Yield-trait performance landscapes: from theory to application in breeding maize for drought tolerance

The effectiveness of breeding strategies to increase drought resistance in crops could be increased further if some of the complexities in gene-to-phenotype (G → P) relations associated with epistasis, pleiotropy, and genotype-by-environment interactions could be captured in realistic G → P models, and represented in a quantitative manner useful for selection. This paper outlines a promising methodology. First, the concept of landscapes was extended from the study of fitness landscapes used in evolutionary genetics to the characterization of yield-trait-performance landscapes for agricultural environments and applications in plant breeding. Second, the E(NK) model of trait genetic architecture was extended to incorporate biophysical, physiological, and statistical components. Third, a graphical representation is proposed to visualize the yield-trait performance landscape concept for use in selection decisions. The methodology was demonstrated at a particular stage of a maize breeding programme with the objective of improving the drought tolerance of maize hybrids for the US Western Corn-Belt. The application of the framework to the genetic improvement of drought tolerance in maize supported selection of Doubled Haploid (DH) lines with improved levels of drought tolerance based on physiological genetic knowledge, prediction of test-cross yield within the target population of environments, and their predicted potential to sustain further genetic progress with additional cycles of selection. The existence of rugged yield-performance landscapes with multiple peaks and intervening valleys of lower performance, as shown in this study, supports the proposition that phenotyping strategies, and the directions emphasized in genomic selection can be improved by creating knowledge of the topology of yield-trait performance landscapes.     

Captive Breeding Programs Based on Family Groups in Polyploid Sturgeons

In species with long life cycles and discontinuous availability of individuals to reproduction, implementing a long-term captive breeding program can be difficult or impossible. In such cases, managing diversity among familiar groups instead of individuals could become a suitable approach to avoid inbreeding and increase the possibility to accomplish a breeding scheme. This is the case of several sturgeon species including the Adriatic sturgeon, whose recovery depends on the management of a few captive stocks directly descended from the same group of wild parents. In the present study, relatedness among 445 potential breeders was inferred with a novel software for pedigree reconstruction in tetraploids (“BreedingSturgeons”). This information was used to plan a breeding scheme considering familiar groups as breeding units and identifying mating priorities. A two-step strategy is proposed: a short-term breeding program, relying on the 13 remaining F0 individuals of certain wild origin; and a long-term plan based on F1 families. Simulations to evaluate the loss of alleles in the F2 generation under different pairing strategies and assess the number of individuals to breed, costs and logistical aquaculture constraints were performed. The strategy proposed is transferable to the several other tetraploid sturgeon species on the brink of extinction.     

Advances and prospects: biotechnologically improving crop water use efficiency

Bio-water saving can be defined as the reduction of crop water consumption employing biological measures. This is the focus of efforts to save water in agriculture. Different levels of water-use efficiency (WUE) have been developed. The genetic diversity of WUE has been confirmed in several crops. WUE is the basis of bio-watering and physiological WUE is the key. The degree to develop physiological WUE potential decides the performance of bio-watering in the field. During this process, fine management is important. Thus bio-watering is closely related to WUE. Crop WUE has improved and evolved as a result of breeding programs. Many WUE genes have been located in different genomic and aneuploid materials and have been mapped by various molecular markers in a number of crops. Two genes, (Erecta and alx8), which control water use efficiency; have been cloned in Arabidopsis thaliana. Eleven WUE genes have been identified by microarray analysis. Six genes associated with drought resistance and photosynthesis have been transfered into crops which have resulted in improving WUE and drought resistance. WUE is important on the basis of functional identification of more drought resistant gene resources. The popularity on the industrial-scale of transgenic plants is still in its infancy and one of the reasons for this is the lack of knowledge regarding molecular mechanisms and it is a very immature technology. Enhanced agricultural practices and the theoretical aspects of improving crop WUE have been developed and are discussed in this review paper. Rapid progress will be made in bio-water savings and that crop WUE can be substantially improved under both favorable and unfavorable water-limited environments. This will be achieved by a combination of traditional breeding techniques and the introduction of modern biotechnology.     

