Mobile carbohydrates in Himalayan treeline trees I. Evidence for carbon gain limitation but not for growth limitation

To test whether the altitudinal distribution of trees is determined by a carbon shortage or an insufficient sugar fraction (sugar:starch ratio) in treeline trees, we studied the status of nonstructural carbohydrates (NSC) and their components (total soluble sugars and starch) in Abies fabri (Mast.) Craib and Picea balfouriana var. hirtella Rehd. et Wils. trees along three elevational gradients, ranging from lower elevations to the alpine treeline, on the eastern edge of the Tibetan Plateau. For comparison, we investigated a low-altitude species (Tsuga yunnanensis (Franch.) Pritz.) which served as a warm-climate reference because it is distributed in closed montane forests below 3100 m a.s.l. in the study area. The carbon status of T. yunnanensis responded to altitude differently from that of the treeline species. At the species level, total NSC was not consistently more abundant in treeline trees than in trees of the same species growing at lower elevations. Thus there was no consistent evidence for carbon limitation of growth in treeline trees. For the three treeline species studied (P. balfouriana and A. fabri in the Kang-Ding Valley and A. fabri in the Mo-Xi Valley), winter NSC concentrations in treeline trees were significantly lower than in lower-elevation trees of the same species, suggesting that, in winter, carbon is limited in treeline trees. However, in no case was there total overwinter depletion of NSC or its components in treeline trees. Treeline and low-altitude species had similar sugar:starch ratios of about three at their upper-elevational limits in April. We conclude that survival and growth of trees at the elevational or latitudinal climate limit depend not only on NSC concentration in perennial tissues, but also on the maintenance of an overwintering sugar:starch ratio greater than three.     

长白山岳桦体内碳素供应状况

低温条件常常驱动的碳缺乏导致高海拔处植物生长缓慢,生长季的高寒低温更是限制高海拔地区树木向上分布的关键因素之一。高海拔地区的不利     

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration and not by growth rate, which differs from previous studies. The observed between-species difference at the intra-annual scale is key information for anticipating suitability of future species in temperate forests.     

Climate and seasonal rainfall anomalies along an elevational gradient in the El Sira Mountains,Peru, and their impacts on tree radial growth

The explicit purpose of this study was to characterize climate and vegetation along the western slope of the El Sira Mountains(Peru) and evaluate radial tree growth in response to seasonal rainfall anomalies. From May 2011 until September 2015, we monitored radial stem growth of 67 trees using point dendrometers and measured climate within five sites along an altitudinal gradient. The transect extends from lowland terra firme forests, over submontane forests, late and mid successional montane cloud forests up to exposed elfin forests. Monthly rainfall estimates by the TRMM PR satellite(product 3B42) were highly correlated with our rain gauge observations but underestimate rainfall at high altitudes. Different intra-annual tree growth patterns could be identified within each elevational forest type, showing species with strictly seasonal growth, continuous growth or alternating growth patterns independent of the seasons. Stem growth at each site was generally larger during rainy seasons, except for the elfin forest. The rainy season from October 2013 to March 2014 was extraordinarily dry, with only 73% of long-term mean precipitation received, which resulted in reduced radial growth, again with the exception of the elfin forest. This indicates that montane tropical rain forests may suffer from prolonged droughts, while exposed ridges with elfin forests still receive plenty of precipitation and benefit from receiving more solar radiation for photosynthesis.     

