( Arkadiusz Kosmala Kopecký David Eds.) Festulolium from the Nature to Modern Breeding Merging two genomes from distant species provides an unprec- edented opportunity to combine agricultural traits from both parents Festu into a single organism in breeding programs, but also to uncover basic mechanisms of plant evolution. Festulolium, a forage grass hybrid originated from the cross of fescue (Festuca sp.) and rye- grass (Lolium sp.) species, combines high yield from ryegrasses Festulolium and abiotic stress tolerance from fescues. Moreover, ability to distinguish parental chromosomes and existence of homoeologous lolium chromosome pairing and recombination allows tracking evolutionary changes associated with wide hybridization. This book collects cur- from the Nature rent knowledge in breeding and research of Festulolium hybrids. It may serve as a platform providing breeders and researchers with new scienti c information on agricultural performance, genetics, to Modern physiology, ecology and evolution of this outstanding grass hybrid. Breeding Modern Breeding to the Nature from David Kopecký and Arkadiusz Kosmala Editors Selected Papers from Biologia Plantarum Sponsors Institute of Experimental Botany AS CR, Olomouc, Czech Republic olomouc.ueb.cas.cz/en Institute of Plant Genetics, Polish Academy of Sciences, Poznan, Poland www.igr.poznan.pl/en/home-en DLF Seeds, s.r.o., Hladké Životice, Czech Republic www.dlf.com Centre of the Region Haná for Biotechnological and Agricultural Research cr-hana.eu/en EUCARPIA Fodder Crops and Amenity Grasses Section www.forages-eucarpia.org Festulolium from the Nature to Modern Breeding David Kopecký and Arkadiusz Kosmala Editors Selected Papers from Biologia Plantarum All the articles are published under the terms of the Creative Commons BY-NC-ND Licence (creativecommons.org/licenses/by/4.0/). 1st edition © Y. Akiyama, J. Aper, J. Baert, J. Balek, S. Barth, M. Bělínová, J. Bojarszczuk, B. Boller, B. Borawska-Jarmułowicz, W. Borucki, S. L. Byrne, V. Černoch, N. Curran, J. Doležel, J. Eitzinger, J. Fuchs, A. Ghesquiere, M. Glombik, B. Golińska, P. Goliński, J. Grant, D. Grogan, M. Hanley, E. Harasim, J. Harper, P. Hlavinka, T. R. Hodkinson, M. W. Humphreys, K. Hura, K. Jaškūnė, V. Kemešytė, D. Kopecký, A. Kosmala, P. Kraska, A. Kubota, C. Kwiatkowski, V. Mahelka, J. Majka, M. Majka, E. Małuszyńska, K. Marzec-Schmidt, G. Mastalerczuk, D. Milbourne, E. O’riordan, J. E. Olesen, D. Perlikowski, A. Płażek, Y. Sanada, M. A. Semenov, D. Semerádová, A. Schaumberger, M. Staniak, G. Statkevičiūtė, K. Tamura, K. Tase, M. Trnka, K. Van Laere, C. Van Waes, E. Willner, P. Zahradníček, C. Zhao, Z. Zwierzykowski, Z. Žalud, 2021 © Palacký University Olomouc, 2021 ISBN 978-80-244-5912-7 (print) ISBN 978-80-244-5913-4 (online: iPDF) Table of content Preface ...............................................................vii Humphreys and Zwierzykowski: Festulolium, a century of research and breeding and its increased relevance in meeting the requirements for multifunctional grassland agriculture. 1 Boller et al.: Spontaneous natural formation of interspecific hybrids within the Festuca-Lolium complex ............................................... 27 Majka et al.: Cytogenetic insights into Festulolium. 52 Baert et al.: Breeding and genetics of two new amphiploid Festulolium synthetics with improved yield and digestibility .......................................... 63 Kemešytė et al.: Festulolium field performance under fluctuating growing conditions in Lithuania . 77 Curran et al.: Persistency, yield, and silage quality of Festulolium cultivars over a consecutive five-year period under a mild Atlantic climate . 89 Borawska-Jarmułowicz et al.: Morphological diversity of seeds of Polish Festulolium cultivars depending on weather conditions. 109 Tamura et al.: Quantitative trait loci for winter hardiness and other agronomic traits in the progeny of two amphiploid Festulolium hybrids (Festuca pratensis × Lolium perenne) backcrossed twice to tetraploid Lolium perenne ............122 Perlikowski and Kosmala: Mechanisms of drought resistance in introgression forms of Lolium multiflorum/Festuca arundinacea .......................................146 Staniak et al.: Prolonged drought stress induced changes in yield and physiological processes of Trifolium repens and Festulolium braunii .....................158 Černoch and Kopecký: Drought tolerance and regrowth capacity revealed in the Festuca-Lolium complex ............................................................172 vi Marzec-Schmidt et al.: Rapid increases in β-1,3-glucanase and chitinase activities are markers of resistance to Microdochium nivale in grasses of the Lolium-Festuca complex. 