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Comparison of the growth traits of a commercial pioneer tree species, paper mulberry (Broussonetia papyrifera L. Vent.), with those of shade-tolerant tree species: investigation of the ecophysiological mechanisms underlying shade-intolerance

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In rural areas of northern Laos, a commercially valuable pioneer tree species, paper mulberry (Broussonetia papyrifera L. Vent.) has been recommended for intercropping under plantations of commercial trees. However, less is understood about the growth of this pioneer tree species in the understorey and the mechanism underlying the shade intolerance. We measured growth characteristics for seedlings of paper mulberry under four light intensities. We compared the relative growth rates in aboveground biomass and standing leaf area (RGRmass and RGRleaf), light-capture efficiency, and seeding-level mass-based daily photosynthetic rates (A mass) with those of field-grown seedlings of eight shade-tolerant species to identify factors potentially responsible for shade-intolerance. Most growth traits of the paper mulberry seedlings did not differ consistently from those of the shade tolerant species. The ecophysiological–architectural model software showed higher A mass and RGRmass capacity in paper mulberry than in shade-tolerant species. Despite their higher RGRmass, paper mulberry seedlings had negative RGRleaf under shaded conditions due to short leaf lifespan. The linear RGRmass–RGRleaf relationship for paper mulberry had a high RGRmass intercept, indicating that a high RGRmass was required to provide positive RGRleaf. Progressive decreases in standing leaf area with time, and possibly photosynthesis, appear to be responsible for the shade-intolerance of paper mulberry. Although intercropping of paper mulberry has been suggested in the species’ native region, understorey cultivation of paper mulberry would only be possible with relatively open canopies.

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We thank the anonymous reviewers who gave valuable comments on the earlier version of this manuscript. We are grateful for the advice and kind support provided during our field survey of natural paper mulberry forests and paper mulberry plantations in northern Laos by Dr. Andreas Neef of Kyushu University, Dr. Ulrich Schuler of the German Federal Institute for Geosciences and Natural Resources (BGR), Dr. Wolfram Spreer of the University of Hohenheim, Mr. Peter Elstner of Chiang Mai University, and Mr. Bernhard Mohns of Deutsche Gesellschaft für Technische Zusammenarbeit. This study was carried out as part of the master’s thesis of CM, who was the recipient of an MSc grant financed by the Japanese Grant Aid for Human Resource Development Scholarship (JDS). This study was also supported by the JSPS to Y Miyazawa (20–7278, 25850108) and by JSPS Core-to-Core Program, B. Asia-Africa Science Platforms.

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Correspondence to Yoshiyuki Miyazawa.



See Figs. 4, 5 and 6.

Fig. 4
figure 4

Three-dimensional images of paper mulberry seedlings in the four light-intensity treatments produced using the Y-plant software

Fig. 5
figure 5

Temporal changes in a aboveground biomass and b standing leaf area. Values represent means (n = 5 per treatment) and standard deviations

Fig. 6
figure 6

Biomass allocation of paper mulberry at the time of harvest to leaves (white), stems (gray), and roots (black)

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Miyazawa, Y., Manythong, C., Fukuda, S. et al. Comparison of the growth traits of a commercial pioneer tree species, paper mulberry (Broussonetia papyrifera L. Vent.), with those of shade-tolerant tree species: investigation of the ecophysiological mechanisms underlying shade-intolerance. Agroforest Syst 88, 907–919 (2014).

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