Plant functional traits have been widely used to study the linkage between environmental drivers, trade-offs among different functions within a plant, and ecosystem structure and functioning. Here, the whole-plant traits, leaf morphological and physiological traits of two dominant species, Leymus chinensis (C 3 perennial rhizome grass) and Cleistogenes squarrosa (C 4 perennial bunchgrass), were studied in the Inner Mongolia grassland of China, with a grazing experiment including five stocking rates (0, 3.0, 4.5, 7.5, and 9.0 sheep/ha) in 2008 (wet year) and 2009 (dry year). Our results demonstrated that, for both species, the effects of stocking rate, year, and stocking rate×year on whole-plant traits and leaf morphological and physiological traits were highly significant in most cases. The differential responses of plant trait to variation in precipitation were caused by trait trade-offs between the wet and dry years. L. chinensis adopted the high N content and net photosynthetic rate (P n ) in the wet year but both the low N content and P n in the dry year under grazed conditions. The trait trade-offs of C. squarrosa were characterized by high specific leaf area (SLA) and P n in the dry year vs. low SLA and P n in the wet year. Our findings also indicate that C. squarrosa is more resistant to grazing than L. chinensis in terms of avoidance and tolerance traits, particularly under heavy grazing pressure and in the dry year.

Summary1. Large grazing herbivores have been reported to determine the structure and function of grassland ecosystems. However, the ecological linkages between structure and functioning components have yet been thoroughly explored. 2. Here, we test the hypothesis of the impact of grazing on soil nematode community (e.g. structure and composition) and linkages to ecosystem functioning (e.g. soil N mineralization and ANPP) via changes in pathways of plant community, soil nutrients and soil environment using a field experiment maintained for 5 years with seven levels of grazing intensity in the Inner Mongolian grassland. 3. A structural equation model (SEM) with nematode abundances as response variables showed that plant-feeding and fungal-feeding nematodes were driven by changes in the plant community, and bacterial-feeding nematodes were affected by soil abiotic nutrients and environment, while omnivorous + carnivorous nematodes were altered by soil environment and bacterial-feeding nematodes. This indicates that the top-down control by grazing leads to bottom-up control in the soil food web. 4. We found that grazing affected the ecosystem functioning via different pathways. Grazing effects soil N mineralization by changing plant community, soil nutrients, soil environment and nematodes community structure, while it affects ANPP by altering soil N mineralization and soil environment. 5. Our findings could provide a better understanding of the responses of plants and soils to grazing and the linkages between structure and functioning of above-ground and belowground in the semi-arid steppe.

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