Island-Model Genomic Selection for Long-Term Genetic Improvement of Autogamous Crops

Acceleration of genetic improvement of autogamous crops such as wheat and rice is necessary to increase cereal production in response to the global food crisis. Population and pedigree methods of breeding, which are based on inbred line selection, are used commonly in the genetic improvement of autogamous crops. These methods, however, produce a few novel combinations of genes in a breeding population. Recurrent selection promotes recombination among genes and produces novel combinations of genes in a breeding population, but it requires inaccurate single-plant evaluation for selection. Genomic selection (GS), which can predict genetic potential of individuals based on their marker genotype, might have high reliability of single-plant evaluation and might be effective in recurrent selection. To evaluate the efficiency of recurrent selection with GS, we conducted simulations using real marker genotype data of rice cultivars. Additionally, we introduced the concept of an “island model” inspired by evolutionary algorithms that might be useful to maintain genetic variation through the breeding process. We conducted GS simulations using real marker genotype data of rice cultivars to evaluate the efficiency of recurrent selection and the island model in an autogamous species. Results demonstrated the importance of producing novel combinations of genes through recurrent selection. An initial population derived from admixture of multiple bi-parental crosses showed larger genetic gains than a population derived from a single bi-parental cross in whole cycles, suggesting the importance of genetic variation in an initial population. The island-model GS better maintained genetic improvement in later generations than the other GS methods, suggesting that the island-model GS can utilize genetic variation in breeding and can retain alleles with small effects in the breeding population. The island-model GS will become a new breeding method that enhances the potential of genomic selection in autogamous crops, especially bringing long-term improvement.     

Population-Wide Failure to Breed in the Clark’s Nutcracker (Nucifraga columbiana)

In highly variable environments, conditions can be so stressful in some years that entire populations forgo reproduction in favor of higher likelihood of surviving to breed in future years. In two out of five years, Clark’s nutcrackers (Nucifraga Columbiana) in the Greater Yellowstone Ecosystem exhibited population-wide failure to breed. Clark’s nutcrackers at the study site experienced substantial interannual differences in food availability and weather conditions, and the two nonbreeding years corresponded with low whitebark pine (Pinus albicaulis) cone crops the previous autumn (≤ an average of 8 ± 2 cones per tree versus ≥ an average of 20 ± 2 cones per tree during breeding years) and high snowpack in early spring (≥ 61.2 ± 5.5 cm versus ≤ 51.9 ± 4.4 cm during breeding years). The average adult body condition index during the breeding season was significantly lower in 2011 (-1.5 ± 1.1), a nonbreeding year, as compared to 2012 (6.2 ± 2.0), a breeding year. The environmental cues available to the birds prior to breeding, specifically availability of cached whitebark pine seeds, may have allowed them to predict that breeding conditions would be poor, leading to the decision to skip breeding. Alternatively, the Clark’s nutcrackers may have had such low body energy stores that they chose not to or were unable to breed. Breeding plasticity would allow Clark’s nutcrackers to exploit an unpredictable environment. However, if large-scale mortality of whitebark pines is leading to an increase in the number of nonbreeding years, there could be serious population-level and ecosystem-wide consequences.     

Harvesting the Promising Fruits of Genomics: Applying Genome Sequencing Technologies to Crop Breeding

Next generation sequencing (NGS) technologies are being used to generate whole genome sequences for a wide range of crop species. When combined with precise phenotyping methods, these technologies provide a powerful and rapid tool for identifying the genetic basis of agriculturally important traits and for predicting the breeding value of individuals in a plant breeding population. Here we summarize current trends and future prospects for utilizing NGS-based technologies to develop crops with improved trait performance and increase the efficiency of modern plant breeding. It is our hope that the application of NGS technologies to plant breeding will help us to meet the challenge of feeding a growing world population.

This article is part of the PLOS Biology Collection “The Promise of Plant Translational Research.”

