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1.  Spatial and seasonal variations in mobile carbohydrates in Pinus cembra in the timberline ecotone of the Central Austrian Alps 
European journal of forest research  2011;130(2):173-179.
To test whether the altitudinal limit of tree growth is determined by carbons shortage or by a limitation in growth we investigated non structural carbohydrates and their components starch and total soluble sugars in Pinus cembra trees along an elevational gradient in the timberline ecotone of the Central Austrian Alps. NSC contents in needles, branches, stems, and coarse roots were measured throughout an entire growing season. At the tissue level NSC contents were not significantly more abundant in treeline trees as compared to trees at lower elevations. Along our 425 m elevational transect from the closed forest to the treeline we failed to find a stable elevational trend in the total NSC pool of entire trees and observed within season increases in the tree’s NSC pool that can be attributed to an altitudinal increase in leaf mass as needles contained the largest NSC fraction of the whole tree NSC pool. Furthermore, whole tree NSC contents were positively correlated with net photosynthetic capacity. Although our observed NSC characteristics do not support the hypothesis that tree life at their upper elevational limit is determined by an insufficient carbon balance we found no consistent confirmation for the sink limitation hypothesis.
doi:10.1007/s10342-010-0419-7
PMCID: PMC3191523  PMID: 22003357
Non structural carbohydrates; seasonal variation; elevational gradient; timberline ecotone; treeline formation; treelife limitation
2.  Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients 
Annals of Botany  2013;112(3):623-631.
Background and Aims
The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations.
Methods
Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra).
Key Results
Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood.
Conclusions
Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.
doi:10.1093/aob/mct127
PMCID: PMC3718216  PMID: 23788748
Carbon supply; elevational gradient; Larix decidua; Nothofagus betuloides; Nothofagus pumilio; Patagonia; Pinus cembra; Pinus sylvestris; Swiss Alps; Alpine treeline
3.  Seasonal Dynamics of Mobile Carbon Supply in Quercus aquifolioides at the Upper Elevational Limit 
PLoS ONE  2012;7(3):e34213.
Many studies have tried to explain the physiological mechanisms of the alpine treeline phenomenon, but the debate on the alpine treeline formation remains controversial due to opposite results from different studies. The present study explored the carbon-physiology of an alpine shrub species (Quercus aquifolioides) grown at its upper elevational limit compared to lower elevations, to test whether the elevational limit of alpine shrubs (<3 m in height) are determined by carbon limitation or growth limitation. We studied the seasonal variations in non-structural carbohydrate (NSC) and its pool size in Q. aquifolioides grown at 3000 m, 3500 m, and at its elevational limit of 3950 m above sea level (a.s.l.) on Zheduo Mt., SW China. The tissue NSC concentrations along the elevational gradient varied significantly with season, reflecting the season-dependent carbon balance. The NSC levels in tissues were lowest at the beginning of the growing season, indicating that plants used the winter reserve storage for re-growth in the early spring. During the growing season, plants grown at the elevational limit did not show lower NSC concentrations compared to plants at lower elevations, but during the winter season, storage tissues, especially roots, had significantly lower NSC concentrations in plants at the elevational limit compared to lower elevations. The present results suggest the significance of winter reserve in storage tissues, which may determine the winter survival and early-spring re-growth of Q. aquifolioides shrubs at high elevation, leading to the formation of the uppermost distribution limit. This result is consistent with a recent hypothesis for the alpine treeline formation.
doi:10.1371/journal.pone.0034213
PMCID: PMC3316670  PMID: 22479567
4.  No evidence for depletion of carbohydrate pools in Scots pine (Pinus sylvestris L.) under drought stress 
The physiological mechanisms leading to Scots pine (Pinus sylvestris L.) decline in the dry inner Alpine valleys are still unknown. Testing the carbon starvation hypothesis, we analysed the seasonal course of mobile carbohydrate pools (NSC) of Scots pine growing at a xeric and a dry-mesic site within an inner Alpine dry valley (750 m a.s.l., Tyrol, Austria) during the year 2009, which was characterized by exceptional soil dryness. Although, soil moisture content dropped to c. 10% at both sites during the growing season, NSC concentrations were rising in all tissues (branch, stem, root) till end of July, except in needles where maxima were reached around bud break. NSC concentrations were not significantly different in the analysed tissues at the xeric and the dry-mesic site. At the dry-mesic site NSC concentrations in the above ground tree biomass were significantly higher during the period of radial growth. An accumulation of NSC in roots at the end of July indicates a change in carbon allocation after an early cessation in above ground growth, possibly due to elevated below ground carbon demand. In conclusion our results revealed that extensive soil dryness during the growing season did not lead to carbon depletion. However, even though C-reserves were not exhausted, a sequestration of carbohydrate pools during drought periods might lead to deficits in carbon supply that weaken tree vigour and drive tree mortality.
doi:10.1111/j.1438-8677.2011.00467.x
PMCID: PMC3427021  PMID: 21974742
non-structural carbohydrates; Scots pine; drought; dry inner Alpine valley; carbon starvation; tree mortality
5.  Do elevations in temperature, CO2, and nutrient availability modify belowground carbon gain and root morphology in artificially defoliated silver birch seedlings? 
