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No description available.

<p> Fruit size is a polygenic trait controlled by multiple genomic regions each with small effect. The complex nature of fruit size regulation makes it challenging to dissect individual genes responsible for phenotypic variation. Though recent advances in high-throughput genome sequencing technology in conjunction with improved statistical and computational methods empowered science to explicitly understand the genetic basis underlying multiple fruit quality traits, much of the work that has been done through classical quantitative trait loci (QTL) approach resulted in reduced resolution and instability when evaluating in different genetic backgrounds and different environments. To increase the precision and improve the stability of QTL analyses and to identify genes controlling fruit size, we performed a set of multiple quantitative and molecular genetic analyses to elucidate the underlying genetic architecture of fruit mass. A total of nine genomic regions associated with fruit mass were identified, two of which are novel to this study; markers Md14_26050918 and Md14_26050904. Detected QTLs explained ~ 42% of the total genetic variation of which ~ 20% is explained by the two novel QTLs. Regions responsible for fruit mass variation appear to be under strong additive and epistatic genetic control. These regions exhibited high stability across-family as well as across-years and showed accurate genomic prediction across-family. Additionally, we identified the apple gene family of putative fw2.2 orthologs, naming them Cell Number Regulators (CNRs) genes (MdCNRs). Three CNRs (MdCNR1-3) showed increased expression at early fruit growth in small-fruited crabapple, associating with reduced relative cell production rate (RCPR), suggesting that alteration in cell number that leads to a subsequent reduction in fruit size is probably due to reduced cell division most likely due to changes in CNRs regulation. Furthermore, our study revealed that reduced fruit size is partially due to the shortened cell expansion period after which cell expansion ceases in the small-fruited crabapple species. Together, these data will advance our understanding of dissecting fruit mass genetic architecture and have high potential to be deployed for marker-assisted selection and further breeding approaches. </p>

[Truncated abstract] In order to expand the gene pool of canola-quality rapeseed (Brassica napus) reciprocal interspecific crosses were made between B. napus cv. Mystic and near canola-quality B. juncea breeding line JN29. F1 progeny from these crosses were used to make backcrosses to both parents in all possible combinations and directions, and were selfed to form F2-derived lines. The highest frequencies of viable F2 and BC1 progeny were obtained when B. napus was the maternal parent of the interspecific hybrid. BC1 and F2 progeny (and subsequent generations) were grown under field conditions to identify agronomic improvements over the parents. Transgressive segregation was observed in F2 and BC1 and in subsequent generations for agronomic traits (seed yield under high or low rainfall conditions, plant biomass, harvest index, height, branching and days to anthesis) and seed quality traits (oil, protein, glucosinolates, oleic acid). The majority of progeny conformed to B. napus morphology, and a minority segregated to B. juncea morphology in subsequent generations. Some of the B. juncea morphotypes had lower glucosinolates and higher oleic acid than the parent JN29, with no detectable erucic acid, and thereby conformed to canola quality. Methods were developed for tracing B-genome in interspecific progeny. A repetitive DNA sequence pBNBH35 from B. nigra (genome BB, 2n = 16) was used to identify B-genome chromosomes and introgressions in interspecific progeny. Specific primers were designed for pBNBH35 in order to amplify the repetitive sequence by PCR. A cloned sub-fragment of 329 bp was confirmed by sequencing as part of pBNBH35. PCR and hybridisation techniques were used on an array of Brassica species to confirm that the pBNBH35 subfragment was Brassica B-genome specific. Fluorescence in situ hybridisation (FISH) in B nigra, B. juncea (AABB, 2n=36) and B. napus (AACC, 2n=38) showed that the pBNBH35 sub-fragment was present on all eight Brassica Bgenome chromosomes and absent from A- and C-genome chromosomes. The pBNBH35 repeat was localised to the centromeric region of each B-genome chromosome. FISH clearly distinguished the B-genome chromosomes from the A-genome chromosomes in the amphidiploid species B. juncea. This is the first known report of a B-genome repetitive marker that is present on all Brassica Bgenome chromosomes. ... The results suggest that novel B. napus genotypes have been generated containing introgressions of B-genome chromatin from B. juncea chromosomes. B. juncea morphology occurred in interspecific progeny with a chromosome complement similar to B. napus (2n = 38) and without the entire Bgenome present. It also is highly likely that recombination has occurred between the A-genome of the two Brassica species. This research has demonstrated that the secondary gene pool of B. napus may be accessed by selfing interspecific hybrids, and without sacrificing canola quality, if the B. juncea parent is near canola-quality. Interspecific progeny may be screened to enhance the proportion with B-genome positive signals. Some progeny with B. junceatype morphology had improved seed quality over the JN29 parent.

