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Plant resorption of multiple nutrients during leaf senescence has been established but stoichiometric changes among N, P and K during resorption and after fertilization are poorly understood. We anticipated that increased N supply would lead to further P limitation or co-limitation with N or K [i.e. P-(co)limitation], decrease N resorption and increase P and K resorption, while P and K addition would decrease P and K resorption and increase N resorption. Furthermore, Ca would accumulate while Mg would be resorbed during leaf senescence, irrespective of fertilization. We investigated the effect of N, P and K addition on resorption in two evergreen shrubs ( Chamaedaphne calyculata and Rhododendron groenlandicum) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada. In general, N addition caused further P-(co)limitation, increased P and K resorption efficiency but did not affect N resorption. P and K addition did not shift the system to N limitation and affect K resorption, but reduced P resorption proficiency. C. calyculata resorbed both Ca and Mg while R. groenlandicum resorbed neither. C. calyculata showed a higher resorption than R. groenlandicum, suggesting it is better adapted to nutrient deficiency than R. groenlandicum. Resorption during leaf senescence decreased N:P, N:K and K:P ratios. The limited response of N and K and the response of P resorption to fertilization reflect the stoichiometric coupling of nutrient cycling, which varies among the two shrub species; changes in species composition may affect nutrient cycling in bogs. [ABSTRACT FROM AUTHOR]

Plants in nutrient-poor environments typically have low foliar nitrogen (N) concentrations, long-lived tissues with leaf traits designed to use nutrients efficiently, and low rates of photosynthesis. We postulated that increasing N availability due to atmospheric deposition would increase photosynthetic capacity, foliar N, and specific leaf area (SLA) of bog shrubs. We measured photosynthesis, foliar chemistry and leaf morphology in three ericaceous shrubs ( Vaccinium myrtilloides, Ledum groenlandicum and Chamaedaphne calyculata) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada, with a background deposition of 0.8 g N m a. While biomass and chlorophyll concentrations increased in the highest nutrient treatment for C. calyculata, we found no change in the rates of light-saturated photosynthesis ( A), carboxylation ( V) or SLA with nutrient (N with and without PK) addition, with the exception of a weak positive correlation between foliar N and A for C. calyculata, and higher V in L. groenlandicum with low nutrient addition. We found negative correlations between photosynthetic N use efficiency (PNUE) and foliar N, accompanied by a species-specific increase in one or more amino acids, which may be a sign of excess N availability and/or a mechanism to reduce ammonium (NH) toxicity. We also observed a decrease in foliar soluble Ca and Mg concentrations, essential minerals for plant growth, but no change in polyamines, indicators of physiological stress under conditions of high N accumulation. These results suggest that plants adapted to low-nutrient environments do not shift their resource allocation to photosynthetic processes, even after reaching N sufficiency, but instead store the excess N in organic compounds for future use. In the long term, bog species may not be able to take advantage of elevated nutrients, resulting in them being replaced by species that are better adapted to a higher nutrient environment. [ABSTRACT FROM AUTHOR]

The boreal forest is of particular interest to climate change research due to its large circumpolar distribution and accumulated soil carbon pool. Carbon uptake in this ecosystem is nitrogen (N)-limited, therefore factors affecting carbon or nitrogen dynamics in the boreal forest can have consequences for global climate. We used a 2-year field experiment to investigate the response of biotic nitrogen fixation by cyanobacteria associated with boreal forest bryophytes, in a factorial experiment combining simulated climate change with habitat fragmentation treatments. We simulated climate change conditions using open-top greenhouse chambers in the field, which increased mean and maximum temperatures, and created a precipitation gradient from ambient levels in the center to extreme drought conditions at the periphery of the chamber. The dry patches near the chamber walls exhibited almost no N-fixation, despite having similar densities of cyanobacteria (predominantly Stigonema sp.) as other patches. Rates of N-fixation were best explained by a model containing moisture, fragmentation, cyanobacteria density and time; warming was not a significant variable affecting N-fixation. There was no significant interaction between warming and fragmentation. These results suggest that cyanobacteria responded physiologically to drought by reducing N-fixation activity long before any changes in density. Ecosystem processes, such as N-fixation, can respond in the short term to environmental change much more rapidly than changes in the underlying community structure. Such rapid physiological responses may occur faster than demographic insurance effects of biodiversity. [ABSTRACT FROM AUTHOR]

