Perspective: 50 years of plant chromosome biology.

The past 50 years has been the greatest era of plant science discovery, and most of the discoveries have emerged from or been facilitated by our knowledge of plant chromosomes. At last we have descriptive and mechanistic outlines of the information in chromosomes that programs plant life. We had almost no such information 50 years ago when few had isolated DNA from any plant species. The important features of genes have been revealed through whole genome comparative genomics and testing of variants using transgenesis. Progress has been enabled by the development of technologies that had to be invented and then become widely available. Arabidopsis (Arabidopsis thaliana) and rice (Oryza sativa) have played extraordinary roles as model species. Unexpected evolutionary dramas were uncovered when learning that chromosomes have to manage constantly the vast numbers of potentially mutagenic families of transposons and other repeated sequences. The chromatin-based transcriptional and epigenetic mechanisms that co-evolved to manage the evolutionary drama as well as gene expression and 3-D nuclear architecture have been elucidated these past 20 years. This perspective traces some of the major developments with which I have become particularly familiar while seeking ways to improve crop plants. I draw some conclusions from this look-back over 50 years during which the scientific community has (i) exposed how chromosomes guard, readout, control, recombine, and transmit information that programs plant species, large and small, weed and crop, and (ii) modified the information in chromosomes for the purposes of genetic, physiological, and developmental analyses and plant improvement.

Metabolic profiling of historical and modern wheat cultivars using proton nuclear magnetic resonance spectroscopy.

To determine changes in the grain components between historical and modern wheat (Triticum aestivum L.) cultivars, wholemeal flours from 19 wheat cultivars and 2 landraces released or introduced between 1870 and 2013 and grown over two crop years were extracted using hydroalcoholic solution and analyzed using one dimensional 1H NMR spectral profiling. Grain yield, grain volume weight (GVW), and grain protein concentration were also measured. Grain yield increased while protein concentration decreased by release year (p < 0.001). Increasing trends (p < 0.01) were observed for tryptophan, sum of the measured amino acids, chlorogenic acid, ferulic acid, vanillic acid, and sum of the measured phenolic acids. Grain yield, phenolic acids, and tryptophan were mainly associated with modern cultivars, whereas grain protein concentration and GVW were associated with historical cultivars. The findings from this study showed changes in concentration of grain components over a century of breeding that may have implications for grain quality and human health.

Wheat (Triticum aestivum L.) Breeding from 1891 to 2010 Contributed to Increasing Yield and Glutenin Contents but Decreasing Protein and Gliadin Contents.

Epidemiologic studies suggest an increasing prevalence of celiac disease and non-celiac gluten/wheat sensitivity. With wheat proteins being the main triggers, changes in wheat protein composition are discussed as a potential cause. The goals of breeding toward increased yield and resistance might have inadvertently contributed to a higher immunostimulatory potential of modern wheat cultivars compared to old wheat cultivars. Therefore, agronomic characteristics, protein content, and gluten composition of 60 German winter wheat cultivars first registered between 1891 and 2010 grown in 3 years were analyzed. While plant height and spike density decreased over time, yield and harvest index increased. The protein and gliadin contents showed a decreasing trend, whereas glutenin contents increased, but there were no changes in albumin/globulin and gluten contents. Overall, the harvest year had a more significant effect on protein composition than the cultivar. At the protein level, we found no evidence to support an increased immunostimulatory potential of modern winter wheat.

Haplotype structure in commercial maize breeding programs in relation to key founder lines.

KEY MESSAGE: High-density haplotype analysis revealed significant haplotype sharing between ex-PVPs registered from 1976 to 1992 and key maize founders, and uncovered similarities and differences in haplotype sharing patterns by company and heterotic group. Proprietary inbreds developed by the private seed industry have been the major source for driving genetic gain in successful North American maize hybrids for decades. Much of the history of industry germplasm can be traced back to key founder lines, some of which were pivotal in the development of prominent heterotic groups. Previous studies have summarized pedigree-based relationships, genetic diversity and population structure among commercial inbreds with expired Plant Variety Protection (ex-PVP). However, less is known about the extent of haplotype sharing between historical founders and ex-PVPs. A better understanding of the relationships between founders and ex-PVPs provides insight into the haplotype and heterotic group structure among industry germplasm. We performed high-density haplotype analysis with 11.3 million SNPs on 212 maize inbreds, which included 157 ex-PVPs registered 1976-1992 and 55 public lines relevant to PVPs. Among these lines were 12 key founders identified in literature review: 207, A632, B14, B37, B73, LH123HT, LH82, Mo17, Oh43, OH7, PHG39 and Wf9. Our results revealed that, on average, 81.6% of an ex-PVP's genome is shared with at least 1 of these 12 founder lines and more than half when limited to B73, Mo17 and 207. Quantifiable similarities and contrasts among heterotic groups and major US seed industry companies were also observed. The results from this study provide high-resolution haplotype data on ex-PVP germplasm, confirm founder relationship trends observed in previous studies, uncover region-specific haplotype structure differences and demonstrate how haplotype sharing analysis can be used as a tool to explore germplasm diversity.