     

Do soil depth and plant community composition interact to modify the resistance and resilience of grassland ecosystem functioning to drought?

While the effect of drought on plant communities and their associated ecosystem functions is well studied, little research has considered how responses are modified by soil depth and depth heterogeneity. We conducted a mesocosm study comprising shallow and deep soils, and variable and uniform soil depths, and two levels of plant community composition, and exposed them to a simulated drought to test for interactive effects of these treatments on the resilience of carbon dioxide fluxes, plant functional traits, and soil chemical properties. We tested the hypotheses that: (a) shallow and variable depth soils lead to increased resistance and resilience of ecosystem functions to drought due to more exploitative plant trait strategies; (b) plant communities associated with intensively managed high fertility soils, will have more exploitative root traits than extensively managed, lower fertility plant communities. These traits will be associated with higher resistance and resilience to drought and may interact with soil depth and depth heterogeneity to amplify the effects on ecosystem functions. Our results showed that while there were strong soil depth/heterogeneity effects on plant‐driven carbon fluxes, it did not affect resistance or resilience to drought, and there were no treatment effects on plant‐available carbon or nitrogen. We did observe a significant increase in exploitative root traits in shallow and variable soils relative to deep and uniform, which may have resulted in a compensation effect which led to the similar drought responses. Plant community compositions representative of intensive management were more drought resilient than more diverse “extensive” communities irrespective of soil depth or soil depth heterogeneity. In intensively managed plant communities, root traits were more representative of exploitative strategies. Taken together, our results suggest that reorganization of root traits in response to soil depth could buffer drought effects on ecosystem functions.     

抗旱相关基因在烟草中的应用研究进展

干旱是影响烟草正常生长、发育、产量和烟叶品质的一个重要逆境因子。在干旱胁迫下,植物体内会通过激发一些抗旱基因的表达来增强植物的抗旱能力。目前,很多抗旱相关的功能蛋白基因和调控蛋白基因已被克隆并在烟草中实现了遗传转化,外源抗旱基因的表达提高了转基因烟草的抗旱能力。抗旱基因的克隆为烟草抗旱新品种的培育奠定了良好的分子基础,系统深入地研究抗旱相关基因在干旱胁迫条件下的表达与调控,可为通过基因工程手段提高烟草的抗旱能力开辟新途径,同时也能为其他农作物的抗旱分子育种和品种改良提供基因资源。     

Evidence-based policymaking when evidence is incomplete: The case of HIV programme integration

Jan Hontelez and co-authors discuss the use of different types of evidence to inform HIV program integration.

Summary points

• Sustainable Development Goal 3 aims to “ensure healthy lives and promote well-being for all at all ages” and has set a target of achieving global universal health coverage, representing a major policy shift away from mostly disease-specific “vertical programmes”.
• While health service integration can be a promising strategy to improve healthcare coverage, health outcomes, and efficiency, the exact impact of integration in different settings is hard to predict, and policy makers need to choose from a large variety of integration strategies and opportunities with varying levels of scientific evidence.
• Using the case of health service integration for HIV in low- and middle-income countries, we outline implementation strategies for integration opportunities with lacking or scarce high-level causal evidence, based on existing frameworks and methodologies from within and beyond healthcare and implementation science.
• Proper use of scientific evidence in other contexts requires adequate and systematic assessments of the transportability of an intervention. Several methods exist that allow for judging transferability and comprehensively identifying key context-specific indicators across studies that can affect the reported impact of interventions.
• When (transferable) evidence is absent, we propose that by drawing on well-established design and implementation methodologies—underpinned by ongoing learning and iterative improvement of local service delivery strategies—countries could substantially improve decision-making even in the absence of scientific evidence.
• Reaching the goal of making the HIV response an integral part of a larger, universal, people-centred health system that meets the needs and requirements of citizens can be facilitated by applying lessons learned from implementation science and novel design methodologies.

