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Fine root respiration is a significant component of carbon cycling in forest ecosystems. Although fine roots differ functionally from coarse roots, these root types have been distinguished based on arbitrary diameter cut-offs (e.g., 2 or 5 mm). Fine root morphology is directly related to physiological function, but few attempts have been made to understand the relationships between morphology and respiration of fine roots. To examine relationships between respiration rates and morphological traits of fine roots (0.15-1.4 mm in diameter) of mature Quercus serrata Murr., we measured respiration of small fine root segments in the field with a portable closed static chamber system. We found a significant power relationship between mean root diameter and respiration rate. Respiration rates of roots<0.4 mm in mean diameter were high and variable, ranging from 3.8 to 11.3 nmol CO2 g(-1) s(-1), compared with those of larger diameter roots (0.4-1.4 mm), which ranged from 1.8 to 3.0 nmol CO2 g(-1) s(-1). Fine root respiration rate was positively correlated with specific root length (SRL) as well as with root nitrogen (N) concentration. For roots<0.4 mm in diameter, SRL had a wider range (11.3-80.4 m g(-1)) and was more strongly correlated with respiration rate than diameter. Our results indicate that a more detailed classification of fine roots<2.0 mm is needed to represent the heterogeneity of root respiration and to evaluate root biomass and root morphological traits.

AbstractScots pine (Pinus sylvestris L.) is one of the most important conifers in Northern Europe. In boreal forests, over one-third of net primary production is allocated to roots. Pioneer roots expand the horizontal and vertical root systems and transport nutrients and water from belowground to aboveground. Fibrous roots, often colonized by mycorrhiza, emerge from the pioneer roots and absorb water and nutrients from the soil. In this study, we installed three flatbed scanners to detect the daily growth of both pioneer and fibrous roots of Scots pine during the growing season of 2018, a year with an unexpected summer drought in Southern Finland. The growth rate of both types of roots had a positive relationship with temperature. However, the relations between root elongation rate and soil moisture differed significantly between scanners and between root types indicating spatial heterogeneity in soil moisture. The pioneer roots were more tolerant to severe environmental conditions than the fibrous roots. The pioneer roots initiated elongation earlier and ceased it later than the fibrous roots. Elongation ended when the temperature dropped below the threshold temperature of 4 °C for pioneer roots and 6 °C for fibrous roots. During the summer drought, the fibrous roots halted root surface area growth at the beginning of the drought, but there was no drought effect on the pioneer roots over the same period. To compare the timing of root production and the aboveground organs’ production, we used the CASSIA model, which estimates the aboveground tree carbon dynamics. In this study, root growth started and ceased later than growth of aboveground organs. Pioneer roots accounted for 87% of total root productivity. We suggest that future carbon allocation models should separate the roots by root types (pioneer and fibrous), as their growth patterns are different and they have different reactions to changes in the soil environment.

Our knowledge of the nature of belowground competition for moisture and nutrients is limited. In this study, we used an earth impedance method to determine the root absorbing area of Sitka spruce (Picea sitchensis (Bong.) Carr.) trees, making measurements in stands of differing density (2-, 4- and 6-m inter-tree spacing). We compared absorbing root area index (RAI(absorbing); based on the impedance measure) with fine root area index (RAI(fine); based on estimates of total surface area of fine roots) and related these results to investment in conductive roots. Root absorbing area was a near-linear function of tree stem diameter at 1.3 m height. At the stand level, RAI(absorbing), which is analogous to and scaled with transpiring leaf area index (maximum stomatal pore area per unit ground area; LAI(transpiring)), increased proportionally with basal area across the three stands. In contrast, RAI(fine) was inversely propotional to basal area. The ratio of RAI(absorbing) to LAI(transpiring) ranged from 7.7 to 17.1, giving an estimate of the relative aboveground versus belowground resource exchange areas. RAI(absorbing) provides a way of characterizing ecosystem functioning as a physiologically meaningful index of belowground absorbing area.

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.

