Seasonal dynamics of non-structural carbon pools and their relationship to growth in two boreal conifer tree species.

In an attempt to comprehensively study the dynamics of non-structural carbon compounds (NCCs), we measured the seasonal changes of soluble sugars, starch, lipids and sugar alcohols in the leaves, branches, stem and roots of the fast-growing Pinus contorta (Loudon) (pine) and slow-growing Picea glauca (Moench) Voss (spruce) trees growing in a boreal climate. In addition to measuring the seasonal concentrations of these compounds, the relative contribution of these compounds to the total NCC pool within the organs of trees (~8 m tall) was estimated and compared across different phenological and growth stages. Both species showed large seasonal shifts from starch to sugars from spring to fall in nearly all organs and tissues; most likely an adaptation to the cold winters. For both species, the total fluctuation of sugar + starch across the year (i.e., the difference between the minimum and maximum observed across collection times) was estimated to be between 1.6 and 1.8 kg for all NCCs. The fluctuation, however, was 1.40 times greater than the minimum reserves in pine, while only 0.72 times the minimum reserves in spruce. By tissue type, NCC fluctuations were greatest in the roots of both species. Roots showed a large build-up of reserves in late spring, but these reserves were depleted over summer and fall. Storage reserves in needles and branches declined over the summer, and this decline may be linked to the sink strength of the stem during diameter growth. Some notable highlights of this holistic study: a late winter build-up of sugars in the stem xylem of both species, but especially spruce; and an increase in sugar alcohols in the bark of spruce in very late winter, which could indicate mobilization to support early growth in spring and high lipid reserves in the bark of pine, which appeared not to be impacted by seasonal changes between summer and winter. Collectively, these observations point toward a more conservative NCC reserve strategy in spruce compared with pine, which is consistent with its stress tolerance and greater longevity.

Functional xylem anatomy of aspen exhibits greater change due to insect defoliation than to drought.

The study of tree rings can reveal long-term records of a tree's response to the environment. This dendroecological approach, when supplemented with finer-scale observations of the xylem anatomy, can provide novel information about a tree's year-to-year anatomical and hydraulic adjustments. Here we use this method in aspen (Populus tremuloides Michx.) to identify xylem response to drought and insect defoliation. Surprisingly, we found that precipitation influenced vessel diameter mostly in the trees' youth, while this correlation was less pronounced at maturity. This is likely due to a reduction in stress the stand experiences as it ages, and reflects an ability to mediate drought stress as trees mature. Defoliation events caused consistent and profound changes in fiber anatomy likely leading to reduced structural support to vessels. We therefore expect that in years of defoliation trees may be vulnerable to drought-induced cavitation when leaf area recovers. This study highlights how the inclusion of cellular level measurements in tree ring studies provides additional information on how stress events may alter tree functioning through alterations in structure.

Viewing forests from below: fine root mass declines relative to leaf area in aging lodgepole pine stands.

In the continued quest to explain the decline in productivity and vigor with aging forest stands, the most poorly studied area relates to root system change in time. This paper measures the wood production, root and leaf area (and mass) in a chronosequence of fire-origin lodgepole pine (Pinus contorta Loudon) stands consisting of four age classes (12, 21, 53, and ≥100 years), each replicated ~ five times. Wood productivity was greatest in the 53-year-old stands and then declined in the ≥100-year-old stands. Growth efficiency, the quantity of wood produced per unit leaf mass, steadily declined with age. Leaf mass and fine root mass plateaued between the 53- and ≥100-year-old stands, but leaf area index actually increased in the older stands. An increase in the leaf area index:fine root area ratio supports the idea that older stand are potentially limited by soil resources. Other factors contributing to slower growth in older stands might be lower soil temperatures and increased self-shading due to the clumped nature of crowns. Collectively, the proportionally greater reduction in fine roots in older stands might be the variable that predisposes these forests to be at a potentially greater risk of stress-induced mortality.

Hydraulic conductance in aspen (Populus tremuloides) seedlings exposed to low root temperatures.

Low root temperatures significantly reduced root hydraulic conductivity and increased resistance to water flow through the roots of aspen (Populus tremuloides Michx.) seedlings. Increased resistance to water flow could not be fully explained by the corresponding increase in water viscosity at low temperatures. The shapes of Arrhenius plots of root water flow and the activation energies were dependent on the direction, sequence and extent of temperature change. The Arrhenius plots suggested that the effect of low root temperature on root water flow was mediated by an effect on root metabolism. The low root temperatures tested did not induce root electrolyte leakage normally associated with cell membrane injury. Although a decrease in root temperatures to 7 or 4 degrees C induced a reduction in stomatal conductance, this reduction lagged the decline in root water flow by several hours. In contrast, when soil temperatures were raised from 4 or 7 degrees C to 25 degrees C, root water flow presumably increased, and stomatal conductance responded rapidly and was temporarily higher than before the cold treatment was imposed.

Photosynthesis and carbon allocation of six boreal tree species grown in understory and open conditions.

