Mark C. Meacham

Biofortification of UK food crops with selenium

Se is an essential element for animals. In man low dietary Se intakes are associated with health disorders including oxidative stress-related conditions, reduced fertility and immune functions and an increased risk of cancers. Although the reference nutrient intakes for adult females and males in the UK are 60 and 75 microg Se/d respectively, dietary Se intakes in the UK have declined from >60 microg Se/d in the 1970s to 35 microg Se/d in the 1990s, with a concomitant decline in human Se status. This decline in Se intake and status has been attributed primarily to the replacement of milling wheat having high levels of grain Se and grown on high-Se soils in North America with UK-sourced wheat having low levels of grain Se and grown on low-Se soils. An immediate solution to low dietary Se intake and status is to enrich UK-grown food crops using Se fertilisers (agronomic biofortification). Such a strategy has been adopted with success in Finland. It may also be possible to enrich food crops in the longer term by selecting or breeding crop varieties with enhanced Se-accumulation characteristics (genetic biofortification). The present paper will review the potential for biofortification of UK food crops with Se.

Shoot yield drives phosphorus use efficiency in Brassica oleracea and correlates with root architecture traits

The environmental and financial costs of using inorganic phosphate fertilizers to maintain crop yield and quality are high. Breeding crops that acquire and use phosphorus (P) more efficiently could reduce these costs. The variation in shoot P concentration (shoot-P) and various measures of P use efficiency (PUE) were quantified among 355 Brassica oleracea L. accessions, 74 current commercial cultivars, and 90 doubled haploid (DH) mapping lines from a reference genetic mapping population. Accessions were grown at two or more external P concentrations in glasshouse experiments; commercial and DH accessions were also grown in replicated field experiments. Within the substantial species-wide diversity observed for shoot-P and various measures of PUE in B. oleracea, current commercial cultivars have greater PUE than would be expected by chance. This may be a consequence of breeding for increased yield, which is a significant component of most measures of PUE, or early establishment. Root development and architecture correlate with PUE; in particular, lateral root number, length, and growth rate. Significant quantitative trait loci associated with shoot-P and PUE occur on chromosomes C3 and C7. These data provide information to initiate breeding programmes to improve PUE in B. oleracea.

Shoot calcium and magnesium concentrations differ between subtaxa, are highly heritable, and associate with potentially pleiotropic loci in Brassica oleracea

Calcium (Ca) and magnesium (Mg) are the most abundant group II elements in both plants and animals. Genetic variation in shoot Ca and shoot Mg concentration (shoot Ca and Mg) in plants can be exploited to biofortify food crops and thereby increase dietary Ca and Mg intake for humans and livestock. We present a comprehensive analysis of within-species genetic variation for shoot Ca and Mg, demonstrating that shoot mineral concentration differs significantly between subtaxa (varietas). We established a structured diversity foundation set of 376 accessions to capture a high proportion of species-wide allelic diversity within domesticated Brassica oleracea, including representation of wild relatives (C genome, 1n = 9) from natural populations. These accessions and 74 modern F1 hybrid cultivars were grown in glasshouse and field environments. Shoot Ca and Mg varied 2- and 2.3-fold, respectively, and was typically not inversely correlated with shoot biomass, within most subtaxa. The closely related capitata (cabbage) and sabauda (Savoy cabbage) subtaxa consistently had the highest mean shoot Ca and Mg. Shoot Ca and Mg in glasshouse-grown plants was highly correlated with data from the field. To understand and dissect the genetic basis of variation in shoot Ca and Mg, we studied homozygous lines from a segregating B. oleracea mapping population. Shoot Ca and Mg was highly heritable (up to 40%). Quantitative trait loci (QTL) for shoot Ca and Mg were detected on chromosomes C2, C6, C7, C8, and, in particular, C9, where QTL accounted for 14% to 55% of the total genetic variance. The presence of QTL on C9 was substantiated by scoring recurrent backcross substitution lines, derived from the same parents. This also greatly increased the map resolution, with strong evidence that a 4-cM region on C9 influences shoot Ca. This region corresponds to a 0.41-Mb region on Arabidopsis (Arabidopsis thaliana) chromosome 5 that includes 106 genes. There is also evidence that pleiotropic loci on C8 and C9 affect shoot Ca and Mg. Map-based cloning of these loci will reveal how shoot-level phenotypes relate to Ca2+ and Mg2+ uptake and homeostasis at the molecular level.

Genetic analysis of potassium use efficiency in Brassica oleracea.

BACKGROUND AND AIMS Potassium (K) fertilizers are used in intensive and extensive agricultural systems to maximize production. However, there are both financial and environmental costs to K-fertilization. It is therefore important to optimize the efficiency with which K-fertilizers are used. Cultivating crops that acquire and/or utilize K more effectively can reduce the use of K-fertilizers. The aim of the present study was to determine the genetic factors affecting K utilization efficiency (KUtE), defined as the reciprocal of shoot K concentration (1/[K](shoot)), and K acquisition efficiency (KUpE), defined as shoot K content, in Brassica oleracea. METHODS Genetic variation in [K](shoot) was estimated using a structured diversity foundation set (DFS) of 376 accessions and in 74 commercial genotypes grown in glasshouse and field experiments that included phosphorus (P) supply as a treatment factor. Chromosomal quantitative trait loci (QTL) associated with [K](shoot) and KUpE were identified using a genetic mapping population grown in the glasshouse and field. Putative QTL were tested using recurrent backcross substitution lines in the glasshouse. KEY RESULTS More than two-fold variation in [K](shoot) was observed among DFS accessions grown in the glasshouse, a significant proportion of which could be attributed to genetic factors. Several QTL associated with [K](shoot) were identified, which, despite a significant correlation in [K](shoot) among genotypes grown in the glasshouse and field, differed between these two environments. A QTL associated with [K](shoot) in glasshouse-grown plants (chromosome C7 at 62.2 cM) was confirmed using substitution lines. This QTL corresponds to a segment of arabidopsis chromosome 4 containing genes encoding the K+ transporters AtKUP9, AtAKT2, AtKAT2 and AtTPK3. CONCLUSIONS There is sufficient genetic variation in B. oleracea to breed for both KUtE and KUpE. However, as QTL associated with these traits differ between glasshouse and field environments, marker-assisted breeding programmes must consider carefully the conditions under which the crop will be grown.