Ecology and Evolution  2013;3(9):2783-2794.
Climate warming increases the risk of insect defoliation in boreal forests. Losses in photosynthetically active surfaces cause reduction in net primary productivity and often compromise carbon reserves of trees. The concurrent effects of climate change and removal of foliage on root growth responses and carbohydrate dynamics are poorly understood, especially in tree seedlings. We investigated if exposures to different combinations of elevated temperature, CO2, and nutrient availability modify belowground carbon gain and root morphology in artificially defoliated 1-year-old silver birches (Betula pendula). We quantified nonstructural carbohydrates (insoluble starch as a storage compound; soluble sucrose, fructose, and glucose) singly and in combination in fine roots of plants under winter dormancy. Also the total mass, fine root proportion, water content, and length of roots were defined. We hypothesized that the measured properties are lower in defoliated birch seedlings that grow with ample resources than with scarce resources. On average, fertilization markedly decreased both the proportion and the carbohydrate concentrations of fine roots in all seedlings, whereas the effect of fertilization on root water content and dry mass was the opposite. However, defoliation mitigated the effect of fertilization on the root water content, as well as on the proportion of fine roots and their carbohydrate concentrations by reversing the outcomes. Elevation in temperature decreased and elevation in CO2 increased the absolute contents of total nonstructural carbohydrates, whereas fertilization alleviated both these effects. Also the root length and mass increased by CO2 elevation. This confirms that surplus carbon in birch tissues is used as a substrate for storage compounds and for cell wall synthesis. To conclude, our results indicate that some, but not all elements of climate change alter belowground carbon gain and root morphology in defoliated silver birch seedlings.
doi:10.1002/ece3.665
PMCID: PMC3790529  PMID: 24101972
Betula pendula; climate change; fertilization; fine roots; folivory; plant sugar allocation
6.  Root physiological adaptations involved in enhancing P assimilation in mining and non-mining ecotypes of Polygonum hydropiper grown under organic P media 
It is important to seek out plant species, high in phosphorus (P) uptake, for phytoremediation of P-enriched environments with a large amount of organic P (Po). P assimilation characteristics and the related mechanisms of Polygonum hydropiper were investigated in hydroponic media containing various concentrations of Po (1–8 mmol L-1) supplied as phytate. The mining ecotype (ME) showed significantly higher biomass in both shoots and roots compared to the non-mining ecotype (NME) at 4, 6, and 8 m mol L-1. Shoot P content of both ecotypes increased up to 4 mmol L-1 while root P content increased continually up to 8 mmol L-1 for the ME and up to 6 mmol L-1 for the NME. Root P content of the ME exceeded 1% dry weight under 6 and 8 mmol L-1. The ME had significantly higher P accumulation in both shoots and roots compared to the NME supplied with 6 and 8 mmol L-1. The ME showed higher total root length, specific root length, root surface area, root volume, and displayed significantly greater root length, root surface area, and root volume of lateral roots compared to the NME grown in all Po treatments. Average diameter of lateral roots was 0.17–19 mm for the ME and 0.18–0.21 mm for the NME. Greater acid phosphatase and phytase activities were observed in the ME grown under different levels of Po relative to the NME. This indicated fine root morphology, enhanced acid phosphatase and phytase activities might be adaptations to high Po media. Results from this study establish that the ME of P. hydropiper is capable of assimilating P from Po media and is a potential material for phytoremediation of polluted area with high Po.
doi:10.3389/fpls.2015.00036
PMCID: PMC4316707  PMID: 25699065
organic phosphorus (Po); phytate; root morphology; acid phosphatase; phytase; phytoremediation
7.  Effects of climate variables on intra-annual stem radial increment in Pinus cembra (L.) along the alpine treeline ecotone 
Annals of forest science  2009;66(5):503.
Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.
doi:10.1051/forest/2009038
PMCID: PMC3059571  PMID: 21423861
dendrometer; Pinus cembra; radial increment; treeline ecotone; xylem formation
8.  Morphological plasticity of ectomycorrhizal short roots in Betula sp and Picea abies forests across climate and forest succession gradients: its role in changing environments 
Morphological plasticity of ectomycorrhizal (EcM) short roots (known also as first and second order roots with primary development) allows trees to adjust their water and nutrient uptake to local environmental conditions. The morphological traits (MTs) of short-living EcM roots, such as specific root length (SRL) and area, root tip frequency per mass unit (RTF), root tissue density, as well as mean diameter, length, and mass of the root tips, are good indicators of acclimation. We investigated the role of EcM root morphological plasticity across the climate gradient (48–68°N) in Norway spruce (Picea abies (L.) Karst) and (53–66°N) birch (Betula pendula Roth., B. pubescens Ehrh.) forests, as well as in primary and secondary successional birch forests assuming higher plasticity of a respective root trait to reflect higher relevance of that characteristic in acclimation process. We hypothesized that although the morphological plasticity of EcM roots is subject to the abiotic and biotic environmental conditions in the changing climate; the tools to achieve the appropriate morphological acclimation are tree species-specific. Long-term (1994–2010) measurements of EcM roots morphology strongly imply that tree species have different acclimation-indicative root traits in response to changing environments. Birch EcM roots acclimated along latitude by changing mostly SRL [plasticity index (PI) = 0.60], while spruce EcM roots became adjusted by modifying RTF (PI = 0.68). Silver birch as a pioneer species must have a broader tolerance to environmental conditions across various environments; however, the mean PI of all MTs did not differ between early-successional birch and late-successional spruce. The differences between species in SRL, and RTF, diameter, and length decreased southward, toward temperate forests with more favorable growth conditions. EcM root traits reflected root-rhizosphere succession across forest succession stages.
