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The radial growth of trees in alpine timberline is particularly sensitive to climate change. We sampled and disposed tree-ring cores of three coniferous tree species including Juniperus saltuaria , Abies forrestii , and Larix potaninii at alpine timberline in Yading Nature Reserve. The standard tree-ring chronology was used to explore the response of radial growth of different timberline species to climate change. The results showed that radial growth of L. potaninii increased after 2000, while that of A. forrestii declined after 2002, and J. saltuaria showed a significant decreasing growth trend in the past 10 years. Such results indicated divergent growth responses to climate factors among the three tree species at alpine timberline. The radial growth of J. saltuaria was sensitive to temperature, and was positively correlated with the minimum temperature from previous October to current August, the mean tempera-ture from previous November to current April and from current July to October, but was negatively associated with the relative humidity from current July to October. The radial growth of A. forrestii showed negative correlation with mean temperature and the maximum temperature from May to June in the current year, while it exhibited positive association with the relative humidity and the Palmer drought severity index from May to June in the current year. L. potaninii radial growth was positively associated with mean temperature and the maximum temperature of November-December in the previous year, the maximum temperature of current March and mean temperature of current August. The temporal stability of climate-growth relationship varied among different timberline species. The positive correlation between radial growth of A. forrestii and J. saltuaria and temperature gradually decreased, while the posi-tive relationship of L. potaninii radial growth and temperature gradually increased. Under the background of climate warming, rapid rise in surface air temperatures may promote the radial growth of L. potaninii , while inhibit that of J. saltuaria and A. forrestii , which may change the position of regional timberline.

We explored the effects of drought stress on photosynthetic characteristics and non-structural carbohydrate (NSC) accumulation of the timberline tree species Betula ermanii in Changbai Mountain with a drought control experiment. The results showed that drought significantly reduced the net photosynthetic rate and stomatal conductance, but increased water use efficiency (WUE) of B. ermanii seedlings. Drought dramatically improved the contents of soluble sugar and total NSC in leaves, barks, stems, and roots of B. ermanii seedlings, but significantly reduced their starch content. The stomatal conductance, photosynthetic rate and WUE decreased rapidly as the drought continued, whereas the contents of soluble sugar, starch and NSC increased and then declined. At the end of the experiment, 90% of the leaves turned yellow, and the ratios of soluble sugar to starch in the stems, barks and roots under the drought treatment were significantly higher than those in the control. These results demonstrated that B. ermanii might be a drought-avoidance species that could reduce water loss by rapidly reducing stomatal conductance and improving WUE under drought stress. B. ermanii might have evolved priority storage strategy to cope with water deficit through improving the content of soluble sugar in organs and increasing the transformation rate between starch and sugar. With the extension of drought stress, seedlings tended to die, since water stress might exceed the threshold of the plant self-regulation capacity. However, the content of NSC in organs did not decrease, suggesting that the death of B. ermanii under drought stress might not be caused by carbon starvation.

Climate change is a major cause of changes in alpine and polar vegetation, particularly at the edges of distributions. In temperate regions, these changes are expected to occur at the timberline of alpine zones. On Mt. Fuji, the highest mountain in Japan, the timberline is located 2400-2500 m above sea level. Over a 40-year period (1978-2018), we researched changes in the timberline vegetation of Mt. Fuji. A permanent belt transect extending from the upper timberline to subalpine zones was set up in August 1978. Tree diameters and heights were recorded at the establishment of the transect and every 20 years afterwards. Over the 40 years of the study, the timberline advanced rapidly upwards, and the degree of vegetation cover above the timberline increased remarkably. Notably, the expansion of Salix reinii into the upper part of the timberline facilitated the subsequent spread of Larix kaempferi into this zone. Seedlings of L. kaempferi were particularly abundant at the upper timberline and became established on the uppermost part of the slope. The shape of L. kaempferi at the upper timberline changed from a prostrate form to an upright tree form. We conclude that the upward advance of the alpine timberline observed on Mt. Fuji is due to climate change.

In alpine regions, elevational gradients in environmental parameters are reflected by structural and functional changes in plant traits. Elevational changes in plant water relations have also been demonstrated, but comparable information on root hydraulics is generally lacking. We analyzed the hydraulic efficiency (specific hydraulic conductivity k s, entire root system conductance K R) and vulnerability to drought-induced embolism (water potential at 50 % loss of conductivity Ψ 50) of the roots of Pinus cembra trees growing along an elevational transect of 600 m. Hydraulic parameters of the roots were compared with those of the stem and related to anatomical traits {mean conduit diameter (d), wall reinforcement [(t/b)(2)]}. We hypothesized that temperature-related restrictions in root function would cause a progressive limitation of hydraulic efficiency and safety with increasing elevation. We found that both root k s and K R decreased from low (1600 m a.s.l.: k s 5.6 ± 0.7 kg m(-1) s(-1) MPa(-1), K R 0.049 ± 0.005 kg m(-2) s (-1) MPa(-1)) to high elevation (2100 m a.s.l.: k s 4.2 ± 0.6 kg m(-1) s(-1) MPa(-1), K R 0.035 ± 0.006 kg m(-2) s(-1) MPa(-1)), with small trees showing higher K R than large trees. k s was higher in roots than in stems (0.5 ± 0.05 kg m(-1)s(-1)MPa(-1)). Ψ 50 values were similar across elevations and overall less negative in roots (Ψ 50 -3.6 ± 0.1 MPa) than in stems (Ψ 50 -3.9 ± 0.1 MPa). In roots, large-diameter tracheids were lacking at high elevation and (t/b)(2) increased, while d did not change. The elevational decrease in root hydraulic efficiency reflects a limitation in timberline tree hydraulics. In contrast, hydraulic safety was similar across elevations, indicating that avoidance of hydraulic failure is important for timberline trees. As hydraulic patterns can only partly be explained by the anatomical parameters studied, limitations and/or adaptations at the pit level are likely.

It has been frequently stressed that at distributional boundaries, like at the Alpine timberline and within dry inner Alpine environments, tree growth will be affected first by changing climate conditions. Climate in 2007 was characterized by the occurrence of exceptionally mild temperatures in spring (3.4 and 2.7 °C above long-term mean (LTM) at timberline and the valley sites, respectively) with an almost continuous drought period recorded in April and slightly warmer than average temperatures throughout summer (1.3 °C above LTM at both sites). We compared temporal dynamics of cambial activity and xylem cell development in Pinus cembra at the Alpine timberline (1950 m a.s.l.) and Pinus sylvestris at a xeric inner Alpine site (750 m a.s.l.) by repeated cellular analyses of micro-cores (n = 5 trees/site). While onset of wood formation in P. sylvestris and P. cembra differed by about two weeks (12 and 27 April, respectively), maximum daily growth rates peaked on 6 May at the valley site and on 23 June at timberline. At both sites maximum tracheid production was reached prior to occurrence of more favourable climatic conditions during summer, i.e. an increase in precipitation and temperature. Xylem formation ended on 31 August and 28 October at the xeric site and at timberline, respectively. This study demonstrates the plasticity of tree-ring formation along an altitudinal transect in response to water availability and temperature. Whether early achievement of maximum growth rates is an adaptation to cope with extreme environmental conditions prevailing at limits of tree growth needs to be analysed more closely by taking belowground carbon allocation into account.