Novel strategies for genomic prediction of untested single-cross maize hybrids using unbalanced historical data.

KEY MESSAGE: Weighted outperformed unweighted genomic prediction using an unbalanced dataset representative of a commercial breeding program. Moreover, the use of the two cycles preceding predictions as training set achieved optimal prediction ability. Predicting the performance of untested single-cross hybrids through genomic prediction (GP) is highly desirable to increase genetic gain. Here, we evaluate the predictive ability (PA) of novel genomic strategies to predict single-cross maize hybrids using an unbalanced historical dataset of a tropical breeding program. Field data comprised 949 single-cross hybrids evaluated from 2006 to 2013, representing eight breeding cycles. Hybrid genotypes were inferred based on their parents' genotypes (inbred lines) using single-nucleotide polymorphism markers obtained via genotyping-by-sequencing. GP analyses were fitted using genomic best linear unbiased prediction via a stage-wise approach, considering two distinct cross-validation schemes. Results highlight the importance of taking into account the uncertainty regarding the adjusted means at each step of a stage-wise analysis, due to the highly unbalanced data structure and the expected heterogeneity of variances across years and locations of a commercial breeding program. Further, an increase in the size of the training set was not always advantageous even in the same breeding program. The use of the two cycles preceding predictions achieved optimal PA of untested single-cross hybrids in a forward prediction scenario, which could be used to replace the first step of field screening. Finally, in addition to the practical and theoretical results applied to maize hybrid breeding programs, the stage-wise analysis performed in this study may be applied to any crop historical unbalanced data.

Construction, characteristics and high throughput molecular screening methodologies in some special breeding populations: a horticultural perspective.

Advanced marker technologies are widely used for evaluation of genetic diversity in cultivated crops, wild ancestors, landraces or any special plant genotypes. Developing agricultural cultivars requires the following steps: (i) determining desired characteristics to be improved, (ii) screening genetic resources to help find a superior cultivar, (iii) intercrossing selected individuals, (iv) generating genetically hybrid populations and screening them for agro-morphological or molecular traits, (v) evaluating the superior cultivar candidates, (vi) testing field performance at different locations, and (vii) certifying. In the cultivar development process valuable genes can be identified by creating special biparental or multiparental populations and analysing their association using suitable markers in given populations. These special populations and advanced marker technologies give us a deeper knowledge about the inherited agronomic characteristics. Unaffected by the changing environmental conditions, these provide a higher understanding of genome dynamics in plants. The last decade witnessed new applications for advanced molecular techniques in the area of breeding,with low costs per sample. These, especially, include next-generation sequencing technologies like reduced representation genome sequencing (genotyping by sequencing, restriction site-associated DNA). These enabled researchers to develop new markers, such as simple sequence repeat and single- nucleotide polymorphism, for expanding the qualitative and quantitative information onpopulation dynamics. Thus, the knowledge acquired from novel technologies is a valuable asset for the breeding process and to better understand the population dynamics, their properties, and analysis methods.

Ancient Indian history: What do we know and how?

When and where was the Rigveda (Rv) composed? How are the Vedic people related to the vast Harappan archaeological tradition? These quintessential questions have no direct answers. At our current level of understanding, archaeology and sacred texts constitute two distinct streams which do not intersect. We must therefore collate evidence from different sources and try to produce a synthesis. It is particularly important to take note of archaeological evidence from Central Asia, because it has not received the attention it deserves. What is well known in science must be kept in mind in the case of history also. A theory to be valid must explain each and every fact (known at present or to be known in future) in a selfconsistent manner. Conversely, even if there is one piece of evidence that a theory is unable to explain, it should be put on hold, modified or even rejected.