CAPES / Os pêssegos e nectarinas são muito apreciados pelos consumidores, mas são frutos climatéricos, tendo disponibilidade no mercado curto espaço de tempo. Dessa forma, é necessário investir na obtenção de genótipos com frutos de qualidade e menor perecibilidade ou que apresentem menos distúrbios fisiológicos após armazenamento. Assim, os objetivos deste trabalho foram i) avaliar a divergência genética entre 40 acessos de pessegueiros e nectarineiras com base na qualidade pós-colheita, selecionando-se possíveis genitores; ii) verificar a susceptibilidade a danos por frio em pêssegos e nectarinas após armazenamento refrigerado; iii) avaliar divergência de pêssegos e nectarinas com base na susceptibilidade aos danos por frio e selecionar acessos superiores; iv) realizar estudo das correlações entre qualidade e susceptibilidade a danos por frio de pêssegos e nectarinas; v) selecionar genitores com a combinação de menor susceptibilidade aos danos por frio após armazenamento e maior qualidade dos frutos. O estudo foi conduzido na EEAD-CSIC, Zaragoza-Espanha, durante o ciclo produtivo 2013/2014. Um total de 40 acessos de pêssegos e nectarinas da coleção de germoplasma foram avaliados. Características de qualidade como, firmeza de polpa, teor de sólidos solúveis totais, acidez titulável, pH, índice de maturação e parâmetros de coloração do mesocarpo foram determinadas. Os frutos foram submetidos ao armazenamento refrigerado de 0º e 5ºC, com umidade relativa média de 95%. Após 14 e 28 dias foram avaliados os sintomas de lanosidade através da presença de polpa farinhenta, granulosa, coriácea, alterações de cor na polpa, como escurecimento e avermelhamento interno e, ausência de sabor. Como critério de seleção adotou-se 20% dos genótipos que apresentaram maior frequência de superioridade quanto às características de qualidade, de susceptibilidade aos danos por frio e combinando ambas. Para as característica de qualidade apresentaram maior divergência os acessos ‘Queen Giant’, ‘Sudanel Blanco’ e ‘Borracho de Jarque’. E, com base na qualidade foram selecionados os oito genótipos ‘Andross’, ‘San Jaime’, ‘San Lorenzo’, ‘Borracho de Jarque’, ‘Sudanell 1’, ‘Carson’, ‘Baby Gold 6’ e ‘Stanford’. Todos os 40 acessos apresentaram susceptibilidade a um ou mais sintomas causados pelo armazenamento refrigerado, quando mantidos por 28 dias, independentemente da temperatura. Para 14 dias, os acessos ‘Baby Gold 6’, ‘Flavortop’ e ‘Queen Giant’ não apresentaram nenhuma desordem fisiológica causada pelo frio. De maneira geral, a temperatura de 0 ºC favoreceu para conservação pós-colheita dos frutos, causando menor incidência e severidade dos sintomas oriundos do armazenamento refrigerado. O armazenamento por 14 dias contribuiu para menor incidência de danos sobre os acessos estudados. Para os 14 dias, em ambas as temperaturas, observou-se divergência para os acessos ‘Queen Giant’, ‘Sudanell Blanco’, ‘Baby Gold 6’, ‘GF3’,‘Baby Gold 8’, ‘Campiel’ e ‘Campiel Rojo’. Para 28 dias a 5 ºC, ‘Queen Giant’, ‘Big Top’, ‘Flavortop’ e ‘Redhaven’ apresentaram-se divergentes. Com base na suscetibilidade a danos por frio a 0 ºC, foram selecionados os oito acessos, ‘Queen Giant’, ‘Keimoes’, ‘Flavortop’, ‘Big Top’, ‘Redhaven’, ‘Sudanell 3’, ‘Bonet I’ e ‘Carson’. Os parâmetros de qualidade índice de maturação, sólidos solúveis, firmeza de polpa e acidez total apresentaram correlações entre si. Os mesmos, correlacionaram-se com polpa granulosa e escurecimento interno, indicando que frutos com maior amadurecimento tendem a desenvolver mais estes sintomas. Os distúrbios escurecimento interno, polpa farinhenta, polpa granulosa e ausência de sabor foram correlacionados em ambos os tempos e temperaturas, corroborando que esses sintomas são os principais distúrbios causados pelo armazenamento refrigerado. Combinando as características de qualidade e suscetibilidade aos danos por frio foram