Nitrogen forms in bark, wood and foliage of nitrogen-fertilized Pinus sylvestris

Cycling of soluble non-protein N compounds is thought to be indicative of the N-nutritional status of trees. We determined the major N forms in bark, wood and foliage and estimated the dependence of prevalent N forms on N availability in Pinus sylvestris L. trees from northern Sweden. Trees subjected to severe N limitation and trees that had been fertilized with an average 64 kg N ha(-1) year(-1) for 25 years were analyzed. Bark and wood samples were collected by tangentially cryo-sectioning the trunk into 30-microm thick sections, from the bark to the functional xylem. Soluble amino compounds were extracted from the sections for analysis. Sap samples from twigs were obtained by centrifugation, and bark samples from twigs were obtained by tissue extraction. In both needles and bark, arginine dominated the amino-N pool. Because arginine concentrations in needles increased with N fertilization, arginine dominance of the amino-N pool in needles was higher in N-fertilized trees than in control trees. In bark, N fertilization resulted in a large increase in glutamine concentration, so that glutamine accounted for a larger proportion of the amino-N pool in bark in N-fertilized trees than in control trees. Glutamine dominated the amino-N pool in wood of control trees. Nitrogen fertilization resulted in an increased proportion of arginine in the wood amino-N pool. We conclude that the composition of the amino-N pools in bark, wood and foliage is highly sensitive to N supply. The composition of the amino-N pools can contribute to the regulation of tree N-nutritional status, which is mediated by shoot to root signalling by long-distance transport of amino compounds.     

Phenological and water-use patterns underlying maximum growing season length at the highest elevations: implications under climate change

Consequences of climate change on tree phenology are readily observable, but little is known about the variations in phenological sensitivity to drought between populations within a species. In this study, we compare the phenological sensitivity to temperature and water availability in Abies pinsapo Boiss., a drought-sensitive Mediterranean fir, across its altitudinal distribution gradient. Twig growth and needle fall were related to temperature, precipitation and plant water status on a daily scale. Stands located at the top edge of the distributional range showed the most favourable water balance, maximum growth rates and little summer defoliation. Towards higher elevations, the observed delay in budburst date due to lower spring temperatures was overcome by a stronger delay in growth cessation date due to the later onset of strong water-deficit conditions in the summer. This explains an extended growing season and the greatest mean growth at the highest elevation. Conversely, lower predawn xylem water potentials and early partial stomatal closure and growth cessation were found in low-elevation A. pinsapo trees. An earlier and higher summer peak of A. pinsapo litterfall was also observed at these water-limited sites. Our results illustrate the ecophysiological background of the ongoing altitudinal shifts reported for this relict tree species under current climatic conditions.     

A study on the relation between soil and needle content of nutrient elements in the Korean pine plantation

Through the content analysis of nutrient elements in the soil and in the needle at the different site classes and periods of growth ofP. koraiensis, and in a single growing season, some conclusions can be drawn below: Most of the soil nutrient elements were higher at the good sites than at the poor ones in an annual average concentration of them, i.e., site class 1 > site class 2> site class 3. And forP. koraiensis tree at every site class, the nutrient elements in their needles did the same. However, each of the elements in the soil or in the need was seldom at the same level of concentration on the good or on the poor site during different periods of growth. There are no an inevitable relationship between the contents of the soil nutrient elements and the needle’s. And whether these elements are high or low within a tree body may depend on the physiogical requirement of the tree, but not on their contents in the environment. Some nutrient elements required forP. koraiensis varied with growth periods during a whole growing season. N, P and K, for example, were greatly needed only at the fast growth period, while Ca at the initial growth period; and on the contrary, Ca was needed less at the fast growth period. In intensive forest management, therefore, fertilizers with N, P and K should much be applied at the fast growth period, while fertilizer with Ca at the initial growth period to meet requirements of growth ofP. koraiensis trees.     

Acclimation to low irradiance in Picea abies: influences of past and present light climate on foliage structure and function

