No abstract

At eight European field sites, the impact of loss of plant diversity on primary productivity was simulated by synthesizing grassland communities with different numbers of plant species. Results differed in detail at each location, but there was an overall log-linear reduction of average aboveground biomass with loss of species. For a given number of species, communities with fewer functional groups were less productive. These diversity effects occurred along with differences associated with species composition and geographic location. Niche complementarity and positive species interactions appear to play a role in generating diversity-productivity relationships within sites in addition to sampling from the species pool.

Aim  At a coarse scale, the treelines of the world's mountains seem to follow a common isotherm, but the evidence for this has been indirect so far. Here we aim at underpinning this with facts. Location  We present the results of a data‐logging campaign at 46 treeline sites between 68° N and 42° S. Methods  We measured root‐zone temperatures with an hourly resolution over 1–3 years per site between 1996 and 2003. Results  Disregarding taxon‐, landuse‐ or fire‐driven tree limits, high altitude climatic treelines are associated with a seasonal mean ground temperature of 6.7 °C (±0.8 SD; 2.2 K amplitude of means for different climatic zones), a surprisingly narrow range. Temperatures are higher (7–8 °C) in the temperate and Mediterranean zone treelines, and are lower in equatorial treelines (5–6 °C) and in the subarctic and boreal zone (6–7 °C). While air temperatures are higher than soil temperatures in warm periods, and are lower than soil temperatures in cold periods, daily means of air and soil temperature are almost the same at 6–7 °C, a physics driven coincidence with the global mean temperature at treeline. The length of the growing season, thermal extremes or thermal sums have no predictive value for treeline altitude on a global scale. Some Mediterranean (Fagus spp.) and temperate South Hemisphere treelines (Nothofagus spp.) and the native treeline in Hawaii (Metrosideros) are located at substantially higher isotherms and represent genus‐specific boundaries rather than boundaries of the life‐form tree. In seasonal climates, ground temperatures in winter (absolute minima) reflect local snow pack and seem uncritical. Main conclusions  The data support the hypothesis of a common thermal threshold for forest growth at high elevation, but also reflect a moderate region and substantial taxonomic influence.

In this review I ®rst compile data for the worldwide position of climate-driven alpine treelines. Causes for treeline formation are then discussed with a global perspective. Available evidence suggests a combination of a general thermal boundary for tree growth, with regionally variable``modulatory'' forces, including the presence of certain taxa. Much of the explanatory evidence found in the literature relates to these modulatory aspects at regional scales, whereas no good explanations emerged for the more fundamental global pattern related to temperature per se, on which this review is focused. I hypothesize that the life form``tree'' is limited at treeline altitudes by the potential investment, rather than production, of assimilates (growth as such, rather than photosynthesis or the carbon balance, being limited). In shoots coupled to a cold atmosphere, meristem activity is suggested to be limited for much of the time, especially at night. By reducing soil heat¯ux during the growing season the forest canopy negatively a ects root zone temperature. The lower threshold temperature for tissue growth and development appears to be higher than 3°C and lower than 10°C, possibly in the 5.5±7.5°C range, most commonly associated with seasonal means of air temperature at treeline positions. The physiological and developmental mechanisms responsible have yet to be analyzed. Root zone temperature, though largely unknown, is likely to be most critical.

Summary 1The ongoing enrichment of the atmosphere with CO 2 raises the question of whether growth of forest trees, which represent close to 90% of the global biomass carbon, is still carbon limited at current concentrations of close to 370 p.p.m. As photosynthesis of C3 plants is not CO 2 -saturated at such concentrations, enhanced 'source activity' of leaves could stimulate 'sink activity' (i.e. growth) of plants, provided other resources and developmental controls permit. I explore current levels of non-structural carbon in trees in natural forests in order to estimate the potential for a carbon-driven stimulation of growth. 2 The concentration of non-structural carbohydrates (NSC) in tree tissues is considered a measure of carbon shortage or surplus for growth. A periodic reduction of NSC pools indicates either that carbon demand exceeds con-current supply, or that both source and sink activity are low. A steady, very high NSC concentration is likely to indicate that photosynthesis fully meets, or even exeeds, that needed for growth (surplus assimilates accumulate). 3 The analysis presented here considers data for mature trees in four climatic zones: the high elevation treeline (in Mexico, the Alps and Northern Sweden), a temperate lowland forest of central Europe, Mediterranean sclerophyllous woodland and a semideciduous tropical forest in Panama. 4 In all four climatic regions, periods of reduced or zero growth show maximum Cloading of trees (source activity exceeding demand), except for dry midsummer in the Mediterranean. NSC pools are generally high throughout the year, and are not significantly affected by mass fruiting episodes. 5 It is concluded that, irrespective of the reason for its periodic cessation, growth does not seem to be limited by carbon supply. Instead, in all the cases examined, sink activity and its direct control by the environment or developmental constraints, restricts biomass production of trees under current ambient CO 2 concentrations. 6 The current carbohydrate charging of mature wild trees from the tropics to the cold limit of tree growth suggests that little (if any) leeway exists for further CO 2 -fertilization effects on growth.

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of nonstructural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad-leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring-porous versus diffuse-porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris . The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues ('fat trees'; Tilia , Pinus , Picea , Larix ), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO 2 enrichment. The classical view that deciduous trees rely more on C-reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO 2 concentrations over the last 150 years.

In most temperate tree species, phenological events such as flowering and autumnal cessation of growth are not primarily controlled by temperature.

Contents  Summary 393 A traditionally scarce resource becomes abundant 394 Photosynthesis is not saturated at current CO2 concentrations  395 The fate of extra carbon 396 Co drivers of plant growth responses to elevated CO2 397 Plant CO2 responses as a function of time 399 Plant CO2 responses per unit land area, a matter of definition  401 CO2 effects on biomass carbon stores depend on tree demography  402 Biomass responses to elevated CO2 in steady state  and expanding systems  403 Conclusions 405  Acknowledgements 406  References 406 Summary In this review I am drawing attention to some constraints and biases in CO2 enrichment experiments and the analysis of data in the literature. Conclusions drawn from experimental works differ when the data are grouped in a way such that the relative frequency of test conditions does not determine the emerging trends, for instance unrealistically strong CO2–‘fertilization’ effects, which are in conflict with some basic ecological principles. I suggest separating three test conditions: uncoupled systems (plants not depending in a natural nutrient cycle) (I); expanding systems, in which plants are given ample space and time to explore otherwise limited resources (II); and fully coupled systems in which the natural nutrient cycling governs growth at steady‐state leaf area index (LAI) and fine root renewal (III). Data for 10 type III experiments yield rather moderate effects of elevated CO2 on plant biomass production, if any. In steady‐state grassland, the effects are water‐related; in closed tree stands, initial effects decline rapidly with time. Plant–soil coupling (soil conditions) deserves far greater attention than plant–atmosphere coupling (CO2 enrichment technology).