doi:10.3389/fpls.2013.00335
PMCID: PMC3759007  PMID: 24032035
plasticity; ectomycorrhizal roots; morphological characteristics; root acclimation; silver birch; Norway spruce; forest succession; climate gradient
9.  Interacting Microbe and Litter Quality Controls on Litter Decomposition: A Modeling Analysis 
PLoS ONE  2014;9(9):e108769.
The decomposition of plant litter in soil is a dynamic process during which substrate chemistry and microbial controls interact. We more clearly quantify these controls with a revised version of the Guild-based Decomposition Model (GDM) in which we used a reverse Michaelis-Menten approach to simulate short-term (112 days) decomposition of roots from four genotypes of Zea mays that differed primarily in lignin chemistry. A co-metabolic relationship between the degradation of lignin and holocellulose (cellulose+hemicellulose) fractions of litter showed that the reduction in decay rate with increasing lignin concentration (LCI) was related to the level of arabinan substitutions in arabinoxylan chains (i.e., arabinan to xylan or A∶X ratio) and the extent to which hemicellulose chains are cross-linked with lignin in plant cell walls. This pattern was consistent between genotypes and during progressive decomposition within each genotype. Moreover, decay rates were controlled by these cross-linkages from the start of decomposition. We also discovered it necessary to divide the Van Soest soluble (labile) fraction of litter C into two pools: one that rapidly decomposed and a second that was more persistent. Simulated microbial production was consistent with recent studies suggesting that more rapidly decomposing materials can generate greater amounts of potentially recalcitrant microbial products despite the rapid loss of litter mass. Sensitivity analyses failed to identify any model parameter that consistently explained a large proportion of model variation, suggesting that feedback controls between litter quality and microbial activity in the reverse Michaelis-Menten approach resulted in stable model behavior. Model extrapolations to an independent set of data, derived from the decomposition of 12 different genotypes of maize roots, averaged within <3% of observed respiration rates and total CO2 efflux over 112 days.
doi:10.1371/journal.pone.0108769
PMCID: PMC4181322  PMID: 25264895
10.  Radial Growth of Qilian Juniper on the Northeast Tibetan Plateau and Potential Climate Associations 
PLoS ONE  2013;8(11):e79362.
There is controversy regarding the limiting climatic factor for tree radial growth at the alpine treeline on the northeastern Tibetan Plateau. In this study, we collected 594 increment cores from 331 trees, grouped within four altitude belts spanning the range 3550 to 4020 m.a.s.l. on a single hillside. We have developed four equivalent ring-width chronologies and shown that there are no significant differences in their growth-climate responses during 1956 to 2011 or in their longer-term growth patterns during the period AD 1110–2011. The main climate influence on radial growth is shown to be precipitation variability. Missing ring analysis shows that tree radial growth at the uppermost treeline location is more sensitive to climate variation than that at other elevations, and poor tree radial growth is particularly linked to the occurrence of serious drought events. Hence water limitation, rather than temperature stress, plays the pivotal role in controlling the radial growth of Sabina przewalskii Kom. at the treeline in this region. This finding contradicts any generalisation that tree-ring chronologies from high-elevation treeline environments are mostly indicators of temperature changes.
doi:10.1371/journal.pone.0079362
PMCID: PMC3828375  PMID: 24244488
11.  Climate Warming and the Recent Treeline Shift in the European Alps: The Role of Geomorphological Factors in High-Altitude Sites 
Ambio  2010;40(3):264-273.
Global warming and the stronger regional temperature trends recently recorded over the European Alps have triggered several biological and physical dynamics in high-altitude environments. We defined the present treeline altitude in three valleys of a region in the western Italian Alps and reconstructed the past treeline position for the last three centuries in a nearly undisturbed site by means of a dendrochronological approach. We found that the treeline altitude in this region is mainly controlled by human impacts and geomorphological factors. The reconstruction of the altitudinal dynamics at the study site reveals that the treeline shifted upwards of 115 m over the period 1901–2000, reaching the altitude of 2505 m in 2000 and 2515 m in 2008. The recent treeline shift and the acceleration of tree colonization rates in the alpine belt can be mainly ascribed to the climatic input. However, we point out the increasing role of geomorphological factors in controlling the future treeline position and colonization patterns in high mountains.
doi:10.1007/s13280-010-0096-2
PMCID: PMC3357808  PMID: 21644455
Climate change; Treeline; Geomorphology; Tree rings; Larix decidua; European Alps
12.  No evidence of carbon limitation with tree age and height in Nothofagus pumilio under Mediterranean and temperate climate conditions 
Annals of Botany  2011;108(5):907-917.