Leaf retention time increases with decreasing irradiance, providing an effective way of amortizing the costs of foliage construction over time. To elucidate the physiological mechanisms underlying this dependence, I studied needle life span, morphology, and concentrations of carbon, nitrogen and nonstructural carbohydrates along a gradient of relative irradiance in understory trees of Picea abies (L.) Karst. Maximum needle life span was greater in shaded trees than in sun-exposed trees. However, irrespective of irradiance, needles with maximum longevity were situated in the middle rather than the bottom of the canopy, suggesting that needle life span is determined by the irradiance to which needles are exposed during their primary growth. Morphology and chemistry of current-year needles were adapted to prevailing light conditions. Current-year needles exposed to high irradiances had greater packing of foliar biomass per unit area than shaded needles, whereas shaded needles maximized foliar area to capture more light. Nitrogen concentrations were higher in shaded needles than in sun-exposed needles. This nitrogen distribution pattern was related to the high nitrogen cost of light interception and was assumed to improve light absorptance per needle mass of shaded needles. In contrast, in both 1- and 2-year-old needles, morphology was independent of prevailing light conditions; however, needle nitrogen concentrations were adjusted toward more effective light interception in 2-year-old foliage but not in 1-year-old foliage, indicating that acclimation of sun-adapted needles to shading takes more than one year. At the same time, needle aging was accompanied by accumulation of nonstructural carbohydrates (NSC), and increasing concentrations of needle carbon, suggesting a shift in the balance between photosynthesis and photosynthate export. The accumulation of NSC and carbon resulted in a dilution of the concentrations of other needle chemicals and explained the decline in needle nitrogen concentrations with increasing age. Thus, although morphological inadequacy to low light availabilities may partly be compensated for by modifications in needle chemistry, age-related changes in needle stoichiometric composition progressively lessen the potential for acclimation to low irradiance. A conceptual model, advanced to explain how environmental factors and age-related changes in the activities of needle xylem and phloem transport affect needle longevity, predicted that adaptation of needle morphology to irradiance during the primary growth period largely determines the fate of needles during subsequent tree growth and development.     

Importance of soil extractable phosphorus distribution for mature Norway spruce nutrition and productivity

Phosphorus is an essential nutrient for forest growth. In this study, we assessed the impact of soil extractable phosphorus using two simple extraction methods on nutrition and productivity of Norway spruce in sixteen mature forest stands on different bedrocks and soils in Bavaria, Southern Germany. Representative trees were sampled for needles, twigs, branches, stem bark, and stem wood. Total phosphorus content in the tree parts and soil phosphorus stock extractable with citric acid and sodium bicarbonate up to a soil depth of 80 cm were determined. We found that easily soil extractable phosphorus is a suitable indicator for estimating phosphorus uptake and stand productivity in Norway spruce. In contrast, organic layer phosphorus showed no significant correlation with aboveground biomass phosphorus contents. In the biomass, the highest phosphorus contents were measured in young needles and twigs, but the highest correlation with soil phosphorus was detected for phosphorus contents in needles and bark. The stock of phosphorus extracted by citric acid down to 40 cm soil depth revealed the best correlation with phosphorus in needles and bark. Therefore, as a supplemental or alternative method to needle analysis, our study suggests the use of phosphorus contents in stem bark to evaluate tree phosphorus nutrition. These results highlight the suitability of the citric acid soil extraction method to characterize plant available phosphorus in Norway spruce ecosystems.     

Altitudinal range shift detected through seedling survival of Ceiba aesculifolia in an area under the influence of an urban heat island

Changes in the position along the elevational gradient for plant species are a predicted consequence of global and local climate change. Within the area of influence of cities it is necessary, for conservation and ecological restoration, to understand the effect on plant communities of climate change and urban heat islands, that can increase the temperature around cities as much as 8 °C when compared with peri-urban rural areas. We explored patterns of seedling survival of Ceiba aesculifolia, a tropical tree species, along an altitudinal gradient in a slope facing the city of Morelia, in Michoacán, México, because the city has followed a trend of increasing mean annual temperatures with a difference of at least 4 °C when compared with the surrounding rural areas. The highest survival was found between 2200 and 2230 m a.s.l. (100% after 1 year of planting and 75% after 2 years), 160 m higher than the altitudinal limit of the remnant adult trees in the study area, and close to the highest limit reported for the species at a regional scale (2200 m a.s.l.). There was a significant effect of altitude on survival among experimental plots (P < 0.0001) and the interaction between elevation and distance from the north side of the experimental area was significant (P = 0.017). For restoring populations of C. aesculifolia within our study region, assisted migration through the establishment of populations at the elevational limit or higher than its historical range might be necessary in areas close to cities that already are under the effects of increased temperatures, but might be necessary also in rural areas for ameliorating the expected effects of global climate change in tropical rural areas.     