Mutation Breeding in Barley: Historical Overview.

The discovery of radioactivity at the end of the nineteenth century played a key role in a series of historical landmarks that would lead to contemporary mutation breeding in agricultural crops. The aim of the earliest experiments was to test the effects of radiation on living organisms beginning with fruit flies. Exposure of plants to X-rays provided the first incontrovertible proof that phenotypic changes could be induced. Chemicals were a second type of mutagen tested from the 1940s and both forms are used today. This chapter is an overview of some of the historical developments that led to the use of mutagenesis in plants, with a focus on barley, a model species for mutation genetics and breeding as well as a major cereal crop. Perhaps the most well-known examples of mutant barley cultivars are Diamant, Golden Promise, and their hybrids.

Retrospective and perspective of rice breeding in China.

Breeding is the art and science of selecting and changing crop traits for the benefit of human beings. For several decades, tremendous efforts have been made by Chinese scientists in rice breeding in improving grain yield, nutrition quality, and environmental performance, achieving substantial progress for global food security. Several generations of crop breeding technologies have been developed, for example, selection of better performance in the field among variants (conventional breeding), application of molecular markers for precise selection (molecular marker assisted breeding), and development of molecular design (molecular breeding by rational design). In this review, we briefly summarize the advances in conventional breeding, functional genomics for genes and networks in rice that regulate important agronomic traits, and molecular breeding in China with focuses on high yield, good quality, stress tolerance, and high nutrient-use efficiency. These findings have paved a new avenue for rational design of crops to develop ideal varieties with super performance and productivity.

[The history and prospect of rice genetic breeding in China].

Rice breeding in China has experienced three major leaps of dwarf breeding, heterosis utilization and green super rice cultivation, accompanied by six important processes: dwarf breeding (the first green revolution), three-line hybrid rice cultivation, two-line hybrid rice cultivation, inter-subspecies heterosis utilization, ideal plant type breeding and green super rice cultivation. The breeding subject ranges from the unique trait of high yield to the complex traits of resistance, high quality and high yield. The breeding concept is gradually upgraded from high yield and quality to the second green revolution concept of "less investment, more output, and better environment". Rice functional genomics achievements have prepared many genes with important utilization values for the second green revolution, and rice breeding is moving towards a new era of design breeding. The genomic selection technology and transgenic technology will help to develop the green super rice for "less pesticides, less fertilizers, water saving and drought tolerance, superior quality and high yield". Here, we summarize the development process of rice genetics and breeding in China, point out advantages and disadvantages of various breeding methods and breeding techniques, systematically introduce the molecular mechanisms on cytoplasmic male sterility, photoperiod-sensitive male genic sterility and indica-japonica hybrid sterility, review the important functional genes related to rice plant architecture, panicle architecture, grain size and nutrient use efficiency, clarify the correlation between yield and heading date, and highlight the important position of China in the rice basic research in the world. In particular, we emphasize the fact that Chinese rice production styles have undergone or are undergoing tremendous changes in recent years, and the breeding concept must also keep pace with the changing production styles. In the future, the hybrid breeding technology should be closely integrated with modern breeding technologies to breed rice varieties that must not only meet the market demand, but also have the natural and healthy characteristics and adapt to the new farming system and methods.

How Mendel's Interest in Inheritance Grew out of Plant Improvement.

Despite the fact that Gregor Mendel is generally respected as the founder of genetics, little is known about the origin of and motivation for his revolutionary work. No primary sources are known that discuss his work during the period of his pea crossing experiments. Here, we report on two previously unknown interconnected local newspaper articles about Mendel's work that predate his famous Pisum lectures by 4 years. These articles describe Mendel as a plant breeder and a horticulturist. We argue that Mendel's initial interests concerned crop improvement, but that with time he became more interested in fundamental questions about inheritance, fertilization, and natural hybridization.

Agrobiotechnology Goes Wild: Ancient Local Varieties as Sources of Bioactives.

