Large protein families are a prominent feature of plant genomes and their size variation is a key element for adaptation. However, gene and genome duplications pose difficulties for functional characterization and translational research. Here we infer the evolutionary history of the DOMAIN OF UNKNOWN FUNCTION (DUF) 26-containing proteins. The DUF26 emerged in secreted proteins. Domain duplications and rearrangements led to the appearance of CYSTEINE-RICH RECEPTOR-LIKE PROTEIN KINASES (CRKs) and PLASMODESMATA-LOCALIZED PROTEINS (PDLPs). The DUF26 is land plant-specific but structural analyses of PDLP ectodomains revealed strong similarity to fungal lectins and thus may constitute a group of plant carbohydrate-binding proteins. CRKs expanded through tandem duplications and preferential retention of duplicates following whole genome duplications, whereas PDLPs evolved according to the dosage balance hypothesis. We propose that new gene families mainly expand through small-scale duplications, while fractionation and genetic drift after whole genome multiplications drive families towards dosage balance.

Institute of Biotechnology University of Helsinki Viikinkaari 5 FI 00014 Helsinki Finland

Organismal and Evolutionary Biology Research Programme Viikki Plant Science Centre VIPS Faculty of Biological and Environmental Sciences University of Helsinki Viikinkaari 1 FI 00014 Helsinki Finland

Present Address Laboratory of Plant Molecular Biology Institute of Biophysics AS CR v v i and CEITEC Central European Institute of Technology Mendel University in Brno Zemědělská 1 613 00 Brno Czech Republic

School of Biological Sciences Nanyang Technological University 60 Nanyang Drive Singapore 637551 Singapore

Structural Plant Biology Laboratory Department of Botany and Plant Biology University of Geneva Geneva Switzerland

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