An isotopic (N-15) assessment of intraclonal regulation in C-4 perennial grasses: ramet interdependence, independence or both?
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1 A series of three experiments was conducted with three C4 perennial grasses, Panicum virgatum (short rhizomes forming distinct clones), Schizachyrium scoparium (caespitose) and Bouteloua gracilis (caespitose), to determine whether intraclonal regulation was characterized by interdependence, independence or a combination of both processes. Specific hypotheses tested were: (i) all ramets within individual clones are physiologically integrated; (ii) ramet hierarchies (several connected generations) within individual clones are capable of acquiring soil nitrogen equally from localized nutrient-rich pulses; and (iii) nitrogen is retained within individual hierarchies through allocation to sequentially developed ramet generations. 2 Mean 15N atom percent excess values within labelled ramet hierarchies were 10-21-fold greater than in immediately adjacent ramet hierarchies, and 105-137-fold greater than in the remaining ramet hierarchies within clones for all three species. This pattern of 15N distribution demonstrates that physiological integration does not occur among all ramets within individual clones and that ramet hierarchies function as integrated physiological individuals. 3 Ramets in closer proximity to a 15N pulse in soils acquired disproportionately greater amounts of the isotope compared to more distant ramets within clones for all three species. Inequitable resource acquisition among ramet hierarchies establishes the potential for asymmetric intraclonal competition below-ground and the development of size inequalities among hierarchies. 4 Approximately 21.7%, 3.5%, 1.7% and 2.3% of the initial 15N pool was allocated from the first ramet generation to the second and third (spring cohort), third (autumn cohort) and fourth generations, respectively. However, the magnitude of nitrogen allocation from parent to juvenile ramet generations was relatively consistent, with 17-32% of the 15N pool within parental ramets allocated to juvenile ramets. Intergenerational 15N allocation in S. scoparium clones therefore demonstrates a high degree of interdependence within individual ramet hierarchies. 5 Clones of the rhizomatous and two caespitose grasses investigated functioned as assemblages of autonomous ramet hierarchies, but ramet generations within individual hierarchies were effectively integrated. Consequently, intraclonal regulation in these temperate, perennial grasses is characterized by both ramet interdependence and independence. The occurrence of ramet independence throughout much of the life history of perennial grasses demonstrates the ecological importance of clonal fragments to genet and population maintenance and emphasizes that this may be the most relevant scale to investigate further the ecology of this important and widely distributed group of clonal plants.
