Effects of neighboring woody plants on target trees with emphasis on effects of understorey shrubs on overstorey physiology in forest communities: a mini-review

Plant-plant interaction plays a key role in regulating the composition and structure of communities and ecosystems. Studies of plant-plant interactions in forest ecosystems have mainly concentrated on growth effects of neighboring plants on target trees. Physiological effects of neighboring plants on target trees, in particular understorey effects on physiology of overstorey trees, have received less attention. It is still unclear what is the physiological mechanisms underlying positive growth effects of understorey removal, although understorey removal has been applied to improve the wood production for hundreds of years worldwide. Only 17.5% of published works dealt with understorey-overstorey interactions and only a few of those researched the understorey effects on the physiology of overstorey trees. Case studies indicated that overstorey Abies faxoniana trees grown with different understorey shrubs showed significantly different levels of tissue nitrogen and mobile carbohydrates. Removal experiment showed that nitrogen and mobile carbohydrates concentrations in Cunninghamia lanceolata trees grown in the absence of understorey shrubs differed significantly (pure stand > mixture) with those in trees grown in the presence of understorey shrubs, in particular during the dry season. This review highlighted that the neighboring woody plants affect Cand N-physiology in overstorey trees. These effects may be mainly resulted from underground competition for soil water rather than for other resources as the effects were more pronounced during the dry season. The present review suggests that positive effects of neighboring removal (e.g., understorey removal, thinning) on overstorey trees can be expected more rapidly and strongly in stressful area (e.g., low rainfall, nutrient-poor site) than in areas with optimal growth conditions. Hence, ecophysiology-based management strategies for dealing with neighboring plants in forest ecosystems should take into account: 1) site conditions, 2) timing, duration and frequency of management practices, and 3) species-specific properties and other aspects such as biodiversity conservation and soil erosion