One-year-old seedlings of Abies balsamea (L.) Mill, Picea glauca (Moench) Voss, Pinus contorta Loudon, Betula papyrifera Marsh., Populus tremuloides Michx. and Populus balsamifera L. were transplanted in the spring, in pots, to the understory of a mixed P. tremuloides-P. balsamifera stand or to an adjacent open site. Growth and leaf characteristics were measured and photosynthetic light response curves determined in mid-August. Overall, the coniferous seedlings showed less photosynthetic plasticity in response to growth conditions than the deciduous species. Abies balsamea, P. glauca and B. papyrifera responded to the understory environment with higher leaf area ratios, and lower photosynthetic light saturation points and area-based leaf respiration relative to values for open-grown seedlings, while they matched or exceeded the height growth of open-grown seedlings. In contrast, seedlings of Pinus contorta, P. tremuloides and P. balsamifera displayed characteristics that were not conducive to survival in the understory. These characteristics included a high light saturation point and leaf dark respiration rate in P. contorta, and lower leaf area variables combined with higher carbon allocation to roots in P. tremuloides and P. balsamifera. By the second growing season, all seedlings of P. tremuloides and P. balsamifera growing in the understory had died.

Seasonal photosynthetic responses to light and temperature in white spruce (Picea glauca) seedlings planted under an aspen (Populus tremuloides) canopy and in the open.

Photosynthetic light and temperature response curves were measured seasonally in seedlings of white spruce (Picea glauca (Moench.) Voss) grown for two years in the understory of aspen (Populus tremuloides Michx.) or in the open in central Alberta. Light-saturated rate of net photosynthesis, the optimum temperature for net photosynthesis, transpiration rate, photochemical efficiency, and stomatal and mesophyll conductances increased from spring to summer and declined thereafter, whereas dark respiration rate and compensation and saturation points were highest in spring. Depression of photosynthetic parameters was greater in open-grown seedlings than in understory seedlings during the periods in spring and autumn when night frosts were common. Net photosynthetic rates were similar in understory and open-grown seedlings in summer, but they were significantly lower in open-grown seedlings in spring and autumn. Significantly lower transpiration rates and stomatal conductances in open-grown seedlings than in understory seedlings were also observed at 15 and 25 degrees C in the autumn. Shoot and needle growth were less in open-grown seedlings than in understory seedlings. In summer, when irradiances were low in the aspen understory, understory white spruce seedlings maintained a positive carbon balance by decreasing their compensation and saturation points and increasing their photochemical efficiency compared to spring and autumn.

Ecophysiology of shade needles of Picea glauca saplings in relation to removal of competing hardwoods and degree of prior shading.

We studied shade needles of Picea glauca (Moench) Voss saplings that were growing in 15, 45 or 90% shade from competing vegetation at northern boreal forest sites in Alberta and Saskatchewan. At each site, in late May or early June 1990, all hardwoods were removed within a 2-m radius of each of eight saplings in each shade treatment (released saplings), and eight saplings in each shade treatment were left as controls. Light-saturated net assimilation (NA), stomatal and mesophyll conductance and water use efficiency of one-year-old needles were measured four times during the 1990 growing season and in the spring of 1991. There was a trend of increased photosynthetic capacity within one week following release. By August 1990 and in the following spring, NA was higher in released trees than in controls. The increase in NA appeared to be related to increased stomatal conductance to water vapor and to increased foliar nitrogen and resulting increases in mesophyll conductance to CO(2). There was no measurable effect of degree of shading prior to release on NA following release. Foliage of the released saplings appeared capable of rapid acclimation to the open conditions.

Seasonal changes in shoot regrowth potential in Calamagrostis canadensis.

A series of laboratory experiments was conducted to examine seasonal change in shoot regrowth potential following disturbance in Calamagrostis canadensis. On several dates during the 1988 and 1989 growing seasons, soil cores were collected from field sites dominated by this grass. Shoot regrowth from cores after clipping at the soil surface was monitored under dark or light laboratory conditions at 20°C. seasonal changes in field concentrations of total nonstructural carbohydrate and nitrogen in rhizomes largely accounted for the observed seasonal change in etiolated regrowth potential of shoots in laboratory experiments. In contrast, shoot regrowth potential in the light showed a very different seasonal pattern. The ratio of shoot biomass regrowth 20 d after clipping in the light versus dark treatment showed a gradual seasonal decrease from 12:1 in the early May experiment to near 1:1 in the September experiment. However, the rate of photosynthesis of regrowing shoots in the light was highest in experiments conducted late in the growing season. This may indicate a strong seasonal decrease in the proportion of current photosynthate of regrowing shoots that is allocated to new shoot growth. Alternatively, mobilization of rhizome carbohydrate reserves for shoot regrowth may have been inhibited during the re-establishment of photosynthesis in the light treatment. Either mechanism would explain why shoot regrowth in the light is poorly correlated with levels of belowground carbohydrate reserves, even under controlled laboratory conditions.

Diurnal variation and interrelations of ecophysiological parameters in three peatland woody species under different weather and soil moisture conditions.

The diurnal patterns of twig xylem water potential, net photosynthesis rate, water use efficiency of photosynthesis, and stomatal and mesophyll conductance to CO2 in tamarack, black spruce and swamp birch growing in a natural peatland in central Alberta, Canada, were examined. The relationships of photosynthesis to other ccophysiological parameters were investigated. Data were collected on three days with different weather and soil moisture conditions in the 1988 growing season. Day 1 was clear and warm and the ground water table was 7 cm above the average peat surface. Day 2 was clear and hot. Day 3 was cloudy but warm. On day 2 and day 3, the water tables were in the normal range for that season. Major findings were: 1) Soil flooding depressed photosynthesis in tamarack and black spruce. 2) Swamp birch was better adapted to flooding than tamarack or black spruce. 3) The trees experienced water stress in the afternoons of the two days with lower water table. 4) Changes in photosynthesis of the three species were primarily affected by changes in mesophyll conductance (gm) and the response of photosynthesis to changes in gm was similar for all three species.