Saponin seed priming improves salt tolerance in quinoa

Quinoa (Chenopodium quinoa Willd.) is a facultative halophyte of great value, and World Health Organization has selected this crop, which may assure future food and nutritional security under changing climate scenarios. However, germination is the main critical stage of quinoa plant phenology affected by salinity. Therefore, two experiments were conducted to improve its performance under salinity by use of saponin seed priming. Seeds of cv. Titicaca were primed in seven different solutions with varying saponin concentrations (i.e. 0%, 0.5%, 2%, 5%, 10%, 15%, 25% and 35%), and then, performances of primed seeds were evaluated based on mean germination time and final germination percentage in germination assays (0 and 400 mM NaCl stress). Saponin solutions of 10%, 15% and 25% concentration were found most effective priming tools for alleviating adverse effects of salt stress during seed germination. Performances of these primed seeds were further evaluated in pot study. At six‐leaf stage, plants were irrigated with saline water having either 0 or 400 mM NaCl. The results indicated that saline irrigation significantly decreased the growth, physiology and yield of quinoa, whereas saponin priming found operative in mitigating the negative effects of salt stress. Improved growth, physiology and yield performance were linked with low ABA concentration, better plant water (osmotic and water potential) and gas relations (leaf photosynthetic rate, stomatal conductance), low Na⁺ and high K⁺ contents in leaves. Our results suggest that saponin priming could be used as an easy‐operated and cost‐effective technology for sustaining quinoa crop growth on salt‐affected soils.     

Effect of high temperature on pollen morphology,plant growth and seed yield in quinoa (Chenopodium quinoa Willd.)

Quinoa (Chenopodium quinoa Willd.) has gained considerable attention worldwide during the past decade due to its nutritional and health benefits. However, its susceptibility to high temperatures has been reported as a serious obstacle to its global production. The objective of this study was to evaluate quinoa growth and pollen morphology in response to high temperatures. Pollen morphology and viability, plant growth and seed set, and several physiological parameters were measured at anthesis in two genotypes of quinoa subjected to day/night temperatures of 22/16°C as a control treatment and 40/24°C as the heat stress treatment. Our results showed that heat stress reduced the pollen viability between 30% and 70%. Although no visible morphological differences were observed on the surface of the pollen between the heat‐stressed and non‐heat‐stressed treatments, the pollen wall (intine and extine) thickness increased due to heat stress. High temperature did not affect seed yield, seed size and leaf greenness. On the other hand, high temperature improved the rate of photosynthesis. We found that quinoa has a high plasticity in response to high temperature, though pollen viability and pollen wall structure were affected by high temperatures in anthesis stage. This study is also the first report of quinoa pollen being trinucleate.     

Influence of the root endophyte Piriformospora indica on the plant water relations,gas exchange and growth of Chenopodium quinoa at limited water availability

This study aimed to evaluate the ability of Piriformospora indica to colonize the root of Chenopodium quinoa and to verify whether this endosymbiont can improve the growth, performance and drought resistance of this species. The study delivered, for the first time, evidence for successful colonization of P. indica in quinoa. Hence, pot experiment was conducted in the greenhouse, where inoculated and non‐inoculated plants were subjected to ample (40%–50% WHC) and deficit (15%–20%WHC) irrigation treatments. Drought adversely influenced the plant growth, leading to decline the total plant biomass by 74%. This was linked to an impaired photosynthetic activity (caused by lower g_s and C_i/C_a ratio; stomatal limitation of photosynthesis) and a higher risk of ROS production (enhanced ETR/A_gross ratio). P. indica colonization improved quinoa plant growth, with total biomass increased by 8% (controls) and 76% (drought‐stressed plants), confirming the growth‐promoting activity of P. indica. Fungal colonization seems to diminish drought‐induced growth hindrance, likely, through an improved water balance, reflected by the higher leaf ψ_w and g_s. Additionally, stomatal limitation of photosynthesis was alleviated (indicated by enhanced C_i/C_a ratio and A_net), so that the threat of oxidative stress was minimized (decreased ETR/A_gross). These results infer that symbiosis with P. indica could negate some of the detrimental effects of drought on quinoa growth, a highly desired feature, in particular at low water availability.     

