Review of wheat improvement for waterlogging tolerance in Australia and India: the importance of anaerobiosis and element toxicities associated with different soils

Background and Aims

The lack of knowledge about key traits in field environments is a major constraint to germplasm improvement and crop management because waterlogging-prone environments are highly diverse and complex, and the mechanisms of tolerance to waterlogging include a large range of traits. A model is proposed that waterlogging tolerance is a product of tolerance to anaerobiosis and high microelement concentrations. This is further evaluated with the aim of prioritizing traits required for waterlogging tolerance of wheat in the field.

Methods

Waterlogging tolerance mechanisms of wheat are evaluated in a range of diverse environments through a review of past research in Australia and India; this includes selected soils and plant data, including plant growth under waterlogged and drained conditions in different environments. Measurements focus on changes in redox potential and concentrations of diverse elements in soils and plants during waterlogging.

Key Results

(a) Waterlogging tolerance of wheat in one location often does not relate to another, and (b) element toxicities are often a major constraint in waterlogged environments. Important element toxicities in different soils during waterlogging include Mn, Fe, Na, Al and B. This is the first time that Al and B toxicities have been indicated for wheat in waterlogged soils in India. These results support and extend the well-known interactions of salinity/Na and waterlogging/hypoxia tolerance.

Conclusions

Diverse element toxicities (or deficiencies) that are exacerbated during waterlogging are proposed as a major reason why waterlogging tolerance at one site is often not replicated at another. Recommendations for germplasm improvement for waterlogging tolerance include use of inductively coupled plasma analyses of soils and plants.Key words: Waterlogging, microelements, toxicity, redox potential, wheat, anaerobiosis

‘No grain is ever produced without water, but too much water tends to spoil the grain and inundation is as injurious to growth as dearth of water.’ Narada Smriti XI, 19; circa 3000 bc.‘Waterlogging’ is defined as a condition of the soil where excess water limits gas diffusion; while ‘waterlogging tolerance’ is defined as survival or the maintenance of high growth rates, biomass accumulation or grain yield under waterlogging relative to non waterlogged (usually drained soil) conditions (Setter and Waters, 2003).

     

Mechanisms of waterlogging tolerance in wheat – a review of root and shoot physiology

We review the detrimental effects of waterlogging on physiology, growth and yield of wheat. We highlight traits contributing to waterlogging tolerance and genetic diversity in wheat. Death of seminal roots and restriction of adventitious root length due to O₂ deficiency result in low root:shoot ratio. Genotypes differ in seminal root anoxia tolerance, but mechanisms remain to be established; ethanol production rates do not explain anoxia tolerance. Root tip survival is short‐term, and thereafter, seminal root re‐growth upon re‐aeration is limited. Genotypes differ in adventitious root numbers and in aerenchyma formation within these roots, resulting in varying waterlogging tolerances. Root extension is restricted by capacity for internal O₂ movement to the apex. Sub‐optimal O₂ restricts root N uptake and translocation to the shoots, with N deficiency causing reduced shoot growth and grain yield. Although photosynthesis declines, sugars typically accumulate in shoots of waterlogged plants. Mn or Fe toxicity might occur in shoots of wheat on strongly acidic soils, but probably not more widely. Future breeding for waterlogging tolerance should focus on root internal aeration and better N‐use efficiency; exploiting the genetic diversity in wheat for these and other traits should enable improvement of waterlogging tolerance.     