Background and Aims
Trees universally decrease their growth with age. Most explanations for this trend so far support the hypothesis that carbon (C) gain becomes limited with age; though very few studies have directly assessed the relative reductions of C gain and C demand with tree age. It has also been suggested that drought enhances the effect of C gain limitation in trees. Here tests were carried out to determine whether C gain limitation is causing the growth decay with tree age, and whether drought accentuates its effect.
Methods
The balance between C gain and C demand across tree age and height ranges was estimated. For this, the concentration of non-structural carbohydrates (NSCs) in stems and roots of trees of different ages and heights was measured in the deciduous temperate species Nothofagus pumilio. An ontogenetic decrease in NSCs indicates support for C limitation. Furthermore, the importance of drought in altering the C balance with ontogeny was assessed by sampling the same species in Mediterranean and humid climate locations in the southern Andes of Chile. Wood density (WD) and stable carbon isotope ratios (δ13C) were also determined to examine drought constraints on C gain.
Key Results
At both locations, it was effectively found that tree growth ultimately decreased with tree age and height. It was found, however, that NSC concentrations did not decrease with tree age or height when WD was considered, suggesting that C limitation is not the ultimate mechanism causing the age/height-related declining tree growth. δ13C decreased with tree age/height at the Mediterranean site only; drought effect increased with tree age/height, but this pattern was not mirrored by the levels of NSCs.
Conclusions
The results indicate that concentrations of C storage in N. pumilio trees do not decrease with tree age or height, and that reduced C assimilation due to summer drought does not alter this pattern.
doi:10.1093/aob/mcr195
PMCID: PMC3177673  PMID: 21852277
Carbon isotope composition; drought; hydraulic limitation hypothesis; Mediterranean climate; non-structural carbohydrates; Nothofagus pumilio; ontogeny; Patagonia
13.  Size-dependent changes in wood chemical traits: a comparison of neotropical saplings and large trees 
AoB Plants  2013;5:plt039.
We have a fundamental and applied understanding of how differences in the wood chemistry of trees affects the durability of wood products. By comparison, relatively little is known about the ecological causes and consequences of species differences in wood chemistry; even less is known about how or why wood chemistry differs within species, across trees of different sizes. In this study we find strong and consistent differences in wood chemistry of saplings and canopy trees, in several tropical hardwood species. These differences point to the importance of pathogens and tree biomechanics as evolutionary causes of size-dependent changes in wood chemistry.
Wood anatomical traits are important correlates of life-history strategies among tree species, yet little is known about wood chemical traits. Additionally, size-dependent changes in wood chemical traits have been rarely examined, although these changes may represent an important aspect of tree ontogeny. Owing to selection for pathogen resistance and biomechanical stability, we predicted that saplings would show higher lignin (L) and wood carbon (Cconv), and lower holocellulose (H) concentrations, compared with conspecific large trees. To test these expectations, we quantified H, L and Cconv in co-occurring Panamanian tree species at the large tree vs. sapling size classes. We also examined inter- and intraspecific patterns using multivariate and phylogenetic analyses. In 15 of 16 species, sapling L concentration was higher than that in conspecific large trees, and in all 16 species, sapling H was lower than that in conspecific large trees. In 16 of 24 species, Cconv was higher in saplings than conspecific large trees. All large-tree traits were unrelated to sapling values and were unrelated to four life-history variables. Wood chemical traits did not show a phylogenetic signal in saplings, instead showing similar values across distantly related taxa; in large trees, only H showed a significant phylogenetic signal. Size-dependent changes in wood chemistry show consistent and predictable patterns, suggesting that ontogenetic changes in wood chemical traits are an important aspect of tree functional biology. Our results are consistent with the hypothesis that at early ontogenetic stages, trees are selected for greater L to defend against cellulose-decaying pathogens, or possibly to confer biomechanical stability.
doi:10.1093/aobpla/plt039
PMCID: PMC4455665
Functional traits; pathogens; phylogenetic analysis; plant defence; resource availability hypothesis; tropical forest; tropical tree; wood economics; wood traits.
14.  Direct comparison of MRI and X-ray CT technologies for 3D imaging of root systems in soil: potential and challenges for root trait quantification 
Plant Methods  2015;11:17.
Background
Roots are vital to plants for soil exploration and uptake of water and nutrients. Root performance is critical for growth and yield of plants, in particular when resources are limited. Since roots develop in strong interaction with the soil matrix, tools are required that can visualize and quantify root growth in opaque soil at best in 3D. Two modalities that are suited for such investigations are X-ray Computed Tomography (CT) and Magnetic Resonance Imaging (MRI). Due to the different physical principles they are based on, these modalities have their specific potentials and challenges for root phenotyping. We compared the two methods by imaging the same root systems grown in 3 different pot sizes with inner diameters of 34 mm, 56 mm or 81 mm.
Results
Both methods successfully visualized roots of two weeks old bean plants in all three pot sizes. Similar root images and almost the same root length were obtained for roots grown in the small pot, while more root details showed up in the CT images compared to MRI. For the medium sized pot, MRI showed more roots and higher root lengths whereas at some spots thin roots were only found by CT and the high water content apparently affected CT more than MRI. For the large pot, MRI detected much more roots including some laterals than CT.