186 Golińska and Goliński: Tensile strength of the rachilla in spikelets of Polish Festulolium braunii cultivars ............................................................199 Trnka et al.: Future agroclimatic conditions and implications for European grasslands. 209 Zwierzykowski and Kopecký: A brief history and (nearly) complete bibliography of Festulolium ............241 Kopecký: List of interspecific hybrids observed within theFestuca -Lolium complex .....280 Appendix ...........................................................283 vii Preface Dear Festulolium community, one hundred years have passed from the first reported artificial hybridization of fescues with ryegrasses and development of the first man-made hybrid, later on named Festulolium. The attempts at interspecific hybridization were not original. In those days, it was well known that species of these two genera can cross with each other and produce hybrids. However, these hybrids were sterile and ap- peared to represented an evolutionary dead-end. Only after the implementation of biotechnology techniques in the 1960’s, such as induced polyploidization and embryo rescue, breeders succeeded in the development of fertile tetraploid hy- brids. Since then, synthetic Festulolium hybrids have been developed in numer- ous breeding programs and some were released as cultivars. The original idea to combine agriculturally beneficial traits, such as yield and nutrition from ryegrass with stress tolerance from fescues, has turned into reality, as for some traits and in some locations Festulolium cultivars outperform their parental species. As such, they became a valuable part of grass mixtures and found their place in seed marketing. Over the years, Festulolium cultivars have become popular among farmers and growers, and their acreage has gradually increased. Predictions of future climate open new utilization of Festulolium hybrids. A higher probability of adverse events in the near future, such as flooding and periods of drought and heat during summer, calls for the development of new genotypes to withstand such conditions and mitigate the effect of these abiotic stresses. Enormous vari- ability of Festulolium and the potential to combine traits from various species is envisaged as the main drive of Festulolium breeding for future conditions. Apart from their breeding value, Festulolium hybrids came into focus of plant evolutionary biologists and geneticists due to their unique features. Chromo- somes of the parental species pair and recombine in the hybrids, generating highly heterogeneous progeny. At the same time, parental chromosomes can be “painted” by in situ hybridization with labeled DNA probes and thus, allow tracking of individual chromosomes in the progeny. These two features opened the way to broaden our knowledge on the principles of meiotic chromosome pairing, evolution of hybrid genomes and competition and collaboration of two distinct genomes in a single organism. Moreover, several studies increased our understanding of the transmission and regulation of specific traits from one species into the other. In 2019, we decided to compile a special issue of Biologia Plantarum entitled “Festulolium – from the nature to modern breeding”. This special issue contains viii 14 articles, including eleven original research papers and three reviews, which are focused on genetics, ecology, physiology, biotechnology and the breeding value of Festulolium and various species within the Festuca-Lolium complex. As a follow-up, we resolved to publish this special issue as a book, to commemorate the anniversary of first targeted Festulolium hybridization. Apart from the papers published in the special issue of Biologia Plantarum, we also include two ad- ditional articles and an Appendix to show the beauty of our amazing crop. One of these articles, by Z. Zwierzykowski and D. Kopecký provides a brief history of the Festulolium research and breeding and as nearly a complete bibliography of Festulolium as possible, while the other, by D. Kopecký, lists all combinations of parental species generated in the development of Festulolium hybrids. We would like to thank all the authors for submitting such a broad range of interesting papers. We are grateful to all reviewers for providing critical feedback, which greatly helped to improve the quality of the manuscripts. Special thanks go to the authors of wonderful photos and figures, which are included in the Appendix. Finally, we would like to express our gratitude to our sponsors and home institutions for a great support, which made publishing this book available. We wish you good reading and enthusiasm and energy in your future research and breeding of our beloved grass. David Kopecký and Arkadiusz Kosmala Editors 1 Original citation: BIOLOGIA PLANTARUM 64: 578–590, 2020 DOI: 10.32615/bp.2020.108