Respiration is instrumental for survival and growth of plants, but increasing costs of maintenance processes with warming have the potential to change the balance between photosynthetic carbon uptake and respiratory carbon release from leaves. Climate warming may cause substantial increases of leaf respiratory carbon fluxes, which would further impact the carbon balance of terrestrial vegetation. However, downregulation of respiratory physiology via thermal acclimation may mitigate this impact. We have conducted a meta-analysis with data collected from 43 independent studies to assess quantitatively the thermal acclimation capacity of leaf dark respiration to warming of terrestrial plant species from across the globe. In total, 282 temperature contrasts were included in the meta-analysis, representing 103 species of forbs, graminoids, shrubs, trees and lianas native to arctic, boreal, temperate and tropical ecosystems. Acclimation to warming was found to decrease respiration at a set temperature in the majority of the observations, regardless of the biome of origin and growth form, but respiration was not completely homeostatic across temperatures in the majority of cases. Leaves that developed at a new temperature had a greater capacity for acclimation than those transferred to a new temperature. We conclude that leaf respiration of most terrestrial plants can acclimate to gradual warming, potentially reducing the magnitude of the positive feedback between climate and the carbon cycle in a warming world. More empirical data are, however, needed to improve our understanding of interspecific variation in thermal acclimation capacity, and to better predict patterns in respiratory carbon fluxes both within and across biomes in the face of ongoing global warming. [ABSTRACT FROM AUTHOR]

C4 plants are rare in cool climates, an ecological pattern attributable to their poor photosynthetic performance at low temperatures relative to C3 species. However, some C4 species are able to persist at high latitudes and high elevations, possibly due to the characteristics of the particular microsites they inhabit in these otherwise unfavourable environments. One such species is Muhlenbergia glomerata, which occurs above 60°N in Canada and is found in the atypical C4 habitat of boreal fens. In this study, we evaluate how microsite features affect the success of M. glomerata in boreal fens. We surveyed 19 populations across northern Ontario during the summers of 1999 and 2000. The ground coverage by woody vegetation was the most important parameter affecting the presence or absence of M. glomerata. Woody plants covered over 50% of the ground area in plots where M. glomerata is absent, but less than 20% where it is present. The minimum light intensity threshold for the presence of the C4 species was about 32% of full-sunlight at plant height. Surprisingly, in boreal fens M. glomerata was largely restricted to the wetter moss hollows, rather than occurring on the dry hummocks where its greater water use efficiency might have been advantageous. Woody species dominated the hummocks, but were uncommon in the hollows. In these cool northern climates M. glomerata apparently persists because sufficient periods of temperatures favourable to C4 photosynthesis occur, but this persistence likely requires some factor that suppresses the woody vegetation. [ABSTRACT FROM AUTHOR]

Boreal black spruce forests typically have a dense ground cover of bryophytes. The two main bryophyte groups in boreal black spruce forests, feathermoss and Sphagnum, have ecophysiological characteristics that influence the biogeochemical cycles of black spruce forests differently. The objective of this study was to examine the environmental controls of ground cover composition and net primary production (NPP) of feathermoss and Sphagnum in a boreal black spruce forest in central Saskatchewan. The fraction of Sphagnum ground cover was positively correlated to canopy photosynthetically active radiation (PAR) transmittance (r2=0.48, P=0.03), but the fraction of feathermoss ground cover was negatively correlated to canopy PAR transmittance in plots where Sphagnum was present (r2=0.87, P<0.0001). Sphagnum presence was inversely correlated (P=0.0001) to water table index, defined as water table depth relative to the peat layer, while feathermoss occurred in a wider range of microenvironments. Average NPP for 1998 was more than three times greater for Sphagnum (77 g C m–2 year–1) than feathermoss (24 g C m–2 year–1), but the average bryophyte NPP for 1998 was 25 g C m–2 year–1 because feathermoss was the dominant ground cover. The large, but differing, peat carbon content of Sphagnum- versus feathermoss-dominated boreal forests and peatlands necessitates the need to accurately quantify fraction ground cover. Additional validation of the empirical models between environmental variables and fraction ground cover of bryophytes is necessary, but the reported relationships offer an approach to model carbon dynamics of bryophytes in boreal forests and peatlands. [ABSTRACT FROM AUTHOR]

