced fitness when feeding on more toxic host plants. Disentangling how specialist insects respond to challenging host plants is a pivotal step in understanding the evolution of specialized diet breadths.     

More than multiple introductions: Multiple taxa contribute to the genesis of the invasive California’s wild artichoke thistle

The history of some invasive species is so complex that their origins can be difficult to determine. One example of such invasive species is the California invasive known as “wild artichoke thistle” (Cynara cardunculus var. sylvestris), found in natural and disturbed ecosystems. Wild artichoke thistle is a Mediterranean native and the progenitor of two domesticated horticultural taxa, artichoke and cardoon. Different hypotheses regarding the origins of California plants have included introductions by 19th century Italian immigrants and the de-domestication (evolutionary reversion to wild-type morphology) of feral (escaped, free-living) cultivars. Using microsatellite markers, we compared the genetic constitutions of 12 artichoke thistle populations in California with possible progenitor populations: 17 Spanish and Italian wild populations and eight different artichoke and cardoon cultivars. Each California population was compared with its putative progenitors using STRUCTURE analysis. Our results suggest that California's artichoke thistle populations are polyphyletic. Surprisingly, two-thirds of California's populations closely matched populations from the Iberian Peninsula. Three populations matched domesticated artichoke. One population appears to have wild and cultivar hybrid ancestry. Alleles specific to Italian populations were found at low frequencies in some California plants, suggesting that Italian wild plants may have been in California, but have left a trivial genetic legacy. Given that the de-domesticated plants in this study appear to be as invasive as the wild taxon, we conclude with a discussion of the role that ferality and de-domestication may have in plant invasions.     

Association between feral pig disturbance and the composition of some alien plant assemblages in Hawaii Volcanoes National Park

Association analysis was used to assess relationships among 25 important alien plant species and their association with feral pig rooting in Hawaii Volcanoes National Park, Hawaii, U.S.A. Results of the association analysis were summarized by means of a simplified, rank-based, polar ordination which yielded three subjective species assemblages. One group was characterized by an association with the endpoint species, Ehrharta stipoides. A second group consisted of species associated with the other endpoint species, Andropogon virginicus. The third group comprised a diverse assemblage of 14 species in the middle of the ordination.Comparison of ordination scores with each species's association with pig-induced soil disturbance revealed that members of the Ehrharta group were strongly positively associated with pig activity, whereas members of the Andropogon group were generally negatively associated. The third group showed no association with pig-induced soil disturbance.These results suggest a strong relationship between feral pig activity and the composition of the alien portion of the plant community. Analysis of the ecologies of both plants and pigs suggests that some species may both encourage pig activity and benefit from it. Likewise, other alien plants appear to neither require nor benefit from pig-induced soil disturbance. While pigs appear to play an important role in the organization of these communities, their removal may have a negligible impact on the success of many of the common weeds in the area. Nomenclature: Wagner, W. L., Herbst, D. R., & Sohmer, S. H. 1990. Manual of the Flowering Plants of Hawai'i. University of Hawaii Press/Bishop Museum Press, Honolulu.     

Occurrence of Transgenic Feral Alfalfa (Medicago sativa subsp. sativa L.) in Alfalfa Seed Production Areas in the United States

The potential environmental risks of transgene exposure are not clear for alfalfa (Medicago sativa subsp. sativa), a perennial crop that is cross-pollinated by insects. We gathered data on feral alfalfa in major alfalfa seed-production areas in the western United States to (1) evaluate evidence that feral transgenic plants spread transgenes and (2) determine environmental and agricultural production factors influencing the location of feral alfalfa, especially transgenic plants. Road verges in Fresno, California; Canyon, Idaho; and Walla Walla, Washington were surveyed in 2011 and 2012 for feral plants, and samples were tested for the CP4 EPSPS protein that conveys resistance to glyphosate. Of 4580 sites surveyed, feral plants were observed at 404 sites. Twenty-seven percent of these sites had transgenic plants. The frequency of sites having transgenic feral plants varied among our study areas. Transgenic plants were found in 32.7%, 21.4.7% and 8.3% of feral plant sites in Fresno, Canyon and Walla Walla, respectively. Spatial analysis suggested that feral populations started independently and tended to cluster in seed and hay production areas, places where seed tended to drop. Significant but low spatial auto correlation suggested that in some instances, plants colonized nearby locations. Neighboring feral plants were frequently within pollinator foraging range; however, further research is needed to confirm transgene flow. Locations of feral plant clusters were not well predicted by environmental and production variables. However, the likelihood of seed spillage during production and transport had predictive value in explaining the occurrence of transgenic feral populations. Our study confirms that genetically engineered alfalfa has dispersed into the environment, and suggests that minimizing seed spillage and eradicating feral alfalfa along road sides would be effective strategies to minimize transgene dispersal.     

