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• W2914989087 abstract "Among the approximately 30 genera and 360 species belonging to the tribe Triticeae (subfamily Pooideae, family Poaceae), wheat (Triticum ssp.) and barley (Hordeum vulgare L.) are economically the most important temperate cereal crops. On a global scale in 2016, wheat was ranked second (749 million tons) and barley fourth (141 million tons) in terms of world production (FAO STAT 2018). A considerable amount of our daily carbohydrates and proteins is contributed by both species. Cereals and bread are the major source of energy for all age groups, providing ∼30% for adults in developed countries, while it can be up to 80% in developing countries (Shewry and Hey 2015). Despite their global importance for food security and supply, yield gains of wheat and barley have remained rather moderate or stagnant within the last two decades, specifically in the high-yielding areas of the world such as Europe (Ray et al. 2012; Grassini et al. 2013). Clearly, this alarming trend has to change, to ensure that future projected food demands can be met in a sustainable manner (Tilman et al. 2011; Garnett et al. 2013). One part of the equation of improving cereal yields is in the way they are bred. Higher genetic gains through accelerated, genomics-informed “smart”-breeding methodologies requires whole-genome or genome-wide information. However, the large and highly repetitive genomes of diploid barley (∼5 Giga bp (Gbp)) and hexaploid bread wheat (∼16 Gbp; T. aestivum L.) greatly hampered previous accessibility to marker or gene information. Only the advent of the latest sequencing technologies over the last ∼1.5 decades in combination with very dedicated scientists from many parts of the world facilitated to ultimately conquer even such large and complex grass genomes. Nowadays physical maps and pseudomolecules are available, providing the first possibility to enter the post-genomics era in wheat (IWGSC 2018) and barley (Mascher et al. 2017) research. Despite these investments in wheat genomics, wheat is notably under-researched and shows a substantial research deficit in comparison to other starch-rich crops, such as maize or rice (Manners and van Etten 2018). To reverse this trend and to attract more research in the areas of wheat and barley biology we would like to highlight exciting current and future research topics in this special issue of the Journal of Integrative Plant Biology. Besides the large-scale genome sequences, the relatively long generation time of 5–6 months (from grain-to-grain) for spring-types is often used as an excuse not to work on wheat or barley. However, after the invention of “speed breeding”, generation times have been substantially reduced to ∼2 months (Ghosh et al. 2018; Watson et al. 2018), which is quite comparable with non-crop, model species. Moreover, “speed breeding” technology can not only be applied for faster population development but certainly also for generation advancement of transgenic plants. Most importantly, wheat and barley are similarly as amenable to gene editing methods as other crop and non-crop species (Kumlehn et al. 2018), providing the important genetic basis for efficient crop breeding and functional gene or allele characterization. The publicly available wheat and barley genomic resources will also be instrumental in identifying historical crop evolutionary and domestication events. In this special issue, Haas et al. (2019) provide insights into advances in sequencing technologies which have and will contribute substantially towards answering unresolved questions in domestication genomics. Authors comprehensively discuss domestication traits, and how they contributed to range expansion. The genomics of crop wild relatives are viewed in the light of allele mining and how new alleles can be introgressed into current breeding programs. In another study, similar authors showed how genomics-assisted genebank management and the utilization of germplasm collections in barley link natural variation to human selection during crop evolution (Milner et al. 2018). Cereal crops, such as wheat and barley, are usually grown as monocultures in fields or strips to enable synchronized maturation and combine harvesting. Dense stands of kin plants in a canopy are therefore the preferred growth situation. The canopy architecture of cultivars is hence a decisive feature of how plants cope with inter-plant competition. While, for example, shade-avoidance responses in cereal crops are hardly understood, the development of so-called “cooperative plant communities” with improved yield potential is still far from being realized (Donald 1968; Denison 2012). As leaf and tiller development are pivotal aspects of canopy and shoot architecture, Shaaf et al. (2019) highlight the most recent genetic achievements for these structural traits. Specifically, estimating and measuring total shoot biomass, leaf area or tiller number was always a struggle in the pre-phenomics era; yet, recent advancements in 2D and 3D imaging and phenotyping techniques have facilitated reliable