The identification and use of species that have best adapted to their growth territory is of paramount importance to preserve biodiversity while promoting sustainable agricultural practices. Parameters including resistance to natural conditions (biotic and abiotic risk factors), biomass and fruit productivity, and phytochemical content with nutraceutical potential, could be used as quantitative markers of the adaptability of plants to wild environments characterized by minimal human impact. Ancient varieties, which are plant varieties growing in regional territories and not destined for market distribution, are a source of unique genetic characters derived from many years of adaptation to the original territory. These plants are often more resistant to biotic and abiotic stresses. In addition, these varieties have a high phytochemical (also known as bioactives) content considered health-beneficial. Notably, the content of these compounds is often lower in commercial cultivars. The use of selected territorial varieties according to the cultivation area represents an opportunity in the agricultural sector in terms of biodiversity preservation, environmental sustainability, and valorization of the final products. Our survey highlights the nutraceutical potential of ancient local varieties and stresses the importance of holistic studies (-omics) to investigate their physiology and secondary metabolism.

Direct archaeological evidence for Southwestern Amazonia as an early plant domestication and food production centre.

Southwestern Amazonia is considered an early centre of plant domestication in the New World, but most of the evidence for this hypothesis comes from genetic data since systematic archaeological fieldwork in the area is recent. This paper provides first-hand archaeobotanical evidence of food production from early and middle Holocene (ca. 9,000-5000 cal. BP) deposits at Teotonio, an open-air site located on a 40 m-high bluff on the south bank of the Madeira river. Such evidence includes the presence of local and exotic domesticates such as manioc (Manihot esculenta), squash (Cucurbita sp.) and beans (Phaseolus sp.), alongside edible fruits such as pequiá (Caryocar sp.) and guava (Psidium sp.) that point to the beginnings of landscape domestication. The results contribute to an ever-growing number of studies that posit southwest Amazonia as an important centre for early crop domestication and experimentation, and which highlight the longue-durée of human impacts on tropical forest biodiversity around the world.

Plant behaviour from human imprints and the cultivation of wild cereals in Holocene Sahara.

The human selection of food plants cannot always have been aimed exclusively at isolating the traits typical of domesticated species today. Each phase of global change must have obliged plants and humans to cope with and develop innovative adaptive strategies. Hundreds of thousands of wild cereal seeds from the Holocene 'green Sahara' tell a story of cultural trajectories and environmental instability revealing that a complex suite of weediness traits were preferred by both hunter-gatherers and pastoralists. The archaeobotanical record of the Takarkori rockshelter in southwest Libya covering four millennia of human occupation in the central Sahara gives us a unique insight into long-term plant manipulation and cultivation without domestication. The success of a number of millets was rooted in their invasive-opportunistic behaviour, rewarded during their coexistence with people in Africa. These wild plants were selected for features that were precious in the past but pernicious for agriculture today. Reconnecting past practices with modern farming strategies can help us to seek out the best resources for the future.

Russia's new Lysenkoism.

During the late 1940s and 1950s, a pseudo-scientific concept based on Marxist-Leninist ideology became internationally known as 'Lysenkoism'. Lysenkoism was a neo-Lamarckian idea, claiming that in crop plants, such as wheat, environmental influences are heritable via all cells of the organism. Lysenkoism was applied to agriculture during the Stalin era with disastrous consequences. Despite the triumphs of modern genetics, and the disproval of Lysenkoism, recent years have seen a 're-thinking' of this doctrine in Russia. This disturbing pro-Lysenko movement, which is accompanied by a growing sympathy for Stalin, claims to have its scientific roots in modern epigenetics, specifically the heritability of variation by mechanisms other than changes in DNA sequence. Based on recent research on the model plant Arabidopsis thaliana, its is clear that Lysenkoism has nothing to do with heritable 'epigenetic' modifications. Biologists should defend science against ideological and political interferences.

Genomic prediction unifies animal and plant breeding programs to form platforms for biological discovery.

The rate of annual yield increases for major staple crops must more than double relative to current levels in order to feed a predicted global population of 9 billion by 2050. Controlled hybridization and selective breeding have been used for centuries to adapt plant and animal species for human use. However, achieving higher, sustainable rates of improvement in yields in various species will require renewed genetic interventions and dramatic improvement of agricultural practices. Genomic prediction of breeding values has the potential to improve selection, reduce costs and provide a platform that unifies breeding approaches, biological discovery, and tools and methods. Here we compare and contrast some animal and plant breeding approaches to make a case for bringing the two together through the application of genomic selection. We propose a strategy for the use of genomic selection as a unifying approach to deliver innovative 'step changes' in the rate of genetic gain at scale.