Conclusions
Both techniques performed equally well for pots with small diameters which are best suited to monitor root development of seedlings. To investigate specific root details or finely graduated root diameters of thin roots, CT was advantageous as it provided the higher spatial resolution. For larger pot diameters, MRI delivered higher fractions of the root systems than CT, most likely because of the strong root-to-soil contrast achievable by MRI. Since complementary information can be gathered with CT and MRI, a combination of the two modalities could open a whole range of additional possibilities like analysis of root system traits in different soil structures or under varying soil moisture.
Electronic supplementary material
The online version of this article (doi:10.1186/s13007-015-0060-z) contains supplementary material, which is available to authorized users.
doi:10.1186/s13007-015-0060-z
PMCID: PMC4359488  PMID: 25774207
X-ray Computed Tomography (CT); Magnetic Resonance Imaging (MRI); Root system architecture; Common bean (Phaseolus vulgaris L.) 3D imaging; Roots in soil; Non-destructive
15.  Warming and neighbor removal affect white spruce seedling growth differently above and below treeline 
SpringerPlus  2015;4:79.
Climate change is expected to be pronounced towards higher latitudes and altitudes. Warming triggers treeline and vegetation shifts, which may aggravate interspecific competition and affect biodiversity. This research tested the effects of a warming climate, habitat type, and neighboring plant competition on the establishment and growth of white spruce (Picea glauca (Moench) Voss) seedlings in a subarctic mountain region. P. glauca seedlings were planted in June 2010 under 4 different treatments (high/control temperatures, with/without competition) in 3 habitats (alpine ridge above treeline/tundra near treeline /forest below treeline habitats). After two growing seasons in 2011, growth, photosynthesis and foliar C and N data were obtained from a total of 156, one-and-a-half year old seedlings that had survived. Elevated temperatures increased growth and photosynthetic rates above and near treeline, but decreased them below treeline. Competition was increased by elevated temperatures in all habitat types. Our results suggest that increasing temperatures will have positive effects on the growth of P. glauca seedlings at the locations where P. glauca is expected to expand its habitat, but increasing temperatures may have negative effects on seedlings growing in mature forests. Due to interspecific competition, possibly belowground competition, the upslope expansion of treelines may not be as fast in the future as it was the last fifty years.
Electronic supplementary material
The online version of this article (doi:10.1186/s40064-015-0833-x) contains supplementary material, which is available to authorized users.
doi:10.1186/s40064-015-0833-x
PMCID: PMC4339320  PMID: 25729635
Picea glauca; Boreal forest; Climate change; Competition; Subarctic; Alaska
16.  Photosynthetic responses of trees in high-elevation forests: comparing evergreen species along an elevation gradient in the Central Andes 
AoB Plants  2015;7:plv058.
Polylepis tarapacana forms the world's highest forest, being able to grow up to 5,200 m a.s.l. At such elevations, low temperatures, high solar radiation and water scarcity severely restrict plant survival. Our study was focused on the photosynthetic responses of Polylepis species and how they are able to cope with such a challenging environment. We performed all measurements and samplings in their natural environment. This strategy allowed us to observe unexpected patterns of daily adjustments in photosynthetic pigments, which reflect major changes in the structure and organization of the photosynthetic apparatus.
Plant growth at extremely high elevations is constrained by high daily thermal amplitude, strong solar radiation and water scarcity. These conditions are particularly harsh in the tropics, where the highest elevation treelines occur. In this environment, the maintenance of a positive carbon balance involves protecting the photosynthetic apparatus and taking advantage of any climatically favourable periods. To characterize photoprotective mechanisms at such high elevations, and particularly to address the question of whether these mechanisms are the same as those previously described in woody plants along extratropical treelines, we have studied photosynthetic responses in Polylepis tarapacana Philippi in the central Andes (18°S) along an elevational gradient from 4300 to 4900 m. For comparative purposes, this gradient has been complemented with a lower elevation site (3700 m) where another Polylepis species (P. rugulosa Bitter) occurs. During the daily cycle, two periods of photosynthetic activity were observed: one during the morning when, despite low temperatures, assimilation was high; and the second starting at noon when the stomata closed because of a rise in the vapour pressure deficit and thermal dissipation is prevalent over photosynthesis. From dawn to noon there was a decrease in the content of antenna pigments (chlorophyll b and neoxanthin), together with an increase in the content of xanthophyll cycle carotenoids. These results could be caused by a reduction in the antenna size along with an increase in photoprotection. Additionally, photoprotection was enhanced by a partial overnight retention of de-epoxized xanthophylls. The unique combination of all of these mechanisms made possible the efficient use of the favourable conditions during the morning while still providing enough protection for the rest of the day. This strategy differs completely from that of extratropical mountain trees, which uncouple light-harvesting and energy-use during long periods of unfavourable, winter conditions.
doi:10.1093/aobpla/plv058
PMCID: PMC4512032  PMID: 26002745
High mountain plants; light-harvesting; neoxanthin; photosynthesis; xanthophylls; zeaxanthin
17.  Temporal dynamic of wood formation in Pinus cembra along the alpine treeline ecotone and the effect of climate variables 
Trees (Berlin, Germany : West)  2009;23(3):623-635.