Boreal forests are crucial to climate change predictions because of their large land area and ability to sequester and store carbon, which is controlled by water availability. Heterogeneity of these forests is predicted to increase with climate change through more frequent wildfires, warmer, longer growing seasons and potential drainage of forested wetlands. This study aims at quantifying controls over tree transpiration with drainage condition, stand age and species in a central Canadian black spruce boreal forest. Heat dissipation sensors were installed in 2007 and data were collected through 2008 on 118 trees (69 Picea mariana (Mill.) Britton, Sterns & Poggenb. (black spruce), 25 Populus tremuloides Michx. (trembling aspen), 19 Pinus banksiana Lamb. (jack pine), 3 Larix laricina (Du Roi) K. Koch (tamarack) and 2 Salix spp. (willow)) at four stand ages (18, 43, 77 and 157 years old) each containing a well- and poorly-drained stand. Transpiration estimates from sap flux were expressed per unit xylem area, J(S), per unit ground area, E(C) and per unit leaf area, E(L), using sapwood (A(S)) and leaf (A(L)) area calculated from stand- and species-specific allometry. Soil drainage differences in transpiration were variable; only the 43- and 157-year-old poorly-drained stands had ∼ 50% higher total stand E(C) than well-drained locations. Total stand E(C) tended to decrease with stand age after an initial increase between the 18- and 43-year-old stands. Soil drainage differences in transpiration were controlled primarily by short-term physiological drivers such as vapor pressure deficit and soil moisture whereas stand age differences were controlled by successional species shifts and changes in tree size (i.e., A(S)). Future predictions of boreal climate change must include stand age, species and soil drainage heterogeneity to avoid biased estimates of forest water loss and latent energy exchanges.

Increased drought frequency and severity may reshape tree species distribution in arid environments. Dioecious tree species may be more sensitive to climate warming if sex-related vulnerability to drought occurs, since lower performance of one sex may drive differential stress tolerance, sex-related mortality rates and biased sex ratios. We explored the effect of sex and environment on branch hydraulic (hydraulic conductivity and vulnerability to embolism) and trunk anatomical traits in both sexes of the dioecious conifer Juniperus thurifera L. at two sites with contrasting water availability. Additionally, we tested for a trade-off between hydraulic safety (vulnerability to embolism) and efficiency (hydraulic conductivity). Vulnerability to embolism and hydraulic conductivity were unaffected by sex or site at branch level. In contrast, sex played a significant role in xylem anatomy. We found a trade-off between hydraulic safety and efficiency, with larger conductivities related to higher vulnerabilities to embolism. At the anatomical level, females' trunk showed xylem anatomical traits related to greater hydraulic efficiency (higher theoretical hydraulic conductivity) over safety (thinner tracheid walls, lower Mork's Index), whereas males' trunk anatomy followed a more conservative strategy, especially in the drier site. Reconciling the discrepancy between branch hydraulic function and trunk xylem anatomy would require a thorough and integrated understanding of the tree structure-function relationship at the whole-plant level. Nevertheless, lower construction costs and higher efficiency in females' xylem anatomy at trunk level might explain the previously observed higher growth rates in mesic habitats. However, prioritizing efficiency over safety in trunk construction might make females more sensitive to drought, endangering the species' persistence in a drier world.

We investigated the effects of sun- and shade-shoot architecture on the photosynthetic rates of two Picea species by applying light from various angles in the laboratory. Compared with sun shoots, shade shoots were characterized by lower mass allocation per light-intercepting area, less leaf mass per shoot mass, less mutual shading among leaves and more efficient allocation of chlorophyll to photosynthesis. The shoot silhouette to total leaf area ratio (STAR(ϕ)) decreased with increasing shoot inclination angle (ϕ, the shoot axis angle relative to the projection plane) and was consistently higher for the shade shoots. Morphological and physiological characteristics of the shade shoots resulted in maximum rates of net photosynthesis at ϕ = 0° (P(max,0)) similar to that of the sun shoots when expressed on a leaf mass, total leaf area and chlorophyll basis. When the angle of incoming light was varied, P(max,ϕ) per total leaf area (P(max,ϕ )/A(T)) of the shade shoots increased linearly with increasing STAR(ϕ), while P(max,ϕ) per shoot silhouette area did not change. In contrast, the response of the sun shoots was non-linear, and an optimum angle of incoming light was determined. Our results suggest that shade-shoot morphology is adaptive for utilizing diffuse light incoming from various angles, while sun-shoot morphology is adaptive for avoiding the negative effects of strong direct radiation and for enhancing light diffusion into the canopy. We propose that the angle of incoming light should be taken into account when estimating photosynthetic rates of sun shoots of conifer trees in the field.