     

干旱,半干旱地区作物育种的困惑与出路

粮食问题主要取决于一年生谷类作物产量。作物产量低而不稳的原因主要是病虫害及各种胁迫生境,其中干旱缺水为最大的产量限制因素,提高作物生产力的途径有二：其一是改善作物的生长环境,其二是通过育种手段选育在各肿胁迫环境中具有优良表现的基因型（品种）。矮秆化育种手段使水肥充裕区小麦产量有显著的提高,是通过提高收获指数获得的。干旱、半干旱地区育种却未能获得显著效果,要提高干旱、半干旱地区小麦育种的成效,对干旱     

Capturing in-field root system dynamics with RootTracker

Optimizing root system architecture offers a promising approach to developing stress tolerant cultivars in the face of climate change, as root systems are critical for water and nutrient uptake as well as mechanical stability. However, breeding for optimal root system architecture has been hindered by the difficulty in measuring root growth in the field. Here, we describe the RootTracker, a technology that employs impedance touch sensors to monitor in-field root growth over time. Configured in a cylindrical, window shutter-like fashion around a planted seed, 264 electrodes are individually charged multiple times over the course of an experiment. Signature changes in the measured capacitance and resistance readings indicate when a root has touched or grown close to an electrode. Using the RootTracker, we have measured root system dynamics of commercial maize (Zea mays) hybrids growing in both typical Midwest field conditions and under different irrigation regimes. We observed rapid responses of root growth to water deficits and found evidence for a “priming response” in which an early water deficit causes more and deeper roots to grow at later time periods. Genotypic variation among hybrid maize lines in their root growth in response to drought indicated a potential to breed for root systems adapted for different environments. Thus, the RootTracker is able to capture changes in root growth over time in response to environmental perturbations.

RootTracker, a technology that employs impedance touch sensors, enables monitoring in-field root growth over time.     

Drought Stress in Wheat during Flowering and Grain-filling Periods

Drought is a major environmental stress threatening wheat productivity worldwide. Global climate models predict changed precipitation patterns with frequent episodes of drought. Although drought impedes wheat performance at all growth stages, it is more critical during the flowering and grain-filling phases (terminal drought) and results in substantial yield losses. The severity and duration of the stress determine the extent of the yield loss. The principal reasons for these losses are reduced rates of net photosynthesis owing to metabolic limitations—oxidative damage to chloroplasts and stomatal closure—and poor grain set and development. A comprehensive understanding of the impact of terminal drought is critical for improving drought resistance in wheat, with marker-assisted selection being increasingly employed in breeding for this resistance. The limited success of molecular breeding and physiological strategies suggests a more holistic approach, including interaction of drought with other stresses and plant morphology. Furthermore, integration of physiological traits, genetic and genomic tools, and transgenic approaches may also help to improve resistance against drought in wheat. In this review, we describe the influence of terminal drought on leaf senescence, carbon fixation, grain set and development, and explain drought resistance mechanisms. In addition, recent developments in integrated approaches such as breeding, genetics, genomics, and agronomic strategies for improving resistance against terminal drought in wheat are discussed.     

Pre-conditioning the epigenetic response to high vapor pressure deficit increases the drought tolerance of Arabidopsis thaliana

Epigenetic modification of the genome via cytosine methylation is a dynamic process that responds to changes in the growing environment. This modification can also be heritable. The combination of both properties means that there is the potential for the life experiences of the parental generation to modify the methylation profiles of their offspring and so potentially to “pre-condition” them to better accommodate abiotic conditions encountered by their parents. We recently identified high vapor pressure deficit (vpd)-induced DNA methylation at 2 gene loci in the stomatal development pathway and an associated reduction in leaf stomatal frequency.¹ Here, we test whether this epigenetic modification pre-conditioned parents and their offspring to the more severe water stress of periodic drought. We found that 3 generations of high vpd-grown plants were better able to withstand periodic drought stress over 2 generations. This resistance was not directly associated with de novo methylation of the target stomata genes, but was associated with the cmt3 mutant’s inability to maintain asymmetric sequence context methylation. If our finding applies widely, it could have significant implications for evolutionary biology and breeding for stressful environments.     