We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1950 m a.s.l.), treeline (2110 m a.s.l.) and within the krummholz belt (2180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined.
At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5 °C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth.
The rate of xylem cell production determined in two weekly intervals during June through August 2006-2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline.
Results of this study revealed that spatial and temporal (i.e. year-to-year) variability in timing and dynamic of wood formation of Pinus cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.
PMCID: PMC3078619  PMID: 21509148
Cambium; intra-annual growth; Pinus cembra; temperature; tracheid production
18.  Do ectomycorrhizal and arbuscular mycorrhizal temperate tree species systematically differ in root order-related fine root morphology and biomass? 
While most temperate broad-leaved tree species form ectomycorrhizal (EM) symbioses, a few species have arbuscular mycorrhizas (AM). It is not known whether EM and AM tree species differ systematically with respect to fine root morphology, fine root system size and root functioning. In a species-rich temperate mixed forest, we studied the fine root morphology and biomass of three EM and three AM tree species from the genera Acer, Carpinus, Fagus, Fraxinus, and Tilia searching for principal differences between EM and AM trees. We further assessed the evidence of convergence or divergence in root traits among the six co-occurring species. Eight fine root morphological and chemical traits were investigated in root segments of the first to fourth root order in three different soil depths and the relative importance of the factors root order, tree species and soil depth for root morphology was determined. Root order was more influential than tree species while soil depth had only a small effect on root morphology All six species showed similar decreases in specific root length and specific root area from the 1st to the 4th root order, while the species patterns differed considerably in root tissue density, root N concentration, and particularly with respect to root tip abundance. Most root morphological traits were not significantly different between EM and AM species (except for specific root area that was larger in AM species), indicating that mycorrhiza type is not a key factor influencing fine root morphology in these species. The order-based root analysis detected species differences more clearly than the simple analysis of bulked fine root mass. Despite convergence in important root traits among AM and EM species, even congeneric species may differ in certain fine root morphological traits. This suggests that, in general, species identity has a larger influence on fine root morphology than mycorrhiza type.
doi:10.3389/fpls.2015.00064
PMCID: PMC4324066  PMID: 25717334
Acer; Carpinus; Fagus; Fraxinus; mixed stand; root tips; specific root area; Tilia
19.  Nine Years of Irrigation Cause Vegetation and Fine Root Shifts in a Water-Limited Pine Forest 
PLoS ONE  2014;9(5):e96321.
Scots pines (Pinus sylvestris L.) in the inner-Alpine dry valleys of Switzerland have suffered from increased mortality during the past decades, which has been caused by longer and more frequent dry periods. In addition, a proceeding replacement of Scots pines by pubescent oaks (Quercus pubescens Willd.) has been observed. In 2003, an irrigation experiment was performed to track changes by reducing drought pressure on the natural pine forest. After nine years of irrigation, we observed major adaptations in the vegetation and shifts in Scots pine fine root abundance and structure. Irrigation permitted new plant species to assemble and promote canopy closure with a subsequent loss of herb and moss coverage. Fine root dry weight increased under irrigation and fine roots had a tendency to elongate. Structural composition of fine roots remained unaffected by irrigation, expressing preserved proportions of cellulose, lignin and phenolic substances. A shift to a more negative δ13C signal in the fine root C indicates an increased photosynthetic activity in irrigated pine trees. Using radiocarbon (14C) measurement, a reduced mean age of the fine roots in irrigated plots was revealed. The reason for this is either an increase in newly produced fine roots, supported by the increase in fine root biomass, or a reduced lifespan of fine roots which corresponds to an enhanced turnover rate. Overall, the responses belowground to irrigation are less conspicuous than the more rapid adaptations aboveground. Lagged and conservative adaptations of tree roots with decadal lifespans are challenging to detect, hence demanding for long-term surveys. Investigations concerning fine root turnover rate and degradation processes under a changing climate are crucial for a complete understanding of C cycling.
doi:10.1371/journal.pone.0096321
PMCID: PMC4011741  PMID: 24802642
20.  Variation of Maximum Tree Height and Annual Shoot Growth of Smith Fir at Various Elevations in the Sygera Mountains, Southeastern Tibetan Plateau 
PLoS ONE  2012;7(3):e31725.