selecionados os genótipos ‘Baby Gold 6’, ‘Sarell’, ‘Keimoes’, ‘GF3’ ‘San Jaime’, ‘Big Top’, ‘Sudanell 1’, ‘Carson’, ‘Baby Gold 8’, e ‘San Lorenzo’. / The peaches and nectarines are highly appreciated by consumer, but it is climacteric fruits, with availability in the market in small time. It is necessary to invest to obtain genotypes with fruit quality and small perishability or that it presente less physiological disorders after storage. The aims of this work were i) to evaluate the genetic divergence among 40 peach and nectarine trees genotypes based on postharvest quality and select posible parents; ii) to evaluate the susceptibility to chilling injury in peaches and nectarines after cold storage; iii) to evaluate divergence of peaches and nectarines on the basis in the susceptibility for chiling injury and select superior genotypes; iv) evaluate the correlations between quality and susceptibility to chilling injury of peaches and nectarines v) select parents with the combination of lower susceptibility to chilling injury and higher quality fruit. The study was carried out in EEAD-CSIC, Zaragoza - Spain, during the production cycle 2013/2014. A total of 40 peaches and nectarines genotypes from germplasm collection were evaluated. The quality characteristics as flesh firmness, total soluble solids, titratable acidity, pH, rippining index and flesh color parameters were evaluated. The fruits were submitted to cold storage at 0 °C and 5 °C, with 95% average relative humidity. The evaluations were after 14 and 28 days, it being observed the presence of symptoms, such as wooliness through mealiness, flesh grainy, leatheriness and flesh color changes, through browning, bleeding and off flavor. As a selection parameter was adopted 20% of genotypes that had a higher frequency of superiority for quality characteristics, susceptibility to chilling injury and the combining of both. For quality characteristic presented greater divergence the ‘Queen Giant’, ‘Sudanel Blanco’ and ‘Borracho de Jarque’. Based on the quality the eight genotypes were selected, ‘Andross’, ‘San Jaime’, ‘San Lorenzo’, ‘Borracho de Jarque’, ‘Sudanell 1’, ‘Carson’, ‘Baby Gold 6’ and ‘Stanford’. All genotypes studied exhibited susceptibility to one or more symptoms caused by cold storage during 28 days, independent of temperature. For 14 days, the ‘Baby Gold 6’, ‘Flavortop’ and ‘Queen Giant’ genotypes did not show any physiological disorder caused by cold. In general, the temperature of 0 °C favored fruit postharvest conservation, it have a lower incidence and severity of symptoms caused by cold storage. The storage for 14 days contributed for the lower incidence of damage in the genotypes fruits studied. For 14 days, with both temperatures, it was observed divergence for ‘Queen Giant’, ‘Sudanell Blanco’, ‘Baby Gold 6’ ‘GF3’, ‘Baby Gold 8’, ‘Campiel’ and ‘Campiel Rojo’ genotypes. For 28 days, in the 5 °C condition, ‘Queen Giant’, ‘Big Top’, ‘Flavortop’ and ‘Redhaven’ genotypes were divergents. Based on susceptibility to chilling injury at 0 °C, the eight genotypes were selected, it being these, ‘Queen Giant’, ‘Keimoes’, ‘Flavortop’, ‘Big Top’, 'Redhaven', 'Sudanell 3', 'Bonet I' and ‘Carson’. The quality parameters as rippining index, soluble solids, firmness and titratable acidity presented correlation among them. These, also it had correlation with woolines and bowning, what it indicate that fruits with more ripening can have this symptoms more easily. The browning, mealiness, flesh grainy and off flavor variables were correlationed with the time period and temperartures, what it confirm that these symptoms are the main disorders caused by cold storage. The quality characteristics together susceptibility to chilling injury allowed selected ‘Baby Gold 6’, ‘Sarell’, ‘Keimoes’, ‘GF3’ ‘San Jaime’, ‘Big Top’, ‘Sudanell 1’, ‘Carson’, ‘Baby Gold 8’, and ‘San Lorenzo’ genotypes.

