Understanding the mechanism of plant morphological plasticity in response to grazing and clipping of semiarid grassland can provide insight into the process of disturbance-induced decline in grassland productivity. In recent studies there has been controversy regarding two hypotheses: 1) grazing avoidance; and 2) growth limiting mechanisms of morphological plasticity in response to defoliation. However, the experimental evidence presented for the memory response to grazing and clipping of plants has been poorly reported. This paper reports on two experiments that tested these hypotheses in field and in a controlled environment, respectively. We examined the effects of long-term clipping and grazing on the functional traits and their plasticity for Leymus chinensis (Trin.) Tzvelev (the dominate species) in the typical-steppe grassland of Inner Mongolia, China. There were four main findings from these experiments. (i) The majority of phenotypic traits of L. chinensis tended to significantly miniaturize in response to long-term field clipping and grazing. (ii) The significant response of morphological plasticity with and without grazing was maintained in a hydroponic experiment designed to remove environmental variability, but there was no significant difference in L. chinensis individual size traits for the clipping comparison. (iii) Plasticity indexes of L. chinensis traits in a controlled environment were significantly lower than under field conditions indicating that plants had partial and slight memory effect to long-term grazing. (iv) The allometry of various phenotypic traits, indicated significant trade-offs between leaf and stem allocation with variations in plant size induced by defoliation, which were maintained only under grazing in the hydroponic controlled environment experiment. Taken together, our findings suggest that the morphological plasticity of L. chinensis induced by artificial clipping was different with that by livestock grazing. The miniaturization of plant size in long-term grazed grassland may reflect retained characteristics of dwarf memory for adaptation to long-term grazing by large herbivores.
Identifying the linkages between nutrient properties and plant size is important for reducing uncertainty in understanding the mechanisms of plant phenotypic plasticity. Although the positive effects of grazing exclusion on plant morphological plasticity has been well documented, surprisingly little is known about the relationship of nutrient strategies with plant shoot size after long-term grazing exclusion. We experimentally investigated the impacts of grazing exclusion over time (0, 9, 15, and 35 years) on the relationships of nutrient traits (nutrient concentration, allocation, and stoichiometry) of with morphological plasticity in Leymus chinensis, which is a dominant species in grasslands of Inner Mongolia, China. Our results showed that there was a significantly negative correlation between the degrees of plasticity and stability of various morphological traits. Increases in plant size by 126.41, 164.17, and 247.47% were observed with the increase of grazing exclusion time of 9, 15, and 35 years, respectively. Plant size was negatively correlated with nitrogen (N) and phosphorus (P) concentrations, but was positively correlated with carbon (C) concentration. Biomass partitioning and leaf to stem ratios of nutrient concentrations contributed more than 95% of the changes in N, P, and C allocation in L. chinensis leaves and stems induced by grazing exclusions. Nine years’ grazing exclusion rapidly changed the nutrient concentrations (averaged by -34.84%), leaf to stem nutrient allocations (averaged by -86.75%), and ecological stoichiometry (averaged by +46.54%) compared to free-grazing, whereas there was no significant trend of these nutrient traits across the 9, 15, and 35 years’ grazing exclusion in L. chinensis individuals. Our findings suggest that with the increase of the duration of the grazing exclusion, time effects on plant performances gradually weakened both in plant morphological plasticity and nutrient properties. There is a significant negative effect between plant sizes and nutrient traits under long-term grazing exclusion.
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