Boreal peatlands, which contain a large fraction of the world's soil organic carbon pool, may be significantly affected by changes in climate and land use, with attendant feedback to climate through changes in albedo, fluxes of energy or trace gases, and soil carbon storage. The response of peatlands to changing environmental conditions will probably be dictated in part by scale-dependent topographic heterogeneity, which is known to interact with hydrology, vegetation, nutrients, and emissions of trace gases. Because the bryophyte community can contribute the majority of aboveground production in bogs, we investigated how microscale topography affects the response of bryophyte species production and cover to warming (using overhead infrared lamps) and manipulations of water-table height within experimental mesocosms. We removed 27 intact peat monoliths (2.1-m2 surface area, 0.5–0.7 m depth) from a bog in northern Minnesota, USA, and subjected them to three warming and three water-table treatments in a fully crossed factorial design. Between 1994 and 1998, we determined annual production of the four dominant bryophyte taxa within three microtopographic zones (low, medium, and high relative to the water table). We also estimated species cover and calculated changes in topography and roughness of the bryophyte surface through time. Total production of all bryophytes, and production of the individual taxa Polytrichum strictum, Sphagnum magellanicum, and Sphagnum Section Acutifolia, were about 100% greater in low microtopographic zones than in high zones, and about 50% greater in low than in medium zones. Production of bryophytes increased along the gradient of increasing water-table heights, but in most years, total production of bryophytes was negatively correlated with height above the set water table only for the wettest water-table treatment. Although bryophyte production was unaffected by the warming treatments, the bryophyte surface flattened in proportion to the degree of warming. These results indicate that production of bryophytes is driven most strongly by the absolute and relative height of the bryophyte surface above the water table. Predicted changes in water-table height commensurate with changes in surface temperature may thus affect both production and superficial topography of bryophyte communities. [ABSTRACT FROM AUTHOR]

Theories on allelochemical concentrations in plants are often based upon the relative carbon costs and benefits of multiple metabolic fractions. Tests of these theories often rely on measuring metabolite concentrations, but frequently overlook priorities in carbon partitioning. We conducted a pulse-labeling experiment to follow the partitioning of 14CO2-labeled photosynthate into ten metabolic pools representing growth and maintenance (amino acids, organic acids, lipids plus pigments, protein, residue), defense (phenolic glycosides, methanol:water and acetone-soluble tannins/phenolics), and transport and storage (sugars and starch) in source and importing sink leaves of quaking aspen ( Populus tremuloides). The peak period of 14C incorporation into sink leaves occurred at 24 h. Within 48 h of labeling, the specific radioactivity (dpm/mg dry leaf weight) of phenolic glycosides declined by over one-third in source and sink leaves. In addition, the specific radioactivity in the tannin/phenolic fraction decreased by 53% and 28% in source and sink leaves, respectively. On a percent recovery basis, sink leaves partitioned 1.7 times as much labeled photosynthate into phenolic glycosides as source leaves at peak 14C incorporation. In contrast, source leaves partitioned 1.8 times as much 14C-labeled photosynthate into tannins/phenolics as importing sink leaves. At the end of the 7-day chase period, sink leaves retained 18%, 52%, and 30% of imported 14C photosynthate, and labeled source leaves retained 15%, 66%, and 19% of in situ photosynthate in metabolic fractions representing transport and storage, growth and maintenance, and defense, respectively. Analyses of the phenolic fractions showed that total phenolics were twice as great and condensed tannins were 1.7 times greater in sink than in source leaves. The concentration of total phenolics and condensed tannins did not change in source and sink leaves during the 7-day chase period. [ABSTRACT FROM AUTHOR]