Determining the major Bt refuge crops for cotton bollworm in North China

Evaluation of the effectiveness of refuge strategies involved in cotton bollworm Bt resistance management would be aided by technologies that allow monitoring and quantification of key factors that affect the process under field conditions. We hypothesized that characterization of stable carbon and nitrogen isotopes in adult bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) moths may aid in determining the larval host that they developed upon. We found moths reared from larvae fed on peanut, soybean or cotton, respectively, could be differentiated using isotopic analyses that also corresponded to their respective larval host origins. These techniques were then used to classify feral second‐generation bollworm moths caught in Bt cotton (Gossypium hirsutum) fields into different populations based on their isotopic signatures. In 2006–2007 feral moths captured in Bt cotton fields predominantly correlated with the peanut (Arachis hypogea) having served as their larval host, indicating this is the most important refuge crop for Bt‐susceptible bollworm individuals (providing 58%?64% individuals) during independent moth peaks for the second generation in North China. The remaining feral moths correlated with soybean (Glycine max) (0?10%); other C₃ plant (20%?22%) and non‐C₃ plant (12%?14%) host types also provided some Bt‐sensitive moths. Field observations showed that peanut constitutes the primary refuge crop contributing to sustaining Bt‐susceptible moths dispersing into cotton in North China. These results suggest that peanut may be a more effective refuge to sustain Bt‐susceptible bollworm individuals and reduce the risk of development of a Bt‐resistant biotype.     

Integrating genetic information into plant breeding programmes: how will we produce varieties from molecular variation,using bioinformatics?

The development of new varieties of crop plants is ongoing for plant breeders and progress since the Green Revolution has been steady, if not dramatic. With the recent sequencing of Arabidopsis thaliana and of rice the development of both physical and informational resources has entered a new phase. This paper examines the state of plant bioinformatics as it is now and as it is likely to develop in the future. It also looks rather further forward to what crop scientists might want from bioinformatics, before examining the likely physiological targets for sustainability traits and the prospects for their improvement in wheat. Wheat is taken as the focus crop because it is potentially one of the most dif. Cult to work with in molecular terms, both because of its large hexaploid genome size and because of its considerable genetic distance from the most information rich plant species, Arabidopsis. Finally, we examine how these tools might be used to track down the underlying genes controlling sustainability traits and how these may then be exploited in plant breeding programmes using marker‐ assisted selection.     

TISSUE CULTURE IN THE PRODUCTION OF NOVEL DISEASE-RESISTANT CROP PLANTS

1. Plant tissue cultures form the basis of a number of techniques which have been developed to effect genetic changes in plants. Progress is being made in the application of these techniques in breeding new, disease-resistant cultivars. 2. It is possible to induce and select for mutants among populations of cultured plant cells. Novel disease-resistant plants of a small number of species have been regenerated from cells selected in culture for their resistance to toxins produced by pathogens, both with and without prior exposure to mutagens. It is not known whether such procedures are widely applicable, and the nature of the genetic changes involved has not yet been determined. 3. The tissues of plant species which are propagated vegetatively are normally genetic mosaics with regard to many characteristics, including resistance to disease. Thus, some of the plants regenerated from cultured cells of such species are more resistant to pathogens than the parent plants. Novel plants produced in this way are already being used in some breeding programmes. 4. Many attempts have been made to modify the genomes of cultured plant cells by means of exogenous nucleic acids. The evidence for integration and replication of this genetic material is equivocal. The technique, therefore, offers no immediate prospects for the development of novel disease-resistant plants, but may be important in the long term as methods are perfected for using plasmids and other agents as carriers of useful genes. 5. Steady advances are being made in producing somatic hybrids of crop plants by fusion of isolated protoplasts. In the long term it may be possible to use protoplast fusion to transfer desirable disease-resistance traits between related species which cannot be hybridized by conventional breeding methods. 6. The culture of excised embryos may be used to grow interspecific and inter-generic hybrid plants in cases where incompatibility occurs after normal fertilization. The technique is already being used by breeders in the production of disease-resistant hybrids of crop species. 7. It is concluded that tissue culture has a limited but useful role to play in the development of novel disease-resistant crop plants.     

Genetic and molecular basis of grain size and grain number and its relevance to grain productivity in higher plants.

Grain size and grain number constitute 2 important components of grain yield. In particular, the grain size also influences the end-use quality (e.g., flour yield and protein content) and attracts consumer preference. These 2 traits are also the components of the domestication syndrome of crop plants. A number of important studies have recently been conducted to understand the genetic and molecular basis of these 2 important yield-contributing traits. Information generated from these studies was collected and synthesized for the benefit of plant biologists, particularly plant breeders. In the present article, this information is briefly reviewed and the prospects of using this information for improvement of grain productivity in crop plants are discussed.     

Cryopreserved storage of clonal germplasm in the USDA National Plant Germplasm System

The US Department of Agriculture-Agricultural Research Service (USDA-ARS), National Plant Germplasm System (NPGS) plant collections are a critical source of genetic diversity for breeding and selection of improved crops, including vegetatively propagated plants. Information on these collections is readily accessible to breeders and researchers on the internet from the Germplasm Resources Information Network (GRIN). The clonal collections are at risk for loss due in part to their genetic diversity that makes growing them in one location a challenge, but also because it is difficult to have duplicate collections without incurring great expense. The development of cryopreservation techniques during the last two decades provides a low maintenance form of security backup for these collections. National plant collections for vegetatively propagated crop plants and their wild relatives are maintained by the USDA-ARS, NPGS at 15 sites across the country. These sites include various combinations of field, greenhouse, screenhouse, and in vitro collections. Cryopreserved backup collections in liquid nitrogen storage were instituted in the 1990s, increased greatly in the 2000s with the advent of new techniques, and are continuing today. Collections of dormant buds of temperate trees, shoot tips of in vitro cultures of many crops, and embryonic axes of some large seeded or recalcitrant seeded plants are all part of the clonal backup storage system.     