Effects of Increasing Salinity Stress and Decreasing Water Availability on Ecophysiological Traits of Quinoa (Chenopodium quinoa Willd.) Grown in a Mediterranean‐Type Agroecosystem

Quinoa is a native Andean crop for domestic consumption and market sale, widely investigated due to its nutritional composition and gluten‐free seeds. Leaf water potential (Ψ_leaf) and its components and stomatal conductance (g_s) of quinoa, cultivar Titicaca, were investigated in Southern Italy, in field trials (2009 and 2010). This alternative crop was subjected to irrigation treatments, with the restitution of 100 %, 50 % and 25 % of the water necessary to replenish field capacity, with well water (100 W, 50 W, 25 W) and saline water (100 WS, 50 WS, 25 WS) with an electrical conductivity (EC_w) of 22 dS m^?1. As water and salt stress developed and Ψ_leaf decreased, the leaf osmotic potential (Ψ_π) declined (below ?2.05 MPa) to maintain turgor. Stomatal conductance decreased with the reduction in Ψ_leaf (with a steep drop at Ψ_leaf between ?0.8 and 1.2 MPa) and Ψ_π (with a steep drop at Ψ_π between ?1.2 and ?1.4 MPa). Salt and drought stress, in both years, did not affect markedly the relationship between water potential components, RWC and g_s. Leaf water potentials and g_s were inversely related to water limitation and soil salinity experimentally imposed, showing exponential (Ψ_leaf and turgor pressure, Ψ_p, vs. g_s) or linear (Ψ_leaf and Ψ_p vs. SWC) functions. At the end of the experiment, salt‐irrigated plants showed a severe drop in Ψ_leaf (below ?2 MPa), resulting in stomatal closure through interactive effects of soil water availability and salt excess to control the loss of turgor in leaves. The effects of salinity and drought resulted in strict dependencies between RWC and water potential components, showing that regulating cellular water deficit and volume is a powerful mechanism for conserving cellular hydration under stress, resulting in osmotic adjustment at turgor loss. The extent of osmotic adjustment associated with drought was not reflected in Ψ_π at full turgor. As soil was drying, the association between Ψ_leaf and SWC reflected the ability of quinoa to explore soil volume to continue extracting available water from the soil. However, leaf ABA content did not vary under concomitant salinity and drought stress conditions in 2009, while differing between 100 W and 100 WS in 2010. Quinoa showed good resistance to water and salt stress through stomatal responses and osmotic adjustments that played a role in the maintenance of a leaf turgor favourable to plant growth and preserved crop yield in cropping systems similar to those of Southern Italy.     

Antioxidative Response of Quinoa Exposed to Iso‐Osmotic,Ionic and Non‐Ionic Salt Stress

Antioxidants play an important role in adapting plants to abiotic stress by detoxifying reactive oxygen species (ROS). Involvement of antioxidant enzymes in abiotic stress tolerance of highly stress‐tolerant quinoa was studied in a climatic chamber at 6 mOsm (milliosmolar) ionic (300 mm NaCl) and non‐ionic (600 mm mannitol) salts combined with increasing levels of potassium K1 and K2 (6, 12 mm ), respectively. Fifteen days of salt treatment (both ionic and non‐ionic) decreased plant growth (shoot and root fresh weight), stomatal conductance and chlorophyll content index. Furthermore, both forms of salt stress increased the activities of superoxide dismutase, catalase, ascorbate peroxidase and peroxidase up to 2.33‐, 3.98‐, 4.78‐ and 5.55‐folds, respectively, compared to no salt treatment, whereas membrane stability index decreased corresponding to increase in lipid peroxidation (malondialdehyde), with salt treatments compared to non‐stressed plants. However, no significant effect of potassium and salt treatments has been noticed on the maximal photochemical efficiency of PSII. The results suggested that enhanced antioxidant enzymes activity under salt stress could be one of the factors responsible for abiotic stress tolerance in quinoa.     