A global barley panel revealing genomic signatures of breeding in modern Australian cultivars

The future of plant cultivar improvement lies in the evaluation of genetic resources from currently available germplasm. Today’s gene pool of crop genetic diversity has been shaped during domestication and more recently by breeding. Recent efforts in plant breeding have been aimed at developing new and improved varieties from poorly adapted crops to suit local environments. However, the impact of these breeding efforts is poorly understood. Here, we assess the contributions of both historical and recent breeding efforts to local adaptation and crop improvement in a global barley panel by analysing the distribution of genetic variants with respect to geographic region or historical breeding category. By tracing the impact that breeding had on the genetic diversity of Hordeum vulgare (barley) released in Australia, where the history of barley production is relatively young, we identify 69 candidate regions within 922 genes that were under selection pressure. We also show that modern Australian barley varieties exhibit 12% higher genetic diversity than historical cultivars. Finally, field-trialling and phenotyping for agriculturally relevant traits across a diverse range of Australian environments suggests that genomic regions under strong breeding selection and their candidate genes are closely associated with key agronomic traits. In conclusion, our combined data set and germplasm collection provide a rich source of genetic diversity that can be applied to understanding and improving environmental adaptation and enhanced yields.     

Molecular markers in management of ex situ PGR – A case study

Worldwide germplasm collections contain about 7.4 million accessions of plant genetic resources for food and agriculture. One of the 10 largest ex situ genebanks of our globe is located at the Leibniz Institute of Plant Genetics and Crop Plant Research in Gatersleben, Germany. Molecular tools have been used for various gene bank management practices including characterization and utilization of the germplasm. The results on genetic integrity of long-term-stored gene bank accessions of wheat (self-pollinating) and rye (open-pollinating) cereal crops revealed a high degree of identity for wheat. In contrast, the out-pollinating accessions of rye exhibited shifts in allele frequencies. The genetic diversity of wheat and barley germplasm collected at intervals of 40 to 50?years in comparable geographical regions showed qualitative rather than a quantitative change in diversity. The inter- and intraspecific variation of seed longevity was analysed and differences were detected. Genetic studies in barley, wheat and oilseed rape revealed numerous QTL, indicating the complex and quantitative nature of seed longevity. Some of the loci identified were in genomic regions that co-localize with genes determining agronomic traits such as spike architecture or biotic and abiotic stress response. Finally, a genome-wide association mapping analysis of a core collection of wheat for flowering time was performed using diversity array technology (DArT) markers. Maker trait associations were detected in genomic regions where major genes or QTL have been described earlier. In addition, new loci were also detected, providing opportunities to monitor genetic variation for crop improvement.     

北方冬麦区小麦抗旱种质资源遗传多样性分析

遗传多样性分析对于作物资源评价和利用具有重要的意义。本研究以我国北方冬麦区136份小麦抗旱种质资源为材料,分析10个农艺性状及其耐旱指数的相关性,以及抗旱种质的遗传多样性。结果表明:在雨养和灌溉条件下,穗叶距的变异系数最高,分别为42.1%和37.2%,单穗总小穗数的变异系数最低,为6.4%和5.7%;不同水分条件下,植株稳产性主要受单株穗数、有效小穗数及穗下节长的影响;性状耐旱指数的多样性指数在1.95到2.07之间变化,平均值为2.02;根据性状耐旱指数将供试材料分为7个类群,其中第I、第III类群材料表现为对水分条件不敏感,而第II类群材料更适于在干旱条件下种植。材料之间的抗旱性差异可以作为抗旱育种中亲本选配的依据。     

燕麦种质资源形态学性状的遗传多样性

【目的】燕麦种质资源遗传多样性研究不仅有助于种质资源的收集和评价,而且对燕麦生产和育种具有重要指导意义。【方法】研究对260份燕麦种质资源的20个形态学性状多样性、变异及聚类进行分析,评价其形态学性状的遗传变异水平,明确燕麦种质资源的性状特点与遗传多样性,以期为燕麦种质创新和品种改良提供依据。【结果】260份燕麦种质资源形态学性状间存在广泛的遗传多样性,质量性状的遗传多样性指数以粒色最大(1.53),芒色最小(0.76);12个数量性状呈正态分布,数量性状的遗传多样性指数以千粒重最大(2.03),有效分蘖数最小(1.22),变异系数最大的是有效分蘖数(89.02%),最小的是株高(11.19%)。根据燕麦品种(系)间各性状的遗传差异,聚类分析将供试的260份燕麦种质资源分为6类,其中种质群Ⅰ包括42份材料,可作为种用型育种目标的亲本材料;种质群Ⅱ包括31份材料,可作为选育高产饲草品种的亲本材料;种质群Ⅳ包括41份材料,可作为选育大粒专用型品种的育种材料;种质群Ⅴ包括46份材料,可作为燕麦矮化的亲本材料;而种质群Ⅲ包括46份材料,种质群Ⅵ包括54份材料,这两类种质群材料的综合性状表现不突...     