Carbon dioxide exchange rates in excised 2-year-old shoot sections of five common moss species were measured by infrared gas analysis in mosses collected from different stands of mature vegetation near Fairbanks, Alaska. The maximum rates of net photosynthesis ranged from 2.65 mg CO gh in Polytrichum commune Hedw. to 0.25 in Spagnum nemoreum Scop. Intermediate values were found in Sphagnum subsecundum Nees., Hylocomium splendens (Hedw.) B.S.G., and Pleurozium schreberi (Brid.) Mitt. Dark respiration rates at 15°C ranged from 0.24 mg CO gh in S. subsecundum to 0.57 mg CO gh in H. splendens. The dark respiration rates were found to increase in periods of growth or restoration of tissue (i.e., after desiccation). There was a strong decrease in the rates of net photosynthesis during the winter and after long periods of desiccation. Due to increasing amounts of young, photosynthetically active tissue there was a gradual increase in the rates of net photosynthesis during the season to maximum values in late August. As an apparent result of constant respiration rates and increasing gross photosynthetic rates, the optimum temperature for photosynthesis at light saturation and field capacity increased during the season in all species except Polytrichum, with a corresponding drop in the compensation light intensities. Sphagnum subsecundum seemed to be the most light-dependent species. Leaf water content was found to be an important limiting factor for photosynthesis in the field. A comparison between sites showed that the maximum rates of net photosynthesis increased with increasing nutrient content in the soil but at the permafrostfree sites photosynthesis was inhibited by frequent moisture stress. [ABSTRACT FROM AUTHOR]

Plants of the widely distributed species Heliotropium curassavicum L. have a large photosynthetic acclimation potential to temperature. There are, however, some differences among the acclimation potentials of populations occupying dissimilar thermal regimes. Plants of populations originating from a cool maritime climate have a greater acclimation potential than plants of populations originating from a desert habitat, which is characterized by large seasonal changes in temperature. [ABSTRACT FROM AUTHOR]

Soils of three Picea mariana-Cladina stellaris woodland sites, aged 2, 104 and 137 years after fire, were sampled at 3 cm intervals to 9 cm depth for soil microfungi. Mortierella, Mucor, Penicillium and Trichoderma were the most frequently isolated genera. The species composition of the microfungi remained much the same across the age gradient. Direct and indirect gradient analyses of the microfungal species and 14 soil environmental factors indicate that the frequencies of occurrence of the microfungi are influenced by environmental factors primarily related to the build-up of the organic layer and its destruction by fire. [ABSTRACT FROM AUTHOR]

We tested the hypothesis that life forms (trees, shrubs, forbs, and mosses; deciduous or evergreen) can be used to group plants with similar physiological characteristics. Carbon isotope ratios (δ13C) and carbon isotope discrimination (Δ) were used as functional characteristics because δ13C and Δ integrate information about CO2 and water fluxes, and so are useful in global change and scaling studies. We examined δ13C values of the dominant species in three boreal forest ecosystems: wet Picea mariana stands, mesic Populus tremuloides stands, and dry Pinus banksiana stands. Life form groups explained a significant fraction of the variation in leaf carbon isotope composition; seven life-form categories explained 50% of the variation in δ13C and 42% of the variation in Δ and 52% of the variance not due to intraspecific genetic differences ( n=335). The life forms were ranked in the following order based on their values: evergreen trees2 ( ci/ ca) than did evergreen leaves in a similar environment within these boreal ecosystems. We found the same pattern of carbon isotope discrimination in a year with above-average precipitation as in a year with below-average precipitation, indicating that environmental fluctuations did not affect the ranking of life forms. Furthermore, plants from sites near the northern and southern boundaries of the boreal forest had similar patterns of discrimination. We concluded that life forms are robust indicators of functional groups that are related to carbon and water fluxes within boreal ecosystems. [ABSTRACT FROM AUTHOR]