Growth and Physiological Responses of Quinoa to Drought and Temperature Stress

Quinoa (Chenopodium quinoa Willd.), traditionally called the mother of grains, has the potential to grow under high temperatures and drought, tolerating levels regarded as stresses in other crop species. A pot experiment was conducted in a climate chamber to investigate the potential of quinoa tolerance to increasing drought and temperature. Quinoa plants were subjected to three irrigation and two temperature regimes. At low temperature, the day/night climate chamber temperature was maintained at 18/8 °C and 25/20 °C for high temperature throughout the treatment period. The irrigation treatments were full irrigation (FI), deficit irrigation (DI) and alternate root‐zone drying (ARD). FI plants were irrigated daily to the level of the pot's water‐holding capacity. In DI and ARD, 70 % water of FI was applied either to the whole pot or to one side of the pot alternating, respectively. The results indicated that plant height and shoot dry weight significantly decreased by ARD and DI compared to FI treatment both at low and at high temperatures. However, plants in ARD treatment showed significantly higher plant height and shoot dry weight compared to DI especially at higher temperature, which is linked to increased xylem ion content. Higher quinoa plant growth in ARD was associated with increase in water‐use efficiency (WUE_i) due to higher abscisic acid concentration and higher nutrient contents compared to DI. From results, it can be concluded that quinoa plant growth is favoured by high temperature (25/20 °C) and ARD is an effective irrigation strategy to increase WUE in drought prone areas.     

The Situation for Quinoa and Its Production in Southern Bolivia: From Economic Success to Environmental Disaster

In Bolivia, one of the world’s most important centres of plant domestication, there is growing awareness of the value of native Andean crops, both for domestic consumption and for market sale – notably the virtually boom‐like consumer demand for quinoa around the world. The southern altiplano of Bolivia, south of Oruro, relies almost purely on the production of quinoa and breeding of llamas, which have also been selected as the two commodities of priority to the government to increase the income of the country. Presently, however, quinoa is facing increasing problems in production, owing to its increasing export market and price. The flat areas around the salt desert of the southern altiplano, previously characterized by natural vegetation fed by the llamas, are being increasingly sown with quinoa, hence transformed into deserts, because intensive cultivation methods make the soil loose its fertility. Possible solutions to these problems will require extensive efforts in the south, in addition to various strategies, which also include other parts of the Bolivian altiplano and a strengthened focus on other Andean crops.     

The locus for resistance to Asian soybean rust in PI 587855

Asian soybean rust (ASR) caused by Phakopsora pachyrhizi is one of the most serious soybean (Glycine max) diseases in tropical and subtropical areas. A soybean line, PI 587855, showed a resistance phenotype against ASR pathogens in Japan and South America at high frequency; however, little is known of the genetic control of this resistance and chromosomal location of the corresponding locus. Therefore, the aim of this study was to study the inheritance of PI 587855 resistance and map the corresponding locus with SSR markers aiming to use the linked markers in marker‐assisted selection. In the segregating population, resistance to ASR appeared to be controlled by a single dominant gene. The ASR resistance locus was mapped near to the chromosomal region where the resistant loci, Rpp1 and Rpp1‐b, were previously mapped. Comparative genetic mapping and disease reaction profiles of other seven lines carrying Rpp1 or Rpp1‐b to four Brazilian ASR isolates revealed that the resistance reaction exhibited by PI 587855 was similar to that of Rpp1‐b‐carrying varieties which have useful resistance to South American ASR strains.     

Development of gene‐based markers for the Turnip mosaic virus resistance gene retr02 in Brassica rapa

Turnip mosaic virus (TuMV) is responsible for a serious disease that affects the production of Chinese cabbage. Previous studies have cloned a series of TuMV resistance genes and developed molecular markers. In this study, a derived cleaved amplified polymorphism sequence (dCAPS) marker and a Kompetitive Allele Specific PCR (KASP) marker were developed based on a single recessive gene, retr02, which confers broad‐spectrum TuMV resistance in Chinese cabbage by means of an additional G at the junction of exon 1 and intron 1. The two markers were able to detect the retr02 allele in Chinese cabbage accessions used in breeding programmes. Compared with the dCAPS marker, the KASP marker was flexible, cost‐effective and quick to process, which is likely to be beneficial in establishing high‐throughput assays for marker‐assisted selection.     