Heritage genetics for adaptation to marginal soils in barley

Future crops need to be sustainable in the face of climate change. Modern barley varieties have been bred for high productivity and quality; however, they have suffered considerable genetic erosion, losing crucial genetic diversity. This renders modern cultivars vulnerable to climate change and stressful environments. We highlight the potential to tailor crops to a specific environment by utilising diversity inherent in an adapted landrace population. Tapping into natural biodiversity, while incorporating information about local environmental and climatic conditions, allows targeting of key traits and genotypes, enabling crop production in marginal soils. We outline future directions for the utilisation of genetic resources maintained in landrace collections to support sustainable agriculture through germplasm development via the use of genomics technologies and big data.     

Population Structure of Barley Landrace Populations and Gene-Flow with Modern Varieties

Landraces are heterogeneous plant varieties that are reproduced by farmers as populations that are subject to both artificial and natural selection. Landraces are distinguished by farmers due to their specific traits, and different farmers often grow different populations of the same landrace. We used simple sequence repeats (SSRs) to analyse 12 barley landrace populations from Sardinia from two collections spanning 10 years. We analysed the population structure, and compared the population diversity of the landraces that were collected at field level (population). We used a representative pool of barley varieties for diversity comparisons and to analyse the effects of gene flow from modern varieties. We found that the Sardinian landraces are a distinct gene pool from those of both two-row and six-row barley varieties. There is also a low, but significant, mean level and population-dependent level of introgression from the modern varieties into the Sardinian landraces. Moreover, we show that the Sardinian landraces have the same level of gene diversity as the representative sample of modern commercial varieties grown in Italy in the last decades, even within population level. Thus, these populations represent crucial sources of germplasm that will be useful for crop improvement and for population genomics studies and association mapping, to identify genes, loci and genome regions responsible for adaptive variations. Our data also suggest that landraces are a source of valuable germplasm for sustainable agriculture in the context of future climate change, and that in-situ conservation strategies based on farmer use can preserve the genetic identity of landraces while allowing adaptation to local environments.     

Efficiency of cold hardiness induction by desiccation stress in four winter cereals

A number of defined desiccation treatments without low temperature exposure were able to induce freezing tolerance in 20 cultivars of winter cereals. A maximal degree of freezing tolerance was induced in epicotyls at 24°C in 24 hours at 40% relative humidity in rye and wheat, 7 days at 54% RH in barley, and 4 days at 70% RH in oats. Freezing tolerance was not correlated to water content of the plants after desiccation treatment but was related to the genetic capacity of the cultivars to frost harden. Levels of freezing tolerance induced by desiccation were similar to those induced by cold acclimation in rye and wheat, but considerably less in barley and oats. This is associated with a more rapid desiccation injury in barley and oats, precluding the completion of the hardening process.     

Cereal breeding takes a walk on the wild side

Elite cultivated crop gene pools of the Triticeae tribe (wheat, barley and rye) exhibit limited genetic diversity, raising concerns about our ability to increase or simply sustain crop yield and quality in the face of dynamic environmental and biotic threats. Although exploiting their wild relatives as a source of novel alleles is challenging, it has provided notable successes in cereal improvement for >100 years. Increasingly facile gene discovery, improved enabling technologies for genetics and breeding and a better understanding of the factors limiting practical exploitation of exotic germplasm promise to transform existing, and accelerate the development of new, strategies for efficient and directed germplasm utilization.     