Tree- and needle-age-dependent variations in antioxidants and photoprotective pigments in Norway spruce needles at the alpine timberline

To cope with environmental stress, plants are equipped with antioxidative (e.g., ascorbate, glutathione and alpha-tocopherol) and photoprotective (e.g., xanthophyll cycle pigments) defense systems. We investigated the defense capacities of three tree age classes (mature, sapling and seedling) of Norway spruce (Picea abies (L.) Karst.) at a field site near the timberline. Biochemical data were expressed on both a needle dry mass and a surface area basis. Compared with current-year needles, previous-year needles contained higher mass- and area-based concentrations of chlorophylls and alpha-tocopherol, and a larger xanthophyll cycle pool that was in a more epoxidized state. Total glutathione concentration was lower, the glutathione pool was more reduced and the ascorbate pool was more oxidized in previous-year needles than in current-year needles. Needle concentrations of glutathione and alpha-tocopherol increased and chlorophyll concentration decreased with increasing tree age when expressed on a surface area basis. On a dry mass basis, these trends were reversed or nonexistent. The ascorbate pool was more reduced and the glutathione pool was more oxidized in needles of mature trees than in needles of saplings and seedlings. The proportion of protective xanthophyll cycle pigments decreased and the de-epoxidation state increased with increasing tree age. We conclude that tree age and the basis of expression of antioxidant concentration--surface area or dry mass--are important in scaling from seedlings to large trees.     

Canopy position and needle age affect photosynthetic response in field-grown Pinus radiata after five years of exposure to elevated carbon dioxide partial pressure

Photosynthesis of tree seedlings is generally enhanced during short-term exposure to elevated atmospheric CO2 partial pressure, but longer-term studies often indicate some degree of photosynthetic adjustment. We present physiological and biochemical evidence to explain observed long-term photosynthetic responses to elevated CO2 partial pressure as influenced by needle age and canopy position. We grew Pinus radiata D. Don. trees in open-top chambers for 5 years in sandy soil at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressures. The trees were well watered and exposed to natural light and ambient temperature. In the fourth year of CO2 exposure (fall 1997), when foliage growth had ceased for the year, photosynthetic down-regulation was observed in 1-year-old needles, but not in current-year needles, suggesting a reduction in carbohydrate sink strength as a result of increasing needle age (Turnbull et al. 1998). In 5-year-old trees (spring 1997), when foliage expansion was occurring, photosynthetic down-regulation was not observed, reflecting significantly large sinks for carbohydrates throughout the tree. Net photosynthesis was stimulated by 79% in trees growing in elevated CO2 partial pressure, but there was no significant effect on photosynthetic capacity or Rubisco activity and concentration. Current-year needles were more responsive to elevated CO2 partial pressure than 1-year-old needles, exhibiting larger relative increases in net photosynthesis to elevated CO2 partial pressure (98 versus 64%). Lower canopy and upper canopy leaves exhibited similar relative responses to growth in elevated CO2 partial pressure. However, needles in the upper canopy exhibited higher net photosynthesis, photosynthetic capacity, and Rubisco activity and concentration than needles in the lower canopy. Given that the ratio of mature to juvenile foliage mass in the canopy will increase as trees mature, we suggest that trees may become less responsive to elevated CO2 partial pressure with increasing age. We conclude that tree response to elevated CO2 partial pressure is based primarily on sink strength and not on the duration of exposure.     