S‐locus diversity and cross‐compatibility of wild Prunus avium for timber breeding

Prunus avium is primarily cultivated for its fruit, sweet cherries. However, it is also used to produce high‐quality timber. In a P. avium seed orchard, gametophytic self‐incompatibility is a restriction for free pollen flow and should be considered when establishing basic forest materials. In this study, S‐locus diversity and cross‐incompatibility of wild cherry individuals in clonal banks established for breeding for timber production were investigated. Wild cherry trees (140) with outstanding forest growth habit, collected in northern Spain, grafted and planted in two clonal banks, were genotyped at the S‐locus. The self‐incompatibility S‐locus genes, S‐RNase and SFB, were analysed by PCR. Twenty‐two S‐haplotypes, resulting in 72 different S‐genotypes, were identified. The genotypes were grouped into 33 incompatibility groups and 39 unique genotypes. This initial S‐locus analysis revealed large genetic diversity of wild cherry trees from the Spanish northern deciduous forest, and provides useful information for seed orchard design. Wild P. avium displays significantly more genetic diversity than what is detected in local cultivars, revealing a narrowing of genetic diversity during local domestication.     

Dissection of a major QTL for seed colour and fibre content in Brassica napus reveals colocalization with candidate genes for phenylpropanoid biosynthesis and flavonoid deposition

A major quantitative trait locus (QTL) influencing seed fibre and colour in Brassica napus was dissected by marker saturation in a doubled haploid (DH) population from the black‐seeded oilseed rape line ‘Express 617’ crossed with a yellow‐seeded B. napus line, ‘1012–98’. The marker at the peak of a sub‐QTL with a strong effect on both seed colour and acid detergent lignin content lay only 4 kb away from a Brassica (H+)‐ATPase gene orthologous to the transparent testa gene AHA10. Near the peak of a second sub‐QTL, we mapped a copy of the key phenylpropanoid biosynthesis gene cinnamyl alcohol dehydrogenase, while another key phenylpropanoid biosynthesis gene, cinnamoyl co‐a reductase 1, was found nearby. In a cross between ‘Express 617’ and another dark‐seeded parent, ‘V8’, Bna.CCR1 was localized in silico near the peak of a corresponding seed fibre QTL, whereas in this case Bna.CAD2/CAD3 lay nearby. Re‐sequencing of the two phenylpropanoid genes via next‐generation amplicon sequencing revealed intragenic rearrangements and functionally relevant allelic variation in the three parents.     

The generation of novel species hybrids between garden dahlias and Dahlia macdougallii to increase the gene pool for variety breeding

To transfer new traits into the gene pool of garden dahlias (Dahlia variabilis), crosses between garden dahlias (2n = 64) and the epiphytic species Dahlia macdougallii (2n = 32) from the section Epiphytum were conducted. Six hybrid plants were obtained. The hybrid status was verified using three SSR markers. In addition, flow cytometry was performed to determine the genome size of the hybrids and to show that the hybrids were hexaploid as expected from crosses between tetraploids and octoploids. The open pollinated progeny of the hybrids produced four progeny with octoploid genomes. The hybrids exhibited indeterminate vegetative growth and the formation of flowers from axillary buds, similar to the father D. macdougallii. This result is of interest for breeding new varieties of dahlia with traits that are not present in the current gene pool.     

Development of microsatellite markers in sweet passion fruit,and identification of length and conformation polymorphisms within repeat sequences

The aim of this study was to construct and characterize an SSR‐enriched genomic library for Passiflora alata, a fruit species native to the Brazilian plateau and the eastern Amazon region. There is potential to improve this crop, as the fruit is attractive because of its pleasant aroma and flavour characteristics. Of 862 sequences, 391 (45%) were found to have SSRs. We identified 412 microsatellites: 69% were classified as perfect, 23% imperfect, 5% interrupted and 3% were compound microsatellites. The main types of repeat sequences were dinucleotide (62.5%) and trinucleotide (23%) repeats. It was possible to design 312 primer pairs, and 229 of them were synthesized and tested. The amplicons were electrophoresed using denaturing and non‐denaturing gels to screen for divergence between two phenotypically distinct parents of a mapping population of P. alata. Length and conformation polymorphisms within repeat sequences amounted to 35% and 28%, respectively. The importance of the development of SSR markers for sweet passion fruit and the effectiveness of the SSCP approach to increase the available number of polymorphic SSRs are discussed.     