Membrane transporters for nitrogen, phosphate and potassium uptake in plants

Nitrogen, phosphorous and potassium are essential nutrients for plant growth and development. However, their contents in soils are limited so that crop production needs to invest a lot for fertilizer supply. To explore the genetic potentialities of crops (or plants) for their nutrient utilization efficiency has been an important research task for many years. In fact, a number of evidences have revealed that plants, during their evolution, have developed many morphological, physiological,biochemical and molecular adaptation mechanisms for acquiring nitrate, phosphate and potassium under stress conditions.Recent discoveries of many transporters and channels for nitrate, phosphate and potassium up take have opened upopportunities to study the molecular regulatory mechanisms for acquisition of these nutrients. This review aims to briefly discuss the genes and gene families for these transporters and channels. In addition, the functions and regulation of some important transporters and channels are particularly emphasized.     

Illegal gene flow from transgenic creeping bentgrass: the saga continues

Ecologists have paid close attention to environmental effects that fitness‐enhancing transgenes might have following crop‐to‐wild gene flow (e.g. Snow et al. 2003 ). For some crops, gene flow also can lead to legal problems, especially when government agencies have not approved transgenic events for unrestricted environmental release. Creeping bentgrass (Agrostis stolonifera), a common turfgrass used in golf courses, is the focus of both areas of concern. In 2002, prior to expected deregulation (still pending), The Scotts Company planted creeping bentgrass with transgenic resistance to the herbicide glyphosate, also known as RoundUp^®, on 162 ha in a designated control area in central Oregon ( Fig. 1 ). Despite efforts to restrict gene flow, wind‐dispersed pollen carried transgenes to florets of local A. stolonifera and A. gigantea as far as 14 km away, and to sentinel plants placed as far as 21 km away ( Watrud et al. 2004 ). Then, in August 2003, a strong wind event moved transgenic seeds from windrows of cut bentgrass into nearby areas. The company’s efforts to kill all transgenic survivors in the area failed: feral glyphosate‐resistant populations of A. stolonifera were found by Reichman et al. (2006) , and 62% of 585 bentgrass plants had the telltale CP4 EPSPS transgene in 2006 ( Zapiola et al. 2008 ; Fig. 2 ). Now, in this issue, the story gets even more interesting as Zapiola & Mallory‐Smith (2012) describe a transgenic, intergeneric hybrid produced on a feral, transgenic creeping bentgrass plant that received pollen from Polypogon monspeliensis (rabbitfoot grass). Their finding raises a host of new questions about the prevalence and fitness of intergeneric hybrids, as well as how to evaluate the full extent of gene flow from transgenic crops.

Figure 1 Open in figure viewer PowerPoint Creeping bentgrass and rabbitfoot grass occur along canals and irrigation ditches within the designated control area near Madras, Oregon. Photo courtesy of M. L. Zapiola.     

Bioengineering of crop plants and resistant biotype evolution in insects: Counteracting coevolution

The use, as opposed to the procurement, of transgenic crop plants is discussed in this paper. Transgenic crop plants must not be used until appropriate strategies for their use have been designed and not before crop plants with a variety of insect defenses have been developed. The use of a crop plant with a single defense will pose as strong a selection pressure as the use of a single synthetic insecticide, since insect herbivores are able to evolve effective counter-defenses. The defenses of insects in natural plant-insect associations and with regard to synthetic insecticides are described to demonstrate that there is nothing unique about insecticide resistance. It is the inevitable alternative to local extinction in response to a persistent and predictable selection pressure. Plants counteract insect defensive evolution by keeping the selection pressure as variable as possible. This leads to the conclusion that the best use of biotechnology in crop protection is to reintroduce chemical diversity into crop plants.     

Morphological and phenological comparisons of two Hopi maize varieties conserved in situ and ex situ

Over the last twenty-five years, crop genetic resources (CGR) have been preserved in genebanks around the world for use by formal plant breeders. Recently conservation of folk crop varieties for direct use by the farmer-breeders of traditional agricultural communities has been suggested as another purpose for CGR conservation. While both in and ex situ CGR conservation programs have been proposed to meet the needs of formal plant breeders and farming communities, the needs and goals of the two groups are different. Formal breeders seek maximum allelic diversity while farmer-breeders are interested in both diversity and population structure that provide local adaptation. Based on the morphological and phenological data analyzed for this study of two Hopi maize varieties conserved in and ex situ, it appears that both genetic shift and genetic drift have occurred ex situ, and that populations conserved ex situ are different from those maintained in situ. These findings suggest that CGR conservation strategies must be re-evaluated in light of the specific conservation goals that are sought.