trait measurements. An equally important but almost completely neglected research area, in the context of canopy or shoot architecture, is culm length or plant height. Lodging of too tall plants seemed solved, at least in wheat, by the introduction of so-called semidwarfing genes during the Green Revolution in the late 1960s which resulted in wheat cultivars with shorter culms which were sturdy enough to carry the grains and extra yield due to enhanced assimilate partitioning (Hedden 2003). However, the latest findings in barley showed that the usage of such alleles is not without trade-off for yield-relevant organ size (Serrano-Mislata et al. 2017). Therefore, the prevention of lodging in barley is even less satisfyingly resolved, that is why McKim (2019) tackles this important issue from a developmental biologist's perspective in her review article. She convincingly shows how little we actually know about culm growth, development and their underlying mechanisms of controlling subapical and intercalary meristem proliferation in the grasses, including control of the cell division plane and temporal dynamics of differentiation. Her seminal review article may aid in directing more attention towards this unattended research topic, in particular in terms of achieving a better control over culm internode growth and shoot height. Finding the right timing for phase transition and starting reproductive development is a crucial life decision in all plants, including crops (Blumel et al. 2015; Gol et al. 2017). Pre-anthesis reproductive development in wheat and barley encompasses inflorescence, that is spike, initiation, differentiation and growth, culminating in carpel/ovule fertilization (anthesis). This pre-anthesis time from sowing until anthesis is usually considered as “flowering time” and its duration often decides over suitable adaptation and yield potential of the crop. In this special issue, Liu et al. (2019) report about TaZIM-A1, an atypical GATA-like transcriptional repressor of flowering time in wheat. Detailed haplotype and association analyses revealed that TaZIM-A1 and its homoeolog, TaZIM-B1, have undergone strong positive selection during modern wheat breeding, most probably as an effect of earlier flowering and improved 1,000 kernel weight. Moreover, these authors show how allele-specific markers can be utilized for wheat cultivar improvement. Two other reviews appraise recent progress in our understanding of the genes controlling inflorescence development in wheat and barley, with the goal of highlighting the significance of improvements in developmental biology for manipulating the agronomic performance of crop plants. While Koppolu and Schnurbusch (2019) focus in particular on two developmental genetic pathways, namely: (i) the row-type pathway in Hordeum species: and (ii) a simplified model describing a probable mechanism of how Triticeae species form the “unbranched spike”, Gauley and Boden (2019) discuss key domestication-related traits of inflorescence architecture that contributed significantly to our understanding of the molecular processes that regulate inflorescence development in the Triticeae. Both reviews propose, fairly in accordance with similar points made by Trevaskis (2018), to widen the genetic basis for modified inflorescence architectures to provide valuable improvements in the genetic control of spikelet and floret development as well as fertility. Two fitting examples in the latter context were shown for the interactions between row-type genes in barley (Zwirek et al. 2018); and the discovery of the GRAIN NUMBER INCREASE1 (GNI1) locus in wheat, which proves to be an ortholog of barley Six-rowed spike1 (VRS1), an inhibitor of floret development (Sakuma et al. 2019). The presence of such genes that actively inhibit the plant's yield potential may change the long-held paradigm that mostly the shortage of assimilates and/or competition with other plant organs were limiting grain set in wheat and barley plants. Accepting and embracing this idea may lead to the discovery of many other novel loci in the future and might similarly help improvement in grain yields. While floret and spikelet fertility are right at the heart of grain number determination, proper assimilate supply towards florets/grains is almost a complete mystery in our cereal crops. Although there is a useful approximation about how nutrients and metabolites are transported and allocated within and along the rice culm/stem (Yamaji and Ma 2014), knowledge in wheat and barley is absent. This is even more worrisome for the anatomy and morphogenesis of the spike, specifically its main axis—the rachis. The rachis typically consists of small, distichously arranged phytomeric units (nodes and internodes) and provides the actual base for spikelet attachment. Considering that the rachis forms the genuine foundation of the spike, but similarly represents a “transport bottleneck” for all spikelets and florets through which all assimilates have to pass, it is quite disturbing how little we actually know about rachis patterning, cell differentiation, vascularization