Abstract
We examined the effects of artificial warming on physiological, biochemical, and structural changes in leaves of maize plants (Zea mays L.) with a field warming experiment in the North China Plain. Stomatal characters, leaf anatomy and ultrastructure, gas exchange, and carbohydrate and mineral nutrition concentrations were examined using light microscopy, electron microscopy, portable photosynthesis system (Licor-6400), and inductively coupled plasma atomic emission spectroscopy. We found that artificial warming (about 2 °C) increased both the stomatal index and stomatal size, and thus increased net photosynthesis rate (A), stomatal conductance (g s), and transpiration rate (E). Artificial warming also significantly increased the profile area of chloroplast and mitochondria, but decreased leaf width and thickness, mesophyll thickness, and mesophyll cell size (mainly palisade cell size). In addition, artificial warming also significantly increased the foliar C:N ratio and soluble sugar contents (glucose, fructose, and sucrose), but not the mineral nutrients and starch contents. Our findings suggest that future global warming may affect the maize growth and production in northern China due to the direct warming effects on the structures (anatomy and ultrastructure), biochemical properties and gas exchanges of the maize leaves.



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Acknowledgments
We thank the associate editor Dr. Urs Feller and the two anonymous reviewers for their constructive comments and suggestions on revising the manuscript. This research was partially supported by the National Key Research and Development Program of China (2012CB417103 and 2010CB833500). This research was also partially supported by Forestry Department of Sichuan Province (Metering System Development and Application of the Regional Forestry Carbon Sink and Source, 2009-204). The authors gratefully acknowledge Dr. Joshua Caplan and Dr. Patrick Burgess (Rutgers University, USA) for their comments on the draft of the manuscript.
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Zheng, Y., Xu, M., Shen, R. et al. Effects of artificial warming on the structural, physiological, and biochemical changes of maize (Zea mays L.) leaves in northern China. Acta Physiol Plant 35, 2891–2904 (2013). https://doi.org/10.1007/s11738-013-1320-z
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DOI: https://doi.org/10.1007/s11738-013-1320-z