Développer des plantes résistantes aux pathogènes nécessite d’identifier des sources de résistance, de les introduire dans le fonds génétique souhaité et de s’assurer de leurs performance et durabilité. Par l’étude de 2 pathosystèmes distincts, ces travaux de thèse m’ont permis d’appréhender ces différentes étapes.Le pathosystème tomate-bégomovirus présente un intérêt économique important. Une étude par transgénèse suggère que la surexpression de SlNAC1 (codant un facteur de transcription) est bien impliquée dans des mécanismes de résistance au bégomovirus, probablement en activant les mécanismes de résistance systémique acquise. Cette surexpression semble s’accompagner d’un retard de croissance, incompatible avec une utilisation en amélioration des plantes. Le pathosystème tomate-potyvirus permet de comparer au sein d’une même espèce des allèles de résistance naturels et induits, reposant sur des mutations affectant un facteur d’initiation de la traduction eIF4E1. Les travaux menés sur ces allèles ont permis de mettre en lumière une redondance au sein de la petite famille multigénique eIF4E vis-à-vis de la résistance, impliquant eIF4E2 dans les mécanismes de sensibilité et sont en faveur de l’utilisation d’allèles de résistance fonctionnels. De plus, l’introgression de l’allèle naturel dans une lignée de tomate possédant déjà d’autres gènes de résistance aux pathogènes s’accompagne d’une érosion du spectre de résistance dû à un contournement massif par la souche PVY-LYE84.Nos travaux soulignent ainsi les précautions à prendre lors de la mise en place de résistances génétiques ainsi que la nécessité de concilier ces approches avec l’étude du développement de la plante. / The development of plants resistant to need to identify sources of resistance, to introduce them into the desired genetic background and subsequently to monitor their performance and durability. By studying two distinct pathosystems, this thesis work has allowed me to address those three stages.The tomato-begomoviruses pathosystem presents a significant economic interest. Transgenic approach suggests that SlNAC1 (coding for a transcription factor) is involved in resistance mechanisms to begomoviruses, probably by triggering systemic acquired resistance. Besides, the SlNAC1 overexpression seems to be associated with growth retardation, a side-effect incompatible with its use in plant breeding.The tomato-potyviruses pathosystem allowed the comparison within the same species of both natural and induced resistance alleles, associated with mutations affecting the translation initiation factor eIF4E1. Work on those alleles highlighted a redundancy effect within the eIF4E small multigenic family toward resistance, affecting eIF4E2 and favouring the use of functional resistance alleles in resistance. In parallel, the introgression of thebroad-spectrum resistance natural allele was introgressed into a tomato line that already possess other pathogen resistance genes is accompanied by an erosion of the resistance spectrum, due to a massive resistance breakdown by the PVY-LYE84 strain.Altogether, this current work highlight cautions to be taken for introducing resistance genes as well as stressing the importance of combining those approaches with the study of plant development.