or cell types. Which role the rachis might play during source-sink relationships is still completely unexplored, but so are other source-sink relations between various plant organs. Similar things can be said about long-distance signaling between organs (e.g. root-shoot, tiller-main culm), including hormonal regulations and patterning. Yet, first tentative steps in this direction have already been taken and have found rather coarse hormonal distributions along the developing spike in barley (Youssef et al. 2017). Admittedly, this is only the start and there remain many opportunities for future exploration and discoveries. While pre-anthesis reproductive development mainly deals with the establishment and morphogenesis of floral organs, post-anthesis reproductive development is predominantly about filling the developing grain. Thus, grain size or weight is one important component of final grain yield, whereas grain shape represents an essential grain quality character. In this special issue, Shirley et al. (2019) therefore highlight early developmental cues leading to gynoecium and gametophyte formation and endosperm cellularization, with a specific emphasis on auxin signaling during ovule and grain development. Authors thereby provide interesting transcriptional signatures of the auxin-signaling pathway during barley caryopsis development with the view of improving grain. While caryopsis tissue differentiation and cell specifications are crucial during the early grain-filling period, assimilate supply and distribution is more important in the latter stages. For example, how sugars are spatio-temporally dispersed in the developing barley grain could be elegantly observed for the first time using latest micro-imaging technologies (Peukert et al. 2014; Guendel et al. 2018). A highly interesting sugar signaling component, reflecting the plant cell's carbon status, is trehalose-6-phosphate (Tre6P). Importantly, increased Tre6P in wheat promotes biosynthetic pathways associated with grain yield, such as starch synthesis and grain size; whereas decreasing Tre6P induces resource mobilization and alterations in sucrose allocation under stress conditions (Griffiths et al. 2016; Paul et al. 2018). Like all other yield components, grain weight or size is a polygenic character and behaves in a quantitative manner. Therefore, Brinton and Uauy (2019) argue in their review article that a “reductionist approach” is required as a starting point to better understand the gene networks and mechanisms regulating individual sub-components underlying yield. They further suggest that, by understanding these precise mechanisms by which individual genes regulate individual yield components, one will be better positioned to understand, and potentially uncouple, negative correlations between yield components. As one outstanding example in this context, authors discuss the wheat ortholog of rice GRAIN WEIGHT2 (GW2) gene, in which deleterious mutations in multiple homoeologs of TaGW2 had positive additive effects on 1 000 grain weight compared to the wild type (single = +5.3%, double = +10.5%, triple = +20.7%). As this special issue cannot be all-encompassing, many other promising and important future research areas, for example for mitigating the agricultural footprint in the ecosystem, such as improved nutrient use efficiency (Distelfeld et al. 2014; Hawkesford 2017; Avin-Wittenberg et al. 2018), root architecture (Salvi 2017), or root-microbe interactions specifically for nitrogen-fixing cereals (Rogers and Oldroyd 2014; Mus et al. 2016), are not itemized. Similarly, the manipulation of recombination at free-will in our cereal crops (Lambing and Heckmann 2018), or a better molecular understanding of heterosis (Schrag et al. 2018; Seifert et al. 2018) and the introduction of a cost-effective and reliable hybrid seed production system in cereals (Muehleisen et al. 2013; Whitford et al. 2013; Tucker et al. 2017) have not even been touched upon. Thus, many global and societal challenges, but also truly exciting research opportunities, still lie ahead of us. Our successful future is primarily determined by our current actions, and we, as a research community, must make ourselves heard in order to attract the best young talents required to solve these urgent challenges in the area of food supply and food security. To get there, we must be brave and enter new territories to shift known frontiers. The author would like to thank Liese Schnurbusch for critically reading previous versions of the manuscript. T.S. received financial support from the HEISENBERG Program of the German Research Foundation (DFG), grant no. SCHN 768/8-1, and IPK core budget. Thorsten Schnurbusch, Special Issue Editor thor@ipk-gatersleben.de Independent HEISENBERG-Research Group Plant Architecture, Leibniz Institute of Plant Genetics and Crop Plant Research (IPK), Corrensstr. 3, OT Gatersleben, 06466 Seeland, Germany" @default.
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• W2914989087 title "Wheat and barley biology: Towards new frontiers" @default.
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