作物抗旱相关分子标记及其辅助选择的研究进展

分子标记辅助选择育种给作物抗旱育种提供了新的途径。本介绍了国内外在小麦、玉米、水稻、大豆等重要农作物抗旱相关分子标记方面的研究进展。对作物抗旱相关QTL分子标记辅助育种进行了探讨,并对其发展策略提出了一些思考。     

CRISPR/Cas‐based tools for the targeted control of plant viruses

Plant viruses are known to infect most economically important crops and pose a major threat to global food security. Currently, few resistant host phenotypes have been delineated, and while chemicals are used for crop protection against insect pests and bacterial or fungal diseases, these are inefficient against viral diseases. Genetic engineering emerged as a way of modifying the plant genome by introducing functional genes in plants to improve crop productivity under adverse environmental conditions. Recently, new breeding technologies, and in particular the exciting CRISPR/Cas (clustered regularly interspaced short palindromic repeats/CRISPR‐associated proteins) technology, was shown to be a powerful alternative to engineer resistance against plant viruses, thus has great potential for reducing crop losses and improving plant productivity to directly contribute to food security. Indeed, it could circumvent the “Genetic modification” issues because it allows for genome editing without the integration of foreign DNA or RNA into the genome of the host plant, and it is simpler and more versatile than other new breeding technologies. In this review, we describe the predominant features of the major CRISPR/Cas systems and outline strategies for the delivery of CRISPR/Cas reagents to plant cells. We also provide an overview of recent advances that have engineered CRISPR/Cas‐based resistance against DNA and RNA viruses in plants through the targeted manipulation of either the viral genome or susceptibility factors of the host plant genome. Finally, we provide insight into the limitations and challenges that CRISPR/Cas technology currently faces and discuss a few alternative applications of the technology in virus research.     

Beyond landraces: developing improved germplasm resources for underutilized species – a case for Bambara groundnut

The potential for underutilized crops (also known as minor, neglected or orphan crops) to improve food and nutrition security has been gaining prominence within the research community in recent years. This is due to their significance for diversified agricultural systems which is a necessary component of future agriculture to address food and nutritional security concerns posed by changing climate and a growing world population. Developing workable value chain systems for underutilized crop species, coupled with comparative trait studies with major crops, potentially allows us to identify suitable agricultural modalities for such species. Bambara groundnut (Vigna subterranea L. Verdc.), an underutilized leguminous species, is of interest for its reported high levels of drought tolerance in particular, which contributes to environmental resilience in semi-arid environments. Here, we present a synopsis of suitable strategies for the genetic improvement of Bambara groundnut as a guide to other underutilized crop species. Underutilized crops have often been adapted over thousands of years in particular regions by farmers and largely still exist as landraces with little or no genetic knowledge of key phenotypic traits. Breeding in these species is fundamentally different to breeding in major crops, where significant pedigree structures and history allow highly directed improvement. In this regard, deploying new integrated germplasm development approaches for variety development and genetic analysis, such as multi-parent advance generation inter-crosses (MAGIC), within breeding programmes of underutilized species will be important to be able to fully utilize such crops.     

Overexpression of a Harpin-encoding gene hrf1 in rice enhances drought tolerance

Harpin proteins are well known as eliciters that induce multiple responses in plants, such as systemic acquired resistance, hypersensitive response, enhancement of growth, resistance to the green peach aphid, and tolerance to drought. Overexpression of Harpin-encoding genes enhances plant resistance to diseases in tobacco, rice, rape, and cotton; however, it is not yet known whether the expression of Harpin-encoding genes in vivo improves plant tolerance to abiotic stresses. The results of this study showed that overexpression of a Harpin-encoding gene hrf1 in rice increased drought tolerance through abscisic acid (ABA) signalling. hrf1- overexpression induces an increase in ABA content and promotes stomatal closure in rice. The hrf1 transgenic rice lines exhibited a significant increase in water retention ability, levels of free proline and soluble sugars, tolerance to oxidative stress, reactive oxygen species-scavenging ability, and expression levels of four stress-related genes, OsLEA3-1, OsP5CS, Mn-SOD, and NM_001074345, under drought stress. The study confirmed that hrf1 conferred enhanced tolerance to drought stress on transgenic crops. These results suggest that Harpins may offer new opportunities for generating drought resistance in other crops.     

