title: Systemic induced resistance: a risk-spreading strategy in  clonal plant networks?
abstract: Clonal plant networks consist of interconnected individuals (ramets) of different sizes and ages. They represent heterogeneous ramet assemblages with marked differences in quality and attractiveness for herbivores. Here, feeding preferences of a generalist herbivore (Spodoptera exigua) for differently-aged ramets of Trifolium repens were studied, and changes in herbivore preference in response to systemic defense induction were investigated. Dual-choice tests were used to assess the preference of herbivores for young versus mature ramets of induced and uninduced plants, respectively. Additionally, leaf traits related to nutrition, biomechanics and chemical defense were measured to explain variation in tissue quality and herbivore preference. Young ramets were heavily damaged in control plants. After systemic defense induction, damage on young ramets was greatly reduced, while damage on mature ramets increased slightly. Defense induction increased leaf strength and thickness, decreased leaf soluble carbohydrates and substantially changed phenolic composition of undamaged ramets connected to attacked individuals. Systemic induced resistance led to a more dispersed feeding pattern among ramets of different ages. It is proposed that inducible defense acts as a risk-spreading strategy in clonal plants by equalizing herbivore preference within the clone, thereby avoiding extended selective feeding on valuable plant tissues.
authors.affiliation: Experimental Plant Ecology, Radboud University Nijmegen, Nijmegen, Netherlands.|Present address: Biology, Tufts University, Medford, MA, USA.
authors.forename: Sara
authors.initials: S
authors.lastname: Gómez
authors.affiliation: Plant Ecology and Biodiversity, Utrecht University, Utrecht, Netherlands.
authors.forename: Yusuke
authors.initials: Y
authors.lastname: Onoda
authors.affiliation: Department of Chemistry. University of Turku, Turku, Finland.
authors.forename: Vladimir
authors.initials: V
authors.lastname: Ossipov
authors.affiliation: Experimental Plant Ecology, Radboud University Nijmegen, Nijmegen, Netherlands.
authors.forename: Josef F
authors.initials: JF
authors.lastname: Stuefer
mesh_terms: D000818:Animals; D001696:Biomechanical Phenomena; D002241:Carbohydrates / Q000032:analysis; D002244:Carbon / Q000032:analysis; D005518:Food Preferences; D005838:Genotype; D007814:Larva / Q000502:physiology; D009584:Nitrogen / Q000032:analysis; D010636:Phenols / Q000032:analysis; D018515:Plant Leaves / Q000033:anatomy & histology / Q000737:chemistry / Q000502:physiology; D018411:Spodoptera / Q000254:growth & development / Q000502:physiology*; D029921:Trifolium / Q000033:anatomy & histology / Q000737:chemistry / Q000502:physiology*
chemical_list: D002241:Carbohydrates; D010636:Phenols; D002244:Carbon; D009584:Nitrogen