Despite of the documented impacts of the so-called green revolution, food security in the world faces new challenges in terms of population growth, increases in no-agricultural land use (urbanization), and climate change. Trends in food security show that the world community is operating within two limits of food system: (i) the quantity of food that can be produced under a given climate; and (ii) the quantity of food needed by a growing and changing population. Therefore, taking food security successfully into the future requires novel approaches to boost agricultural productivity in order to balance food supply and demand without expanding the agricultural land. To date, progress in wheat yield has been largely the result of the development of dwarf varieties through introgression of reduced height (Rht) genes. The height reductions arising from the presence of these genes increased yield by alteration of partitioning of dry matter and nitrogen in favour of the spike. However, increased partitioning through additional reductions in plant height is not likely; as comparative studies indicate that wheat yield is reduced when plants are shortened beyond a threshold, and most of the modern cultivars have reached the optimal height. Therefore, this dissertation aimed to identify the physiological attributes able to produce yield increases in the Rht genotypes with the optimal heights. Approaches based on physiological understanding of yield are necessary for developing genotypes combining high yielding potential and agronomic traits of superior adaptation, and for understanding yield limiting factors. Yet, direct measurement of physiological variables is often difficult or expensive; as an example, measuring plant water status in the field is problematic, with techniques such as psychrometry generally only being suitable for laboratory studies. Therefore, proxy such as tissue RWC may be a good alternative measure of plant water status. We aimed to address these questions with three components of experimental research :(i) proxy-based screening to increased photosynthetic rate and water use efficiency in wheat; (ii) determinants of increased HI in lines with different Rht genes (b, c) when incorporated into contrasting background wheat genomes (B, D), and the relative effect on N partitioning during grain filling; (iii) analyses of stable isotopes (δ²H, δ¹⁸O, δ¹⁵N and δ¹³C) in an agronomic perspective in alley cropping systems associated with adjacent N₂ fixing trees, in terms of hydraulic redistribution, N availability and crop yields. In this thesis, the proxy-based approach to crop selection was defined as a surrogate-based (proxy and surrogate used interchangeably) screening of cultivars for morphological, anatomical, and physiological traits of performance or crop environmental responses. The research proposed steps for conducting a proxy-based crop selection programme. A comparative screening of 23 Eps cultivars and ranking for traits of photosynthetic and water use efficiency showed the correlative relationships of SLA to An, WUEi, leaf N, Δ¹³C, Kh, leaf RWC, and IVD. Additionally, it was observed that IVD may influence WUE and Amax. It was suggested that these relationships of SLA to traits of photosynthesis possibly resulted from the association of SLA and the leaf biochemical characteristics. Attention was also given to examining the mechanistic foundations that determine the relationship between plant height and yield. The results showed the straw-shortening significantly correlated both with Amax and Kh; and SLA decreased with the level of dwarfing; and the Amax related both Kh and SLA. Therefore, it was proposed that the straw-shortening may affects Amax by exerting a controlling influence over Kh through SLA. Moreover, both the partitioning of N to spike and the flag leaf N were related to plant height and growth stage. Additionally, the increased post-anthesis partitioning of N to grain associated with high N uptake rate and high MRT of N were probably the traits behind increased NUE and NHI. The data also indicated that increased grain number per spike, kernel weight and reduced peduncle length might be the driver of the increased HI in this experiment. The test of the hypothesis that there might be practical application of the analyses of the natural abundance of stable isotopes (δ²H, δ¹⁸O, δ¹³C, and δ¹⁵N) and isotopic mixing model by IsoSource to understand plant interactions in terms of water redistribution and nitrogen transfer and uptake in agroforestry systems, indicated a consistent gradient in depletion of wheat xylem water δ²H, δ¹⁸O, and δ¹⁵N in leaf as moving further away from the tree line. The data also reflected a consistent pattern of isotopic values (δ²H, δ¹⁸O, and δ¹⁵N) in wheat in the proximity of the tree being similar to that of the tree, suggesting they were using the same source of water and N. Similarly, an isotopic mixing model data showed that the crops in the proximity of the trees accessed considerably amounts of the water and nitrogen redistributed by trees. The study also indicated the improvement in water use efficiency, chlorophyll content, grain number per spike, and grain yield for the crops nearest to the trees for a distance up to 5 m. In conclusion, selection for increased HI should shift focus from reduced plant height to include increased grain number and kernel weight, increased partitioning of N to spike, reduced peduncle length, and low SLA. Finally, the hypothesis that efflux of water and N in agroforestry system from tree roots in topsoil and influences a number of physiological functions of neighbouring crops was confirmed by isotopic and physiological data.