Sex and Breed-Dependent Organ Development and Metabolic Responses in Foetuses from Lean and Obese/Leptin Resistant Swine

The present study aimed to determine the effects of breed and sex on growth patterns and metabolic features of advanced-pregnancy foetuses exposed to the same environmental conditions. Thus, at Day 62 of pregnancy, swine foetuses from an obese breed with leptin resistance (Iberian breed) were compared to lean crossbred foetuses (25% Large White ×25% Landrace ×50% Pietrain). There were differential developmental patterns in foetuses with leptin resistance, mainly a higher relative weight of the brain resembling “brain-sparing effect”. Prioritization of brain growth may be protective for the adequate growth and postnatal survival of the Iberian individuals, an ancient breed reared in extensive semi-feral conditions for centuries. There were also clear sex-related differences in foetal development and metabolism in the Iberian breed. Female Iberian foetuses were similar in size and weight to male littermates but had a significantly higher relative liver to body weight ratio resembling “liver-sparing effect” and a trend for a higher relative intestine to body ratio. Moreover, the availability of triglycerides, cholesterol and IL-6 in female Iberian foetuses was similar to that of lean crossbred foetuses. Overall, these features may favour a better postnatal survival and development of females, the sex more critical for the species survival. These findings set the basis for future translational studies aimed at increasing the knowledge on the interaction between genetic and environmental factors in the early programming of the adult phenotype.     

Defining selection criteria to improve yield under drought

The many selection criteria that have been proposed to increase drought resistance of our crops have had little, if any, impact on improving crop yields in dry environments. There are several likely reasons for this lack of success. Some of these are: (i) criteria proposed have been related more to survival mechanisms under drought than to productivity, (ii) criteria are inappropriate to the target environment, and (iii) criteria are temporal and are therefore likely to have minimal impact on growth and yield over the entire lifecycle. Another important reason is that breeders have not been convinced the proposed criteria will be successful as they are too difficult to measure. On the other hand, empirical breeding programmes to improve yield under drought have been successful. Surprisingly, some of the greatest successes have been achieved by breeding in environments where water is non-limiting. This paper reviews breeding approaches to improve yield under drought. It focuses on critical factors that must be considered to identify likely plant attributes that can be targeted. These factors, their link with yield, the nature of the target environment, the level of organisation where the trait is expressed are discussed. Three quite different examples are given to emphasize the above considerations and which show substantial promise in targeting traits to improve yield under drought. They are drought at flowering, improving transpiration efficiency and improving early leaf area development.     

Water stress resilient cereal crops: Lessons from wild relatives

Cereal crops are significant contributors to global diets. As climate change disrupts weather patterns and wreaks havoc on crops, the need for generating stress-resilient, high-yielding varieties is more urgent than ever. One extremely promising avenue in this regard is to exploit the tremendous genetic diversity expressed by the wild ancestors of current day crop species. These crop wild relatives thrive in a range of environments and accordingly often harbor an array of traits that allow them to do so. The identification and introgression of these traits into our staple cereal crops can lessen yield losses in stressful environments. In the last decades, a surge in extreme drought and flooding events have severely impacted cereal crop production. Climate models predict a persistence of this trend, thus reinforcing the need for research on water stress resilience. Here we review: (i) how water stress (drought and flooding) impacts crop performance; and (ii) how identification of tolerance traits and mechanisms from wild relatives of the main cereal crops, that is, rice, maize, wheat, and barley, can lead to improved survival and sustained yields in these crops under water stress conditions.