The first genetic linkage map of crape myrtle (Lagerstroemia) based on amplification fragment length polymorphisms and simple sequence repeats markers

Lagerstroemia (crape myrtle) are famous ornamental plants with large pyramidal racemes, long flower duration and diverse colours. Genetic maps provide an important genomic resource of basic and applied significance. A genetic linkage map was developed by genotyping 192 F₁ progeny from a cross between L. caudata (female) and L. indica (‘Xiang Xue Yun’) (male) with a combination of amplification fragment length polymorphisms (AFLP) and simple sequence repeats (SSR) markers in a double pseudo‐testcross mapping strategy. A total of 330 polymorphic loci consisting of 284 AFLPs and 46 SSRs showing Mendelian segregation were generated from 383 AFLP primer combinations and 150 SSR primers. The data were analysed using JoinMap 4.0 (evaluation version) to construct the linkage map. The map consisted of 20 linkage groups of 173 loci (160 AFLPs and 13 SSRs) covering 1162.1 cM with a mean distance of 10.69 cM between adjacent markers. The 20 linkage groups contained 2–49 loci and ranged in length from 7.38 to 163.57 cM. This map will serve as a framework for mapping QTLs and provide reference information for future molecular breeding work.     

Maternal inheritance of male sterility in the progeny of a natural hybrid between Cajanus lineatus and C. cajan

Adoption of pigeonpea hybrids in central and southern India is showing high impact with on‐farm yield advantages of >30%. The hybrid pigeonpea technology, the first in any legume crop, is based on a cytoplasmic‐nuclear male‐sterility (CMS) system. For a long‐term sustainability of hybrid programme, it is imperative that both nuclear diversity and cytoplasmic diversity are maintained among hybrid parents. In this context, a continuous search for new CMS‐inducing cytoplasms is necessary. This paper reports detection of maternal inheritance of male sterility in the progeny derived from a natural hybrid between a wild relative [Cajanus lineatus (W. & A.) Maesen comb. nov.] of pigeonpea and an unknown pigeonpea [Cajanus cajan (L.) Millsp.] genotype. In the present study, the male sterility was maintained up to BC₇F₁ generation by an advanced breeding pigeonpea line ICPL 99044. This male sterility inducing cytoplasm of C. lineatus was tagged as A₆. In future, this CMS genetic stock can be used to develop a range of new pigeonpea hybrids with high yield and adaptation.     

Origin of early maturing pigeonpea germplasm and its impact on adaptation and cropping systems

Pigeonpea breeding activities started about a century ago and for decades only late maturing cultivars dominated the global cultivation. Historically, no early maturing cultivar was available for a very long time and breeding of such varieties started in the third quarter of 20th century but at a low key. From these efforts, some pigeonpea varieties maturing in 90–150 days were bred. Information gathered from various sources revealed that the first few early maturing genotypes originated through spontaneous mutations in the late maturing field‐grown landraces. In other cases, transgressive segregation and induced mutations also produced early maturing varieties. At present, the high yielding early maturing cultivars are contributing significantly towards widening the adaption barriers and in the diversification of some age‐old cropping systems. In this paper, the authors, besides discussing the importance of early maturing cultivars in present agricultural systems, also summarize information related to the origin of primary sources of earliness.     

