14Author contributions: SK conceived the project with JCB, conducted physiological measurements, 15analyzed and interpreted the data with JCB and RM, and wrote the first draft of the paper. JCB and RM 16designed the FAB experiment, coordinated annual biomass surveys, and contributed to subsequent 17 revisions of the paper.
19Summary 20 Interspecific facilitation is often invoked in explanations of biodiversity-ecosystem function 21 relationships in plant communities, but it is seldom clear how it occurs. Physiological 22 experiments show that excess light causes stress that may depress long-term carbon assimilation. 23If shading by a plant's neighbors reduces light stress, it may facilitate that plant's growth. On the 24 other hand, when light is a limiting factor for growth, shading will often have a net negative, 25 competitive effect. 26 In a tree diversity experiment, we measured growth rates and photosynthetic physiology of 27 broadleaf tree species across a gradient of light availability imposed by their neighbors. At the 28 extremes, trees experienced nearly full sun (monoculture), or were shaded by nearby fast-growing 29 conifers (shaded biculture). 30 Although most species had lower growth with larger neighbors, implying a net competitive effect, 31 the two most shade-tolerant species (Tilia americana and Acer negundo) had positive responses 32 to neighbor size. Compared to the others, these two species were especially susceptible to 33 photoinhibition (reduced dark-acclimated Fv/Fm) in full sun. While most species had lower 34 assimilation rates in shaded bicultures, T. americana had carbon assimilation rates up to 25% 35 higher. T. americana also dropped its leaves 3-4 weeks later in the shaded biculture, extending its 36 growing season. We conclude that although large neighbors can cause light limitation in shade-37 intolerant species, they can also increase growth through abiotic stress amelioration in shade-38 tolerant species. 39 Both positive and negative species interactions in our experiment can be explained by the 40 photosynthetic responses of trees to the light environment created by their neighbors. We show 41 that physiological measurements can deepen our understanding of the species interactions that 42 underlie biodiversity-ecosystem function relationships. 43 44