Effect of Germination on Vomitoxin Level in Grain

The present investigation clarified the effect of enzymes or other substances formed during the germination process on the vomitoxin level of contaminated oats. The studies found that oats containing vomitoxin germinated very poorly; the decrease in toxins was also slight. The amount of pure vomitoxin added to toxin-free grain decreased (barley 53 %, oats 40 %, wheat 20 %) during germination (5 d). In homogenized mixture of germinated grain (2.4 and 7 d) and toxic grain no decrease in toxin amount occurred during a 1–7 day period. In contrast, when germinating toxin free grains and toxic oats in a grain mixture the toxin level decreased at first, but later rose considerably. On the basis of these results, the utilization of germination processes for the improvement of grain containing vomitoxin is of questionable value.     

Prospects for utilising plant-adaptive mechanisms to improve wheat and other crops in drought- and salinity-prone environments

Breeding for adaptation to abiotic stress is extremely challenging due to the complexity of the target environments as well as that of the stress‐adaptive mechanisms adopted by plants. While many traits have been reported in the literature, these must be considered with respect to the type of environment for which a cultivar is targeted. In theory, stress‐adaptive traits can be divided into groups whose genes and/or physiological effects are likely to be relatively independent such that when parents with contrasting traits are crossed, adaptive genes will be pyramided. Currently the following groups of candidate traits are being considered for drought adaptation in wheat: traits relating to: (i) pre‐anthesis growth, (ii) water extraction, (iii) water use efficiency, (iv) photo‐protection. A number of mechanisms relating to root function have potential to ameliorate drought stress. Hydraulic redistribution (HR) of water by roots of dryland shrubs enables even relatively small amounts of rainwater to be moved down into the soil profile actively by the root system before it evaporates from the soil surface. Another example is the symbiotic relationship of plants with mycorrhizal fungi that produce a glycoprotein that has a positive effect on soil structure and moisture characteristics. From an agronomic point of view, crop water use efficiency can be increased by exploiting the stress‐adaptive mechanism whereby leaves reduce transpiration rate in response to a chemical root signal in response to drying soil. While there is limited genetic diversity for adaptation to salinity in wheat, tolerance has been found in the ancestral genomes of polyploid wheat and their relatives associated with sodium exclusion into the xylem. Wide crossing techniques such as production of synthetic hexaploids are being exploited to tap into this source of genetic diversity. Looking further into the future, progress is being made into understanding the regulatory mechanisms that are expressed under abiotic stress to maintain cellular homeostasis, as well as in the ability to genetically transform crop plants with genes from alien species.     

Crop traits and the tolerance of wheat and barley to foliar disease

The relationship between yield loss and disease severity can differ widely between crops. This has given rise to the concept of disease tolerance, with some crops exhibiting a smaller yield loss under a given severity of disease than others. Genetic improvement to minimise yield loss under disease is an attractive goal, as it exerts little or no selection pressure on pathogen populations, and could form a useful component of durable disease management programmes. However, progress towards this end requires a thorough understanding of the phenotypic traits that influence the response of yield to disease, their genetic control and the possible trade-offs involved with other desirable agronomic characteristics. This paper examines the candidate crop traits that may confer tolerance of foliar disease in wheat and barley and reviews evidence of genetic variation in their expression. In wheat grown under the relatively low light conditions of North-West Europe, post-anthesis source (assimilate supply) and grain sink capacity (capacity for dry matter accumulation) appear to be closely balanced. Traits associated with maintaining post-anthesis radiation interception and radiation use efficiency in spite of disease may confer tolerance. The most promising traits include a larger flag leaf and compensatory increases in photosynthetic rate in non-infected parts of leaves. In barley, yield is often more strongly sink limited, and early-season disease management is required to protect the formation of potential grain sites. A wider range of potential traits may influence tolerance including compensatory adjustments in leaf growth and morphology, and differences in the sensitivity of tiller and spikelet mortality to photoassimilate supply. Different methods for quantifying tolerance are suggested depending on the trait of interest.     

