Alpine Botany

, Volume 129, Issue 1, pp 43–52 | Cite as

Responses of photochemical efficiency and shoot growth of alpine dwarf-pine Pinus pumila to experimental warming, shading, and defoliation in Japan

  • Yukihiro AmagaiEmail author
  • Gaku Kudo
Original Article


Global warming accelerates shrub expansion in high-latitude and high-elevation ecosystems. Over the last several decades, alpine dwarf-pine Pinus pumila has expanded its range in northern Japan because of enhanced shoot growth under warm climatic conditions. In alpine regions, local environmental conditions and the length of the growing season, vary depending on the topography, elevation, and snowmelt time. This leads to spatially varying shoot performances that are co-affected by climatic change. We applied a warming, shading, and defoliation treatment to assess how temperature and carbon relations in interaction with habitat type (elevation and snowmelt time) affect shoot growth and photochemical efficiency of needles in this species. Photochemical efficiency (Fv/Fm) was maximized during peak growth in the middle of growing season (mid-July–mid-August), and it increased in the shading and warming treatments especially in the early and late season. Shoot growth increased only in the warming treatment, and was not affected by shading and defoliation. These results indicate that shoot growth of alpine dwarf-pine is limited by low temperature, but not by carbon assimilation, i.e., growth is sink- rather than source-limited. Furthermore, the seasonal trend of photochemical efficiency shifted to the late season at higher elevations, and the recovery time of photochemical efficiency took longer in the late-snowmelt habitat, where the growing season was short. Therefore, warmer summers and longer snow-free periods are likely to enhance the growth and areal expansion of alpine dwarf-pine at the expense of the adjacent, species-rich, low-stature alpine plant communities.


Alpine Pinus pumila Photochemical efficiency Shoot growth Sink demand Snowmelt time 



We are grateful to Yuta Aoshima and Yuki Mizunaga for their assistance with the fieldwork. This study was supported by a Grant-in-Aid from the Ministry of Environment of Japan from the Global Environmental Research Fund (D-0904), JSPS Kakenhi No. 24570015, and the Clarke Memorial Foundation of Hokkaido University.


The study was supported by a grant from the Ministry of Environment of Japan from the Global Environmental Research Fund (D-0904), JSPS Kakenhi No. 24570015, and the Clarke Memorial Foundation of Hokkaido University.

Compliance with ethical standards

Ethical Statement

The authors declare that observance Ethical Standards.

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

The Pinus pumila is not an endangered and endemic species.

Informed consent

The investigation at national park was conducted by obtaining due permission.

Supplementary material

35_2019_217_MOESM1_ESM.pptx (2.6 mb)
Supplemental Fig. S1 Research site locations: Hisago fellfield site (H), Goshikigahara site (G), and southern slope of Mt. Kaun site (K) in the Taisetsu Mountains. Plots were selected along the elevation gradients and snowmelt times in G and K. Photochemical efficiency measurements were conducted at the points marked in gray (PPTX 2621 KB)
35_2019_217_MOESM2_ESM.pptx (505 kb)
Supplemental Fig. S2 Photochemical efficiency (Fv/Fm) of one-year-old needles along the elevation gradients (a) and the snowmelt times (b) during the growing season in 2014 at three locations. (PPTX 505 KB)
35_2019_217_MOESM3_ESM.docx (14 kb)
Supplementary material 3 (DOCX 13 KB)


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Copyright information

© Swiss Botanical Society 2019

Authors and Affiliations

  1. 1.Faculty of Environmental Earth ScienceHokkaido UniversitySapporoJapan
  2. 2.Center for Environmental Biology and Ecosystem StudiesNational Institute for Environmental Studies (NIES)TsukubaJapan

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