The relationship between the needle age and the growth rate in Scots pine (Pinus sylvestris): a retrospective analysis by needle trace method (NTM)

Variation in annual radial and height increments (RI and HI, respectively) as well as the number of needles belonging to different needle sets of successive annual shoots, determined by the needle trace method (NTM), was studied retrospectively in Scots pine stands, one in southern Finland and two in Estonia. The connection between different needle sets and tree growth was studied by correlation and covariance analyses. The numbers of needles of the three youngest needle sets were positively correlated with growth, whereas the number of needles older than 3 years had no significant correlation. The covariance analysis emphasized the negative relationship of older needle sets with tree growth. It is generally concluded that the two or three youngest needle sets are important and all older needles minimally contribute to growth, possibly because of their small quantity. Older needles may remain on the tree if assimilation is impeded for some reason, e.g. due to premature loss in the two youngest needle sets. The trees having attached old needles grew less than those that had already totally lost these needle sets.     

Altitudinal differences in bark stripping by sika deer in the subalpine coniferous forest of Mt. Fuji

Deer expansion is a growing concern for forest ecosystem management. In Japan, upward expansion to subalpine and alpine areas has reached alarming proportions in recent years. We examined bark stripping by sika deer along an altitudinal gradient in the subalpine coniferous forest at three altitude ranges (1800-2000 m, 2000-2200 m, and 2200-2400 m) on the southern slope of Mt. Fuji. We tested differences in densities and diameter at breast height (DBH) of trees and those with bark-stripped stems of all tree species among the three altitude ranges. Then, we compared the relative densities of deer, based on pellet counts, to determine the impact of deer in relation to deer use and forest stand patterns across the altitudinal range studied. The results of the study show that differences in bark stripping by sika deer depended largely on the elevation and the species. Larger stems were sparsely distributed in the lowest elevation zone between 1800 and 2000 m. The relative density of deer was highest in the areas exhibiting high bark-stripping intensity on small regenerating trees of the dominant coniferous species, Abies veitchii, and on broadleaf species. In the highest elevation zone between 2200 and 2400 m, smaller stems were densely distributed, and the relative deer density was lowest where the bark-stripping intensity on small stems was lower for all three species studied. The damage to subalpine tree stems corresponded to the availability of palatable tree species with a small diameter, as reflected by the successional stage along the elevational gradient of subalpine forest on Mt. Fuji. These results suggest that the continuous impact of bark stripping on the dominant tree species might cause severe changes in forest succession.     

Carbon assimilation and nitrogen in needles of fertilized and unfertilized field-grown Scots pine at natural and elevated concentrations of CO2

Effects of elevated CO2 concentration ([CO2]) on carbon assimilation and needle biochemistry of fertilized and unfertilized 25-30-year-old Scots pine (Pinus sylvestris L.) trees were studied in a branch bag experiment set up in a naturally regenerated stand. In each tree, one branch was enclosed in a bag supplied with ambient [CO2] (360 micromol mol(-1)), a second branch was enclosed in a bag supplied with elevated [CO2] (680 micromol(-1)) and a control branch was left unbagged. The CO2 treatments were applied from April 15 to September 15, starting in 1993 for unfertilized trees and in 1994 for fertilized trees, which were treated with N in June 1994. Net photosynthesis, amount and activity of Rubisco, N, starch, C:N ratio and SLA of needles were measured during the growing season of 1995. Light-saturated net photosynthetic rates of 1-year-old and current-year shoots measured at ambient [CO2] were not affected by growth [CO2] or N fertilization. Elevated [CO2] reduced the amount and activity of Rubisco, and the relative proportion of Rubisco to soluble proteins and N in needles of unfertilized trees. Elevated [CO2] also reduced the chlorophyll concentration (fresh weight basis) of needles of unfertilized trees. Soluble protein concentration of needles was not affected by growth [CO2]. Elevated [CO2] decreased the Rubisco:chlorophyll ratio in unfertilized and fertilized trees. Starch concentration was significantly increased at elevated [CO2] only in 1-year-old needles of fertilized trees. Elevated [CO2] reduced needle N concentration on a dry weight or structural basis (dry weight minus starch) in unfertilized trees, resulting in an increase in needle C:N ratio. Fertilization had no effect on soluble protein, chlorophyll, Rubisco or N concentration of needles. The decrease in the relative proportions of Rubisco and N concentration in needles of unfertilized trees at elevated [CO2] indicates reallocation of N resources away from Rubisco to nonphotosynthetic processes in other plant parts. Acclimation occurred in a single branch exposed to high [CO2], despite the large sink of the tree. The responses of 1-year-old and current-year needles to elevation of growth [CO2] were similar.     

Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.     

卧龙自然保护区油竹子（Fargesia angustissima）生理生态特征的海拔变化

全球气候变化（如温度升高）迅速而强劲地影响着高海拔地区的生态环境,致使植物生长发生变化,因此高山地区成为研究植物环境适应性及其对全球气候变化响应的理想区域。为了认清气候变化如何影响竹子的生长和生理生态特性,本文系统调查研究了卧龙自然保护区内大熊猫典型主食竹种油竹子（Fargesia angustissima（Mitford）T.P.Yi）,从其天然分布下限至上限,在卧龙自然保护区沿海拔梯度研究了分株的比叶面积（SpecificLeaf Area,SLA）、基于单位叶面积和单位叶质量的叶氮含量（Narea,Nmass）,以及组织非结构性碳水化合物（NSC）含量及其组分。研究结果表明,各调查因子对海拔的响应均是非线性的,表现为随着海拔升高,各调查值先降后升,高峰值出现在分布上限区域（1 810 m）,而最低值出现在中海拔区域（1 620 m）,反映了环境因子随海拔的非线性变化。分析认为,油竹子的生长受不同海拔环境影响较大。     

Testing Binkley's hypothesis about the interaction of individual tree water use efficiency and growth efficiency with dominance patterns in open and close canopy stands

A well-recognized phenomenon during forest development is that stand level forest growth rate begins to decrease after canopy closure. Binkley [Binkley, D., 2004. A hypothesis about the interaction of tree dominance and stand production through stand development. Forest Ecology and Management 190, 265–271] proposed an interesting hypothesis relating individual contribution of trees to forest growth patterns, considering changes in dominance levels and resource use efficiency (ReUE, biomass growth per unit of resource used) of dominant vs suppressed trees. He stated that “the decline in standlevel growth near canopy closure is driven by increasing dominance of larger trees, leading to declining efficiency of resource use by smaller trees”. This decrease in ReUE of suppressed trees once canopy closes would lead to a general decline in standlevel ReUE and thus, in stand growth. This author with other colleagues found evidence according to this hypothesis, but much more research is needed to prove its generality in forests of different species and environmental conditions. The goal of this study was to test Binkley's hypothesis using information of growth patterns, water use efficiency (WUE) and growth efficiency (GE) in different stands of Pinus ponderosa in N.W. Patagonia. According to the hypothesis, we found no dominance in open stands, and some degree of dominance in closed canopy stands. However, in contrast to predictions, WUE differed between the smallest and the largest trees of each stand both with and without dominance. No differences in GE were observed between different tree size groups, and considering individual trees, the more suppressed ones were those with the highest GE. Based on our results, we proposed another hypothesis relating individual and stand ReUE: a decrease in ReUE is not determined by the establishment of dominance and subsequent decrease in suppressed trees efficiencies, but by the availability of resources for individual trees. In this regard, we postulate that differences in ReUE between trees of a stand can appear before dominance, and that these differences in ReUE lead to differences in size of the trees and later, in its social position. When trees grow, they have access to more resources, increasing at the same time, their efficiency using the resources. This efficiency is going to decrease when availability of resources decreases (for competition or environmental stress). The stand ReUE will be maintained, decreased or increased based on the balance of available resources for the individual trees, which in turn will depend on their growth rates, competitive ability and environmental site conditions.     