Little is known about tree height and height growth (as annual shoot elongation of the apical part of vertical stems) of coniferous trees growing at various altitudes on the Tibetan Plateau, which provides a high-elevation natural platform for assessing tree growth performance in relation to future climate change. We here investigated the variation of maximum tree height and annual height increment of Smith fir (Abies georgei var. smithii) in seven forest plots (30 m×40 m) along two altitudinal transects between 3,800 m and 4,200/4,390 m above sea level (a.s.l.) in the Sygera Mountains, southeastern Tibetan Plateau. Four plots were located on north-facing slopes and three plots on southeast-facing slopes. At each site, annual shoot growth was obtained by measuring the distance between successive terminal bud scars along the main stem of 25 trees that were between 2 and 4 m high. Maximum/mean tree height and mean annual height increment of Smith fir decreased with increasing altitude up to the tree line, indicative of a stress gradient (the dominant temperature gradient) along the altitudinal transect. Above-average mean minimum summer (particularly July) temperatures affected height increment positively, whereas precipitation had no significant effect on shoot growth. The time series of annual height increments of Smith fir can be used for the reconstruction of past climate on the southeastern Tibetan Plateau. In addition, it can be expected that the rising summer temperatures observed in the recent past and anticipated for the future will enhance Smith fir's growth throughout its altitudinal distribution range.
doi:10.1371/journal.pone.0031725
PMCID: PMC3291607  PMID: 22396738
21.  Elevational patterns of Polylepis tree height (Rosaceae) in the high Andes of Peru: role of human impact and climatic conditions 
We studied tree height in stands of high-Andean Polylepis forests in two cordilleras near Cuzco (Peru) with respect to variations in human impact and climatic conditions, and compared air and soil temperatures between qualitatively defined dry and humid slopes. We studied 46 forest plots of 100 m2 of five Polylepis species at 3560–4680 m. We measured diameter at breast height (dbh) and tree height in the stands (1229 trees in total), as well as air and soil temperatures in a subset of plots. The data was analyzed combining plots of given species from different sites at the same elevation (±100 m). There was no elevational decrease of mean maximum tree height across the entire data set. On humid slopes, tree height decreased continuously with elevation, whereas on dry slopes it peaked at middle elevations. With mean maximum tree heights of 9 m at 4530 m on the humid slopes and of 13 m at 4650 m on the dry slopes, we here document the tallest high-elevation forests found so far worldwide. These highest stands grow under cold mean growing season air temperatures (3.6 and 3.8°C on humid vs. dry slopes) and mean growing season soil temperatures (5.1 vs. 4.6°C). Mean annual air and soil temperature both decreased with elevation. Dry slopes had higher mean and maximum growing season air temperatures than humid slopes. Mean annual soil temperatures did not significantly differ and mean annual air temperatures only slightly differed between slopes. However, maximum air temperatures differed on average by 6.6 K between dry and humid slopes. This suggests that the differences in tree height between the two slopes are most likely due to differences in solar radiation as reflected by maximum air temperatures. Our study furthermore provides evidence that alpine Polylepis treelines grow under lower temperature conditions than global high-elevation treelines on average, suggesting that Polylepis species may have evolved special physiological adaptations to low temperatures.
doi:10.3389/fpls.2014.00194
PMCID: PMC4023064  PMID: 24847343
air temperatures; forest structure; soil temperatures; solar radiation; alpine treeline; tropical forest
22.  Differences in Fine-Root Biomass of Trees and Understory Vegetation among Stand Types in Subtropical Forests 
PLoS ONE  2015;10(6):e0128894.
Variation of total fine-root biomass among types of tree stands has previously been attributed to the characteristics of the stand layers. The effects of the understory vegetation on total fine-root biomass are less well studied. We examined the variation of total fine-root biomass in subtropical tree stands at two sites of Datian and Huitong in China. The two sites have similar humid monsoon climate but different soil organic carbon. One examination compared two categories of basal areas (high vs. low basal area) in stands of single species. A second examination compared single-species and mixed stands with comparable basal areas. Low basal area did not correlate with low total fine-root biomass in the single-species stands. The increase in seedling density but decrease in stem density for the low basal area stands at Datian and the quite similar stand structures for the basal-area contrast at Huitong helped in the lack of association between basal area and total fine-root biomass at the two sites, respectively. The mixed stands also did not yield higher total fine-root biomasses. In addition to the lack of niche complementarity between tree species, the differences in stem and seedling densities and the belowground competition between the tree and non-tree species also contributed to the similarity of the total fine-root biomasses in the mixed and single-species stands. Across stand types, the more fertile site Datian yielded higher tree, non-tree and total fine-root biomasses than Huitong. However, the contribution of non-tree fine-root biomass to the total fine-root biomass was higher at Huitong (29.4%) than that at Datian (16.7%). This study suggests that the variation of total fine-root biomass across stand types not only was associated with the characteristics of trees, but also may be highly dependent on the understory layer.
doi:10.1371/journal.pone.0128894
PMCID: PMC4457824  PMID: 26047358
23.  DNA analysis of soil extracts can be used to investigate fine root depth distribution of trees 
AoB Plants  2015;7:plu091.
Knowledge of a tree species or cultivar's fine root distribution is important. However, the time and resource requirements of established soil core based methods where live from dead root determination is required, act as a constraint to large studies. We developed a method to determine live fine root DNA density for mango (Mangifera indica). Soil-root samples had large roots separated by sieving (min. 2 mm aperture) and soil DNA extractions made on the sieved soil containing fine root fragments that had passed through the sieve. We showed that the DNA yields of these samples could determine fine root distribution.