Introgression of useful genes from Thinopyrum intermedium to wheat for improvement of bread‐making quality

To study the influence of genes from Thinopyrum intermedium on traits affecting the bread‐making quality of wheat, two derivatives from a putative disomic addition line in cultivar ‘Vilmorin 27’ were used in cytological, biochemical and molecular characterization. Cytological analysis suggested that one of the derivatives (Line‐1) had a terminal deletion involving the long arm of chromosome 1D (2n = 42, Del‐1DL”), and the other (Line‐2) was a conventional addition line, but also carried the same deletion on chromosome 1D (2n = 44, Thi”+Del‐1DL”). Amplification and sequencing of high‐molecular‐weight glutenin subunit (HMW‐GS) genes coded by the Th. intermedium chromosome in Line‐2 indicated the presence of one x‐type with an extra cysteine and four (rather than one) unique y‐type genes. Rheological studies of Line‐1 showed significantly lower dough strength compared to ‘Vilmorin 27’, confirming the recognized role of Glu‐1D coded HMW‐GSs. Line‐2 showed significantly higher dough strength compared to the background cultivar, indicating a significant potential of Th. intermedium for improvement of bread‐making quality in wheat.     

A genetic linkage map in southern‐by‐spring oat identifies multiple quantitative trait loci for adaptation and rust resistance

We developed 178 recombinant inbred lines from a southern‐by‐spring oat population designated as “TxH.” These lines were genotyped to generate a high‐quality linkage map that resolved 6,902 markers into 21 linkage groups that matched closely with the latest hexaploid oat consensus map. Three major quantitative trait loci (QTLs) affecting heading date were found in locations that are consistent with known QTLs and candidate genes, and two other QTLs affecting heading date were found in novel locations. Five QTLs affecting plant height were found. Both sets of QTLs are responsible for transgressive segregation observed for these two traits. Four QTLs affecting resistance to crown rust, caused by the pathogen Puccinia coronata f. sp. avenae, were identified. Two of these QTLs are consistent with known clusters of rust resistance genes, while two may represent new locations of novel rust resistance genes. A complete set of SNP sequences suitable for generating markers for molecular selection is provided.     

The pattern of genetic diversity in the Andean grain crop quinoa (Chenopodium quinoa Willd). I. Associations between characteristics

Summary A germplasm collection comprising 294 accessions of quinoa was evaluated at Cambridge, England, to assess its potential as a break crop for arable agriculture at temperate latitudes. The origins of the accessions ranged from 40°S to 2°N and from sea level to an altitude of 3800 m, and 19 discrete and continuous characteristics describing the plants' pigmentation and morphology and the duration of developmental phases were noted or measured. There was significant variation among accessions for all the continuous characteristics, and the plant and inflorescence dimensions were all fairly strongly correlated, but the associations between the durations of developmental phases were surprisingly weak, suggesting that there is great scope for manipulation of the pattern of development through breeding. Contingency tables revealed associations between pairs of discrete characteristics. The means and variances of many continuous characteristics differed according to the discrete characteristics: for example accessions with branched plants took longer to germinate and were more variable in this respect than those with unbranched plants. Sometimes one form of a discrete characteristic was associated with a homogeneous group of accessions but the alternative form was not: such a relationship would be easily lost in a multivariate analysis. On the basis of there relationships, and using knowledge of the accessions' origins, they were classified into seven groups. It is concluded that the plant characteristics required for temperate agriculture are present to a large extent in the accessions from near sea level in southern-central Chile, but that the seed characteristics are scattered throughout the germplasm.     

Strategic use of Iranian bread wheat landrace accessions for genetic improvement: Core set formulation and validation

Iranian wheat landrace accessions (IWAs) were collected from country‐wide farm fields and market places in 1935 by a professor at the University of Tehran and shared with University of California at Davis, California. IWAs were further submitted to the genebank of International Maize and Wheat Improvement Center (CIMMYT), Mexico. 2,403 IWAs from CIMMYT’s genebank were assayed by DArT‐seq technology to assess genetic diversity. No apparent ecogeographic patterns related to genetic diversity were detected, probably due to long‐term transport and frequent interchange of landraces among farmers. A multivariate clustering procedure combining genotypic and phenotypic information was used in selecting a core‐set, which represented 15% of the hexaploid wheat accessions included in this study. This subset captured an estimated 93% of rare (frequency <0.05) alleles. Multisite phenotypic data (India, Mexico) validated the ability of the core‐set in detecting useful variants. Potential donor accessions for multiple traits (disease resistance, zinc concentration) were identified from the core‐set for wheat‐breeding. This report illustrates a breeder friendly robust core‐set formulation strategy for harnessing the useful genetic variation stored in the genebanks.