Nitrogen and carbohydrate storage in biennials originating from habitats of different resource availability.

Four biennial species (Arctium tomentosum, Cirsium vulgare, Dipsacus sylvester and Daucus carota) which originate from habitats of different nutrient availability were investigated in a 2-year experiment in a twofactorial structured block design varying light (natural daylight versus shading) and fertilizer addition. The experiment was designed to study storage as reserve formation (competing with growth) or as accumulation (see Chapin et al. 1990). We show that (i) the previous definitions of storage excluded an important process, namely the formation of storage tissue. Depending on species, storage tissue and the filling process can be either a process of reserve formation, or a process of accumulation. (ii) In species representing low-resource habitats, the formation of a storage structure competes with other growth processes. Growth of storage tissue and filling with storage products is an accumulation process only in the high-resource plant Arctium tomentosum. We interpret the structural growth of low-resource plants in terms of the evolutionary history of these species, which have closely related woody species in the Mediterranean area. (iii) The use of storage products for early leaf growth determines the biomass development in the second season and the competitive ability of this species during growth with perennial species. (iv) The high-resource plant Arctium has higher biomass development under all conditions, i.e. plants of low-resource habitats are not superior under low-resource conditions. The main difference between high- and low-resource plants is that low-resource plants initiate flowering at a lower total plant internal pool size of available resources.