Perspectives of genomics for genetic conservation of livestock

Genomics provides new opportunities for conservation genetics. Conservation genetics in livestock is based on estimating diversity by pedigree relatedness and managing diversity by choosing those animals that maximize genetic diversity. Animals can be chosen as parents for the next generation, as donors of material to a gene bank, or as breeds for targeting conservation efforts. Genomics provides opportunities to estimate diversity for specific parts of the genome, such as neutral and adaptive diversity and genetic diversity underlying specific traits. This enables us to choose candidates for conservation based on specific genetic diversity (e.g. diversity of traits or adaptive diversity) or to monitor the loss of diversity without conservation. In wild animals direct genetic management, by choosing candidates for conservation as in livestock, is generally not practiced. With dense marker maps opportunities exist for monitoring relatedness and genetic diversity in wild populations, thus enabling a more active management of diversity.     

小麦芽期和苗期耐盐性综合评价

土壤盐渍化严重影响小麦生产,提高小麦耐盐性是应对土壤盐渍化的主要途径之一,耐盐种质资源是耐盐性遗传改良的材料基础。本研究以小麦为材料,筛选芽期和苗期耐盐性鉴定评价的适宜Na Cl浓度,明确了小麦芽期耐盐性鉴定的最适Na Cl溶液浓度为1.2%,苗期耐盐性鉴定的最适土壤Na Cl浓度为0.8%。用该盐浓度胁迫处理321份小麦材料,获得芽期高耐盐材料21份,占供试材料的6.5%;苗期高耐盐材料18份,占供试材料的5.6%;芽期和苗期均为高耐盐的材料2份,分别是中作60115和冀麦一号。     

Improving salinity tolerance in crop plants: a biotechnological view

Salinity limits the production capabilities of agricultural soils in large areas of the world. Both breeding and screening germplasm for salt tolerance encounter the following limitations: (a) different phenotypic responses of plants at different growth stages, (b) different physiological mechanisms, (c) complicated genotype × environment interactions, and (d) variability of the salt-affected field in its chemical and physical soil composition. Plant molecular and physiological traits provide the bases for efficient germplasm screening procedures through traditional breeding, molecular breeding, and transgenic approaches. However, the quantitative nature of salinity stress tolerance and the problems associated with developing appropriate and replicable testing environments make it difficult to distinguish salt-tolerant lines from sensitive lines. In order to develop more efficient screening procedures for germplasm evaluation and improvement of salt tolerance, implementation of a rapid and reliable screening procedure is essential. Field selection for salinity tolerance is a laborious task; therefore, plant breeders are seeking reliable ways to assess the salt tolerance of plant germplasm. Salt tolerance in several plant species may operate at the cellular level, and glycophytes are believed to have special cellular mechanisms for salt tolerance. Ion exclusion, ion sequestration, osmotic adjustment, macromolecule protection, and membrane transport system adaptation to saline environments are important strategies that may confer salt tolerance to plants. Cell and tissue culture techniques have been used to obtain salt tolerant plants employing two in vitro culture approaches. The first approach is selection of mutant cell lines from cultured cells and plant regeneration from such cells (somaclones). In vitro screening of plant germplasm for salt tolerance is the second approach, and a successful employment of this method in durum wheat is presented here. Doubled haploid lines derived from pollen culture of F₁ hybrids of salt-tolerant parents are promising tools to further improve salt tolerance of plant cultivars. Enhancement of resistance against both hyper-osmotic stress and ion toxicity may also be achieved via molecular breeding of salt-tolerant plants using either molecular markers or genetic engineering.     

