1. Department of Genetics and Genome Biology, University of Leicester, Leicester LE1 7RH, United Kingdom
2. Independent Research Group Meiosis, Leibniz Institute of Plant Genetics and Crop Plant Research, Gatersleben, D–06466 Stadt Seeland, Germany
3. Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom
4. University of Bristol, Bristol BS8 1TQ, United Kingdom