Understanding the root distribution of trees by soil coring is time-consuming as it requires the separation of roots from soil and classification of roots into particular size classes. This labour-intensive process can limit sample throughput and therefore sampling intensity. We investigated the use of quantitative polymerase chain reaction (qPCR) on soil DNA extractions to determine live fine root DNA density (RDD, mg DNA m−2) for mango (Mangifera indica) trees. The specificity of the qPCR was tested against DNA extracted from 10 mango cultivars and 14 weed species. All mango cultivars and no weeds were detected. Mango DNA was successfully quantified from control soil spiked with mango roots and weed species. The DNA yield of mango root sections stored in moist soil at 23–28 °C declined after 15 days to low concentrations as roots decayed, indicating that dead root materials in moist soil would not cause false-positive results. To separate large roots from samples, a root separation method for field samples was used to target the root fragments remaining in sieved (minimum 2 mm aperture) soil for RDD comparisons. Using this method we compared the seasonal RDD values of fine roots for five mango rootstock cultivars in a field trial. The mean cultivar DNA yields by depth from root fragments in the sieved soil samples had the strongest relationship (adjusted multiple R2 = 0.9307, P < 0.001) with the dry matter (g m−2) of fine (diameter <0.64 mm) roots removed from the soil by sieving. This method provides a species-specific and rapid means of comparing the distribution and concentration of live fine roots of trees in orchards using soil samples up to 500 g.
doi:10.1093/aobpla/plu091
PMCID: PMC4313792  PMID: 25552675
DNA persistence; fine-roots; root DNA concentration; root DNA density (RDD)
24.  Root biomass and soil carbon distribution in hybrid poplar riparian buffers, herbaceous riparian buffers and natural riparian woodlots on farmland 
SpringerPlus  2013;2:539.
The objectives of this study were to compare coarse root (diameter > 2 mm) and fine root (diameter < 2 mm) biomass, as well as distribution of soil carbon stocks in 3 types of riparian land uses across 4 sites located in farmland of southern Québec, Canada: (1) hybrid poplar buffers (9th growing season); (2) herbaceous buffers; (3) natural woodlots (varying in tree species and age). For all land uses most of the root biomass was within the 0–20 cm depth range. Total coarse root biomass, to a 60 cm depth, ranged from 8.8-73.7 t/ha in woodlots, 0.6-1.3 t/ha in herbaceous buffers, and 9.2-27.3 t/ha in poplars. Total fine root biomass ranged from 2.68-8.64 t/ha in woodlots, 2.60-3.29 t/ha in herbaceous buffers, and 1.86-2.62 t/ha in poplars. Total root biomass was similar or higher in poplar buffers compared to a 27 year-old grey birch forest. This indicates that poplar buffers accelerated riparian soil colonisation by roots compared to natural secondary succession. Generally, fine root biomass in the surface soil (0–20 cm) was lower in poplar than in herbaceous buffers; the reverse was observed at greater depth. Highest coarse root biomass in the 40–60 cm depth range was observed in a poplar buffer, highlighting the deep rooted nature of poplars. On average, total soil C stocks (0–60 cm) were greater in woodlots than in riparian buffers. On most sites, soil C stocks tended to be lower in poplar buffers compared to adjacent herbaceous buffers, especially in surface soil, probably because of lower fine root biomass in poplar buffers. Across all sites and land uses, highest soil C stocks at the different soil depths were found in the soil layers of woodlots that also had the greatest fine root biomass. Strong positive linear relationships between fine root biomass and soil C stocks in the 0–20 cm depth range (R2 = 0.79, p < 0.001), and in the whole soil profile (0–60 cm) (R2 = 0.65, p < 0.01), highlight the central role of fine root biomass in maintaining or increasing soil C stocks.
doi:10.1186/2193-1801-2-539
PMCID: PMC3825063  PMID: 24255839
Agroforestry; Afforestation; Agriculture; Land use; Coarse roots; Fine roots; Vertical distribution
25.  Distribution and mixing of old and new nonstructural carbon in two temperate trees 
The New Phytologist  2015;206(2):590-597.
We know surprisingly little about whole-tree nonstructural carbon (NSC; primarily sugars and starch) budgets. Even less well understood is the mixing between recent photosynthetic assimilates (new NSC) and previously stored reserves. And, NSC turnover times are poorly constrained. We characterized the distribution of NSC in the stemwood, branches, and roots of two temperate trees, and we used the continuous label offered by the radiocarbon (carbon-14, 14C) bomb spike to estimate the mean age of NSC in different tissues. NSC in branches and the outermost stemwood growth rings had the 14C signature of the current growing season. However, NSC in older aboveground and belowground tissues was enriched in 14C, indicating that it was produced from older assimilates. Radial patterns of 14C in stemwood NSC showed strong mixing of NSC across the youngest growth rings, with limited ‘mixing in’ of younger NSC to older rings. Sugars in the outermost five growth rings, accounting for two-thirds of the stemwood pool, had a mean age < 1 yr, whereas sugars in older growth rings had a mean age > 5 yr. Our results are thus consistent with a previously-hypothesized two-pool (‘fast’ and ‘slow’ cycling NSC) model structure. These pools appear to be physically distinct.
doi:10.1111/nph.13273
PMCID: PMC4405048  PMID: 25558814
carbohydrates; carbon allocation; Harvard Forest; radiocarbon (14C); storage; tree rings; wood

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