The spatially heterogeneous distribution of soil nutrients is ubiquitous in terrestrial ecosystems and has been shown to promote the performance of plant communities, influence species coexistence, and alter ecosystem nutrient dynamics. Plants interact with diverse soil microbial communities that lead to an interdependent relationship (e.g., symbioses), driving plant community productivity, belowground biodiversity, and soil functioning. However, the potential role of the soil microbial communities in regulating the effect of soil nutrient heterogeneity on plant growth has been little studied. Here, we highlight the ecological importance of soil nutrient heterogeneity and microorganisms and discuss plant nutrient acquisition mechanisms in heterogeneous soil. We also examine the evolutionary advantages of nutrient acquisition via the soil microorganisms in a heterogeneous environment. Lastly, we highlight a three-way interaction among the plants, soil nutrient heterogeneity, and soil microorganisms and propose areas for future research priorities. By clarifying the role of soil microorganisms in shaping the effect of soil nutrient heterogeneity on plant performance, the present study enhances the current understanding of ecosystem nutrient dynamics in the context of patchily distributed soil nutrients.

1 Unlike non-clonal plants, clonal plants can develop a division of labour in which connected ramets specialize to acquire different, locally abundant resources. This occurs as a plastic response to a patchy environment where two resources tend not to occur together and different ramets experience high availabilities of different resources. We hypothesized that if division of labour is an important advantage of clonal growth in such environments in nature, then clones from habitats where resource availabilities are negatively associated should show a greater capacity for division of labour than clones from habitats where resource availabilities are more uniform. 2 To test this, we collected clones of Fragaria chiloensis from sand dune and grassland sites in each of three regions of the central coast of California, grew pairs of connected or severed ramets under low light and high N or under high light and low N, and measured leaf area, chlorophyll content and final dry mass. Given that previous work has indicated that high availabilities of light and N show a stronger tendency not to occur together in the dune than in the grassland sites, we expected that clones from dunes would show greater capacity for division of labour than clones from grasslands. 3 Clones from dunes showed a greater capacity than clones from grasslands to specialize for acquisition of abundant N via high proportional mass of roots. Clones from the two types of habitats showed similar capacity to specialize for acquisition of abundant light via high leaf area and chlorophyll content of leaves. Specialization via leaf area and chlorophyll content took place mainly within the first half of the 60-day experiment. 4 These results provide evidence that division of labour in a clonal plant has been selected for in natural habitats where high levels of different resources tend to be spatially separated. Results also show that division of labour can occur, not just via allocation of mass, but also via physiological traits, and that both morphological and physiological specialization can take place within a few weeks. 5 Clonal plants dominate many habitats and include many highly invasive species. Division of labour is one of the most striking potential advantages of clonal growth, and is a remarkable instance of phenotypic plasticity in plants. This study further suggests that division of labour in clonal plants is an instance of adaptive plasticity and could therefore play a part in their widespread ecological success.

• Background and Aims In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. • Methods In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. • Key Results Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. • Conclusions The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.