关键词: 横断山区, 高山冰缘植物, 特化形态结构, 生态适应机制, 毡毛雪莲

Abstract

Aims: As one of the most prominent highly specialized morphologies evolved in alpine sub-nival plants in Himalayan-Hengduan Mountains, Southwestern China, ‘greenhouse plant’ is characterized by the huge and semi-translucent bracts that cover inflorescence. However, unlike some typical ‘greenhouse plants’, such as Rheum nobile and R. alexandrae (Polyginonaceae) and Saussurea obvallata (Asteraceae) that have their inflorescence covered by bracts throughout the entire growing season, some other plant species, such as S. velutina studied here has their inflorescence covered by bracts in the early part of the growing season only, while in the majority of the growing season, this plant chooses to extrude its inflorescence outside the cover of bracts till fruit maturation. However, the adaptive strategy behind such an alteration in morphology remained largely unknown. 

Methods: In this study we purposely investigated the thermal situation of the inflorescence in S. velutina after it has extruded from the bract covering. Given these bracts gradually changed their colors from green-yellow to deep pink-purple during this period and they are still attached to the inflorescence, we also assessed the optical properties of the deep pink-purple bracts and explored their potential role on attracting pollinators by assessing the visual model of the bracts to bumblebee (Bombus) as compared with bedrock in its natural habitat and the green-colored leaves. In addition, the influence of the deep pink-purple bracts on seed productions was investigated by gently striping these bracts off. 

Results: Our results show that the temperature of inflorescence remains on average 3-5 K (Calvin temperature index, used to indicate the differences of temperature) higher than adjacent air temperature at the same height of the plant individuals (15—20 cm above ground) during both daytime and nighttime, with the greatest deviation (> 20 K) reaching at sunny-days. Under all prevailing weather conditions (sunny, cloudy, rainy), the dark and solid inflorescence exhibits a pronounced heating efficiency (> 40 %) and the buffering capacity of which reached at ca. 100 minutes as compared with air temperature. However, although the deep pink-purple bracts can still exhibit selective filtering (24.4—36 %) of red and infrared wavelength range that are attributed to warming effects, the artificial treatment on inflorescence did not significantly reduce the seed quantity and quality compared with controlled inflorescence with bracts kept their natural morphology. The color distance of Bombus terrestris from the reference of bed rock to bracts (0.09±0.01 CH) was far less than to green leaves (0.22±0.01 CH), likely indicating a small influence of the bracts on pollinator attractions. However, when green leaves were used as reference, the color distance from them to bracts (0.30±0.02 CH) was significantly different from them to B. terrestris, suggesting a function of pollinator attraction of bracts. These seemingly contradictory results call for more in situ observations and experiments, such as examining the impact of bracts on pollination visit rates to unveil the ecological function of these deep pink-purple bracts. 

Conclusion: The warming and buffering effects of the inflorescence of S. velutina was explained by its dark and solid structure. Such a highly specialized morphology is an adaptive strategy of sub-nival plant against the cool climate during the majority of summer monsoon and short period of growing season in their habitat at exceptionally high elevations.

Key words: Himalaya-Hengduan Mountains, sub-nival plants, highly specialized morphology, adaptive strategy, Saussurea veltutina