Screening for genotype and environment effects on nitrate accumulation in 24 species of young lettuce

BACKGROUND Nitrate accumulates in plants in response to N supply, aerial environment (predominantly light), and genotype. This paper characterises the effects of genotype, environment, and their interactions on nitrate accumulation by 24 cultivated and wild lettuce accessions grown hydroponically in winter and summer. The results will inform future strategies for selecting for low-nitrate varieties. RESULTS A preliminary study in which two accessions were sampled for nitrate over time showed largest differences between cultivars in the early-middle period of growth. Sampling the whole population of lettuce at this stage revealed significant effects of genotype, environment (with nitrate concentrations generally higher in winter), and genotype × environment interactions (largely due to a wider range of concentrations in summer). Changes in the ranking of accessions for nitrate accumulation between the two growing seasons were generally small for cultivated morphotypes. Shoot nitrate concentrations and water contents were positively associated, particularly in summer when separate relationships for different cultivated morphotypes (butterhead, cos/Romaine, crisp, leaf, and stem lettuce) were detected. Expressing nitrate concentration on either a shoot fresh or dry matter basis had relatively little effect on the ranking of most cultivated accessions, but not for the wild types. CONCLUSION There is a well-defined sampling window when differences in nitrate accumulation between lettuce genotypes are at a maximum. Delaying sampling may allow morphological changes in head form to mask earlier genotypic differences. Genotype × environment interactions are predominantly of the non-crossover type and have only a small effect on changes in the ranking of accessions between seasons, allowing selections to be made at any time of year. At least part of the genotypic variation in nitrate accumulation is associated with differences in shoot water content.

High-throughput phenotyping (HTP) identifies seedling root traits linked to variation in seed yield and nutrient capture in field-grown oilseed rape (Brassica napus L.)

Background and Aims Root traits can be selected for crop improvement. Techniques such as soil excavations can be used to screen root traits in the field, but are limited to genotypes that are well-adapted to field conditions. The aim of this study was to compare a low-cost, high-throughput root phenotyping (HTP) technique in a controlled environment with field performance, using oilseed rape (OSR; Brassica napus) varieties. Methods Primary root length (PRL), lateral root length and lateral root density (LRD) were measured on 14-d-old seedlings of elite OSR varieties (n = 32) using a ‘pouch and wick’ HTP system (∼40 replicates). Six field experiments were conducted using the same varieties at two UK sites each year for 3 years. Plants were excavated at the 6- to 8-leaf stage for general vigour assessments of roots and shoots in all six experiments, and final seed yield was determined. Leaves were sampled for mineral composition from one of the field experiments. Key Results Seedling PRL in the HTP system correlated with seed yield in four out of six (r = 0·50, 0·50, 0·33, 0·49; P < 0·05) and with emergence in three out of five (r = 0·59, 0·22, 0·49; P < 0·05) field experiments. Seedling LRD correlated positively with leaf concentrations of some minerals, e.g. calcium (r = 0·46; P < 0·01) and zinc (r = 0·58; P < 0·001), but did not correlate with emergence, general early vigour or yield in the field. Conclusions Associations between PRL and field performance are generally related to early vigour. These root traits might therefore be of limited additional selection value, given that vigour can be measured easily on shoots/canopies. In contrast, LRD cannot be assessed easily in the field and, if LRD can improve nutrient uptake, then it may be possible to use HTP systems to screen this trait in both elite and more genetically diverse, non-field-adapted OSR.

Inter-cultivar variation in soil-to-plant transfer of radiocaesium and radiostrontium in Brassica oleracea

Radiocaesium and radiostrontium enter the human food chain primarily via soil-plant transfer. However, uptake of these radionuclides can differ significantly within species (between cultivars). The aim of this study was to assess inter-cultivar variation in soil-to-plant transfer of radiocaesium and radiostrontium in a leafy crop species, Brassica oleracea. This study comprised four independent experiments: two pot experiments in a controlled environment artificially contaminated with radiocaesium, and two field experiments in an area contaminated with radiocaesium and radiostrontium in the Chernobyl Exclusion Zone. Radiocaesium concentration ratios varied 35-fold among 27 cultivars grown in pots in a controlled environment. These 27 cultivars were then grown with a further 44 and 43 other cultivars in the Chernobyl Exclusion Zone in 2003 and 2004, respectively. In the field-grown cultivars radiocaesium concentration ratios varied by up to 35-fold and radiostrontium concentration ratios varied by up to 23-fold. In three of these experiments (one pot experiment, two field experiments) one out of the 27 cultivars was found to have a consistently lower radiocaesium concentration ratio than the other cultivars. The two field experiments showed that, five out of the 66 cultivars common to both experiments had consistently lower radiocaesium concentration ratios, and two cultivars had consistently lower radiostrontium concentration ratios. One cultivar had consistently lower radiocaesium and radiostrontium concentration ratios. The identification of cultivars that have consistently lower radiocaesium and/or radiostrontium concentration ratios suggests that cultivar selection or substitution may be an effective remediation strategy in radiologically contaminated areas. Future research should focus on plant species that are known to be the largest contributors to human dose.