Patterns of polymorphism and linkage disequilibrium in cultivated barley

We carried out a genome-wide analysis of polymorphism (4,596 SNP loci across 190 elite cultivated accessions) chosen to represent the available genetic variation in current elite North West European and North American barley germplasm. Population sub-structure, patterns of diversity and linkage disequilibrium varied considerably across the seven barley chromosomes. Gene-rich and rarely recombining haplotype blocks that may represent up to 60% of the physical length of barley chromosomes extended across the ‘genetic centromeres’. By positioning 2,132 bi-parentally mapped SNP markers with minimum allele frequencies higher than 0.10 by association mapping, 87.3% were located to within 5 cM of their original genetic map position. We show that at this current marker density genetically diverse populations of relatively small size are sufficient to fine map simple traits, providing they are not strongly stratified within the sample, fall outside the genetic centromeres and population sub-structure is effectively controlled in the analysis. Our results have important implications for association mapping, positional cloning, physical mapping and practical plant breeding in barley and other major world cereals including wheat and rye that exhibit comparable genome and genetic features.     

小麦品种资源耐盐性鉴定

按照农业部行业标准NY/PZT001-2002,对882份小麦品种资源进行耐盐性初步鉴定,筛选出芽期耐盐性为一级的品种328份,苗期和芽期都达到中度耐盐的品种43份。这些品种中很多既具有中度或中度以上耐盐性且具有高产优质等优异特性,如小偃22、新曙光1号等,为小麦耐盐育种提供重要信息。相关分析表明,不同耐盐级别的小麦品种其芽期和苗期耐盐性并没有一致的相关关系,二者并没有可比性,在耐盐种质筛选过程中,都有其本身的意义。     

Comprehensive evaluating of wild and cultivated emmer wheat (<Emphasis Type="Italic">Triticum turgidum</Emphasis> L.) genotypes response to salt stress

Emmer wheat as the progenitor of common wheat, holds the genetic potentiality for improvement of wheat yield, quality and stress tolerance such as drought and salt. To comprehensively evaluate the salt tolerance of emmer wheat, a total of 30 traits including growth, physiology and photosynthesis related as well as K⁺ and Na⁺ content of 30 wild emmer and 14 durum wheat accessions were systematically investigated and compared between normal and saline conditions. Salt tolerance index (STI) based on multiple regression analysis of these traits was calculated and five wild emmer accessions showed high salt tolerance, which could be used as valuable resource for wheat salt tolerance improvement. Furthermore, wild emmer genotypes showed wider trait performance variation compared to durum wheat, indicating the higher genetic diversity in wild emmer wheat. Then, shoot Na⁺ content, shoot K⁺/Na⁺ ratio, root length and root surface area were identified as suitable indexes for salt tolerance evaluation. Na⁺ exclusion mechanism was found to be playing an important role in response to salt stress in emmer wheat. The salt tolerance in emmer wheat was systematically characterized here, which not only provided the elite germplasm for wheat improvement, but also provided the efficient method and some useful indexes for salt tolerance assessing.     

Identifying the critical period for waterlogging on yield and its components in wheat and barley

Background and aims

Crop tolerance to waterlogging depends on factors such as species sensitivity and the stage of development that waterlogging occurs. The aim of this study was to identify the critical period for waterlogging on grain yield and its components, when applied during different stages of crop development in wheat and barley.

Methods

Two experiments were carried out (E1: early sowing date, under greenhouse; E2: late sowing date, under natural conditions). Waterlogging was imposed during 15–20 days in 5 consecutive periods during the crop cycle (from Leaf 1 emergence to maturity).

Results

The greatest yield penalties occurred when waterlogging was applied from Leaf 7 appearance on the main stem to anthesis (from 34 to 92 % of losses in wheat, and from 40 to 79 % in barley for E1 and E2 respectively). Waterlogging during grain filling reduced yield to a lesser degree. In wheat, reductions in grain number were mostly explained by reduced grain number per spike while in barley, by variations in the number of spikes per plant.

Conclusions

The time around anthesis was identified as the most susceptible period to waterlogging in wheat and barley. Exposing the crop to more stressful conditions, e.g. delaying sowing date, magnified the negative responses to waterlogging, although the most sensitive stage (around anthesis) remained unchanged.