Minimum cuticular conductance and cuticle features of Picea abies and Pinus cembra needles along an altitudinal gradient in the Dolomites (NE Italian Alps)

Winter desiccation is believed to contribute to stress in coniferous trees growing at the treeline because cuticular conductance increases with altitude. To test whether winter desiccation occurs in high-altitude conifers of the Dolomites (NE Italian Alps), we measured minimum cuticular conductance (g(min)), needle wettability (contact angle) and cuticle thickness in Picea abies (L.) Karst. and Pinus cembra L. needles from December to August. Samples were collected from adult trees along an altitudinal gradient from valley bottom (1050 m a.s.l.) to the treeline (2170 m a.s.l.). The treeline site is one of the highest in the area and is characterized by a generally low wind exposure. Altitude had no effect on g(min) in either species. In P. abies, large seasonal variations in g(min) were recorded but no changes were related to needle age class. Pinus cembra had a low g(min) and appeared to be efficient in reducing needle water losses. There was a significant increase in g(min) with needle aging in P. cembra growing at low altitude that could be related to a shorter needle longevity compared with P. abies. High contact angles (> 110-120 ) suggested the presence of tubular epicuticular waxes on needles of both species. Contact angles were higher (low wettability) in high-altitude needles than in low-altitude needles. By the end of winter, there was no difference in contact angles between needles in the windward and leeward positions. Wax structures transformed toward planar shapes as demonstrated by the decrease in contact angle from winter to summer. In both species, the cuticle was thicker in needles of high-altitude trees than in needles of low-altitude trees and there was no correlation between g(min) and cuticle thickness. Because desiccation resistance did not decrease with altitude in either species, we conclude that they are not susceptible to winter desiccation at the tree line.     

An investigation of hydraulic limitation and compensation in large,old Douglas-fir trees

The hydraulic limitation hypothesis (Ryan and Yoder 1997) proposes that leaf-specific hydraulic conductance (kl) and stomatal conductance (gs) decline as trees grow taller, resulting in decreased carbon assimilation. We tested the hydraulic limitation hypothesis by comparison of canopy-dominant Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) trees in stands that were approximately 15 m (20 years old), 32 m (40 years old) and 60 m (> 450 years old) tall in Wind River, Washington, USA. Carbon isotope discrimination (Delta) declined with tree height (18.6, 17.6 and 15.9 per thousand for stands 15, 32 and 60 m tall, respectively) indicating that gs may have declined proportionally with tree height in the spring months, when carbon used in the construction of new foliage is assimilated. Hydraulic conductance decreased by 44% as tree height increased from 15 to > 32 m, and showed a further decline of 6% with increasing height. The general nonlinear pattern of kl versus height was predicted by a model based on Darcy's Law. Stemwood growth efficiency also declined nonlinearly with height (60, 35 and 28 g C m-2 leaf area for the 15-, 32- and 60-m stands, respectively). Unlike kl and growth efficiency, gs and photosynthesis (A) during summer drought did not decrease with height. The lack of decline in cuvette-based A indicates that reduced A, at least during summer months, is not responsible for the decline in growth efficiency. The difference between the trend in gs and A and that in kl and D may indicate temporal changes (spring versus summer) in the response of gas exchange to height-related changes in kl or it may be a result of measurement inadequacies. The formal hydraulic limitation hypothesis was not supported by our mid-summer gs and A data. Future tests of the hydraulic limitation hypothesis in this forest should be conducted in the spring months, when carbon uptake is greatest. We used a model based on Darcy's Law to quantify the extent to which compensating mechanisms buffer hydraulic limitations to gas exchange. Sensitivity analyses indicated that without the observed increases in the soil-to-leaf water potential differential (DeltaPsi) and decreases in the leaf area/sapwood area ratio, kl would have been reduced by more than 70% in the 60-m trees compared with the 15-m trees, instead of the observed decrease of 44%. However, compensation may have a cost; for example, the greater DeltaPsi of the largest trees was associated with smaller tracheid diameters and increased sapwood cavitation, which may have a negative feedback on kl and gs.