A series of studies are presented on the genetic evaluation of cultivated potato (Solanum tuberosum L.) to improve the accuracy and efficiency of selection at various stages of a breeding programme. The central theme was the use of correlated data, such as relationship information and spatial and across-trial correlations, within a linear mixed modelling framework to enhance the evaluation of candidate genotypes and to improve the genetic response to selection. Analyses focused on several social and economically-important traits for the enhancement of the nutritional value, disease resistance and yield of potato tubers. At the formative stages of a breeding scheme, devising a breeding strategy requires an improved understanding of the genetic control of target traits for selection. To guide a strategy that aims to enhance the micronutrient content of potato tubers (biofortification), univariate and multivariate Bayesian models were developed to estimate genetic parameters for micronutrient tuber content from a breeding population generated from crosses between Andean landrace cultivars. The importance of the additive genetic components and extent of the narrow-sense heritability estimates indicated that genotypic 'individual' recurrent selection based on empirical breeding values rather than family-based selection is likely to be the most effective strategy in this breeding population. The magnitude of genetic correlations also indicated that simultaneous increases in important tuber minerals, iron and zinc, could be achieved. Optimising selection efficiency is an important ambition of plant breeding programmes. Reducing the level of candidate replication in field trials may, under certain circumstances, contribute to this aim. Empirical field data and computer simulations inferred that improved rates of genetic gain with p-rep (partially replicated) testing could be obtained compared with testing in fully replicated trials at the early selection stages, particularly when testing over two locations. P-rep testing was able to increase the intensity of selection and the distribution of candidate entries across locations to account for G×E effects was possible at an earlier stage than is currently practised. On the basis of these results, it was recommended that the full replication of trials (at the first opportunity, when enough planting material is available) at a single location in the early stages of selection should be replaced with the partial replication of selection candidates that are distributed over two locations. Genetic evaluation aims to identify genotypes with high empirical breeding values (EBVs) for selection as parents. Using mixed models, spatial parameters to target greater control of localised field heterogeneity were estimated and variance models to account for across-trial genetic heterogeneity were tested for the evaluation of soil-borne powdery scab disease and tuber yield traits at the early stages of a selection programme. When spatial effects improved model fit, spatial correlations for rows and columns were mostly small for powdery scab, and often small and negative for marketable and total tuber yield suggesting the presence of interplot competition in some years for tuber yield traits. For the evaluation of powdery scab, genetic variance structures were tested using data from 12 years of long-term potato breeding METs (multi-environment trials). A simple homogeneous correlation model for the genetic effects was preferred over a more complex factor analytic (FA) model. Similarly, for the MET evaluation of tuber yield at the early stages, there was little benefit in using more complex FA models, with simple correlation structures generally the most favourable models fitted. The use of less complex models will be more straightforward for routine implementation of potato genetic evaluations in breeding programmes. Evaluations for (marketable) tuber yield were extended to multi-location MET data to characterise both genotypes and environments, allowing a re-evaluation of New Zealand MET selection strategies aimed at broad adaptation. Using a factor analytic mixed model, results indicated that the programme’s two main trial locations in the North and the South Islands optimised differentiation between genotypes in terms of G×E effects. There was reasonable performance stability of genotypes across test locations and evidence was presented for some, but limited, genetic progress of cultivars and advanced clonal selections for tuber marketable yield in New Zealand over recent years. The models and selection strategies investigated and developed in this thesis will allow an improved and more systematic application of genetic evaluations in potato selection schemes. This will provide the basis for well informed decisions to be made on selection candidates for the genetic improvement of potato in breeding programmes.

Maize (Zea mays L.) ranks third as a food crop after wheat and rice and is characterized not only as a cereal crop but also as a vegetable. Maize used as a vegetable is known as “baby corn”. Baby corn consists of unfertilized young ears harvested 2 or 3 days after silk emergence. The present study was implemented in 2009 at Western Kentucky University Agriculture Research and Education Center (36.93 N, 86.47 E) in Bowling Green, Kentucky. The purpose of the study was to compare the effect of different schemes of harvest on baby corn (BC) yield, grain maize (GM) yield, and estimated economic return. Experimental harvest treatments were 1) no BC harvest, only GM harvest, 2) first harvest as BC, final harvest as GM, 3) first and second harvests as BC, final harvest as GM, and 4) first, second, and third harvests as BC, final harvest as GM. Average estimated BC yields (Kg/ha) for Treatments 2, 3, and 4 were 1445.1, 2681.8, and 3437.5; GM yields (Kg/ha) for Treatments 1, 2, and 3 were 12522.2, 8226.5, and 1380.9; respectively. Since few grain kernels were found after three harvests for BC (Treatment 4), no usable GM yield was produced. BC and GM yields were used for evaluating the economic returns. Results indicated that the sequence of best economic returns would be obtained by harvesting BC three times (Treatment 4), first two harvests for BC and the final for GM (Treatment 3), first harvest for BC and subsequent for GM (Treatment 2), and only for GM harvest (Treatment 1). Although the pattern for only BC harvest was the most profitable system, the human labor requirement and critical timing of harvest limited its production. In states similar to Kentucky, BC could only be grown as an additional crop or to supplant a limited amount of traditional GM hectarage.