Measurements of net ecosystem CO exchange by eddy correlation, incident photosynthetically active photon flux density (PPFD), soil temperature, air temperature, and air humidity were made in a black spruce ( Picea mariana) boreal woodland near Schefferville, Quebec, Canada, from June through August 1990. Nighttime respiration was between 0.5 and 1.5 kg C ha h, increasing with temperature. Net uptake of carbon during the day peaked at 3 kg C ha h, and the daily net uptake over the experiment was 12 kg C ha day. Photosynthesis dropped substantially at leaf-to-air vapor pressure deficit (VPD) greater than 7 mb, presumably as a result of stomatal closure. The response of ecosystem photosynthesis to incident PPFD was markedly non-linear, with an abrupt saturation at 600 μmol m s. This sharp saturation reflected the geometry of the spruce canopy (isolated conical crowns), the frequently overcast conditions, and an increase in VPD coincident with high radiation. The ecosystem light-use efficiency increased markedly during overcast periods as a result of a more even distribution of light across the forest surface. A mechanistic model of forest photosynthesis, parameterized with observations of leaf density and nitrogen content from a nearby stand, provided accurate predictions of forest photosynthesis. The observations and model results indicated that ecosystem carbon balance at the site is highly sensitive to temperature, and relatively insensitive to cloudiness. [ABSTRACT FROM AUTHOR]

The effect of N supply on shoot and leaf lifespan was investigated in established stands of four herbaceous Carex species which differed in maximum dry matter production. These species were, in rank order of increasing maximum dry matter production (per unit ground area): Carex diandra≤C. rostrata

This study evaluated the contribution of different ages of foliage to the nutrient and carbon balance of black spruce ( Picea mariana (Mill.) B.S.P.) from a nutrient-poor peatland in Alberta. Seasonal patterns of foliar nitrogen and phosphorus concentration and content were examined in six needle cohorts up to 10 years old. Trees were treated to simulate excess nutrient deficiency (removal of all one-year-old foliage), nutrient excess (fertilized with 250, 50, 100 kg ha NPK split application in June and July), or left as controls. Gas exchange (net assimilation-Na, stomatal conductance-g, mesophyll conductance-g, water-use efficiency-WUE, dark respiration-RS) was measured on six different needle cohorts in several control trees in 1989 and 1990. Nitrogen and phosphorus concentration decreased with needle age. Foliar nutrient concentration fell from April to June and then was stable until September except for the fertilized trees where it increased. There was no evidence of greater than normal retranslocation of nutrients from older needles for defoliated trees or greater than normal nutrient loading in older needles of fertilized trees. NA, g, g, WUE, and RS were similar for all needles up to six or eight years old, these older needles having NA of 65% of current needles and similar RS. The results do not support to conclusion that older needles of black spruce are retained as an adaptation to nutrient stress. It does not appear that older needles serve as a nutrient storage site in conditions of excess nutrient availability or a greater than normal nutrient source during times of excess nutrient deficiency. It appears that the maintenance of long-livedfoliage in black spruce does not provide for greater flexibility in tree nutrient allocation. Their contribution to the carbon balance of the tree seems to be sufficient to explain their retention. [ABSTRACT FROM AUTHOR]

A small caribou herd of 24-77 animals resided on Pic Island (1138 ha) in Lake Superior from 1976 to 1984. Most of the caribou populations on the adjacent mainland had gone extinct earlier in this century. We tested three hypotheses for the persistence of this remanant population: (1) there was more forage available on the island than the mainland, (2) the animals on the island were isolated from white-tailed deer and did not develop the meningeal worm infection, and (3) there was less predation by wolves on the island. Forage was more abundant on the mainland than on Pic Island. The eggs of meningeal worms were not found in the feces of deer on the mainland or Pic island. Wolves seldom visited the island and when they did there was escape habitat available for the caribou. We concluded that the herd persisted because of this reduced predation risk and that the animals were prepared to select a reduced variety and phytomass of forage to remain for long periods in the relatively safe island habitat. When the animals did visit the mainland to feed they sclected forbs that provided a large bite size. On the island food supplies were too meager to select plants that gave a large bite size and the caribou spent long intervals feeding. Caribou by using habitats with a large phytomass and selecting for large bite size should minimize their time feeding which would allow them more time to watch for predators. [ABSTRACT FROM AUTHOR]