CO2 balance of boreal, temperate, and tropical forests derived from a global database

Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome‐specific carbon budgets; to re‐examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO₂ balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 °C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO₂ balance required the introduction of substantial biome‐specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non‐CO₂ carbon fluxes are not presently being adequately accounted for.     

Contemporary carbon accumulation in a boreal oligotrophic minerogenic mire – a significant sink after accounting for all C-fluxes

Based on theories of mire development and responses to a changing climate, the current role of mires as a net carbon sink has been questioned. A rigorous evaluation of the current net C-exchange in mires requires measurements of all relevant fluxes. Estimates of annual total carbon budgets in mires are still very limited. Here, we present a full carbon budget over 2 years for a boreal minerogenic oligotrophic mire in northern Sweden (64°11′N, 19°33′E). Data on the following fluxes were collected: land–atmosphere CO₂ exchange (continuous Eddy covariance measurements) and CH₄ exchange (static chambers during the snow free period); TOC (total organic carbon) in precipitation; loss of TOC, dissolved inorganic carbon (DIC) and CH₄ through stream water runoff (continuous discharge measurements and regular C-concentration measurements). The mire constituted a net sink of 27±3.4 (±SD) g C m⁻² yr⁻¹ during 2004 and 20±3.4 g C m⁻² yr⁻¹ during 2005. This could be partitioned into an annual surface–atmosphere CO₂ net uptake of 55±1.9 g C m⁻² yr⁻¹ during 2004 and 48±1.6 g C m⁻² yr⁻¹ during 2005. The annual NEE was further separated into a net uptake season, with an uptake of 92 g C m⁻² yr⁻¹ during 2004 and 86 g C m⁻² yr⁻¹ during 2005, and a net loss season with a loss of 37 g C m⁻² yr⁻¹ during 2004 and 38 g C m⁻² yr⁻¹ during 2005. Of the annual net CO₂-C uptake, 37% and 31% was lost through runoff (with runoff TOC>DIC≫CH₄) and 16% and 29% through methane emission during 2004 and 2005, respectively. This mire is still a significant C-sink, with carbon accumulation rates comparable to the long-term Holocene C-accumulation, and higher than the C-accumulation during the late Holocene in the region.     

Non-invasive determination of plant biomass with microwave resonators

Non-invasive and rapid determination of plant biomass would be beneficial for a number of research aims. Here, we present a novel device to non-invasively determine plant water content as a proxy for plant biomass. It is based on changes of dielectric properties inside a microwave cavity resonator induced by inserted plant material. The water content of inserted shoots leads to a discrete shift in the centre frequency of the resonator. Calibration measurements with pure water showed good spatial homogeneity in the detection volume of the microwave resonators and clear correlations between water content and centre frequency shift. For cut tomato and tobacco shoots, linear correlations between fresh weight and centre frequency shift were established. These correlations were used to continuously monitor diel growth patterns of intact plants and to determine biomass increase over several days. Interferences from soil and root water were excluded by shielding pots with copper. The presented proof of principle shows that microwave resonators are promising tools to quantitatively detect the water content of plants and to determine plant biomass. As the method is non-invasive, integrative and fast, it provides the opportunity for detailed, dynamic analyses of plant growth, water status and phenotype.     

Storms can cause Europe-wide reduction in forest carbon sink

Disturbance of ecosystems is a major factor in regional carbon budgets, and it is believed to be partly responsible for the large inter-annual variability of the terrestrial part of the carbon balance. Forest fires have so far been considered as the most important disturbance but also other forms of disturbance such as insect outbreaks or wind-throw might contribute significantly to the largely unexplained inter-annual variability, at least in specific regions. The effect of wind-throw has not yet been estimated because of lack of data on how carbon fluxes are affected. The Gudrun storm, which hit Sweden in January 2005, resulted in ca. 66 million m³ of wind-thrown stem wood on an area of ca. 272 000 ha. Using a model (BIOME-BGC) calibrated to CO₂ flux measurements at two sites, the annual net ecosystem productivity during the first year after the storm was estimated to be in the range −897 to −1259 g C m⁻² yr⁻¹. This is a much higher loss compared with harvested (clear-cut) forests in Europe, which ranged between ca. −420 and −100 g m⁻² yr⁻¹. The reduction in the carbon sink scaled to the whole wind-thrown area was estimated at ca. 3 million tons C during the first year. By historical data on wind-throw in Europe combined with modelling, we estimated that the large Lothar storm in 1999 reduced the European carbon balance by ca. 16 million tons C, this is ca. 30% of the net biome production in Europe. We conclude that the impact of increased forest damage by more frequent storms in future climate change scenarios must be considered and that intermittent large wind-throw events may explain a part of the large inter-annual variability in the terrestrial carbon sink.     

Root cooling strongly affects diel leaf growth dynamics,water and carbohydrate relations in Ricinus communis

In laboratory and greenhouse experiments with potted plants, shoots and roots are exposed to temperature regimes throughout a 24 h (diel) cycle that can differ strongly from the regime under which these plants have evolved. In the field, roots are often exposed to lower temperatures than shoots. When the root‐zone temperature in Ricinus communis was decreased below a threshold value, leaf growth occurred preferentially at night and was strongly inhibited during the day. Overall, leaf expansion, shoot biomass growth, root elongation and ramification decreased rapidly, carbon fluxes from shoot to root were diminished and carbohydrate contents of both root and shoot increased. Further, transpiration rate was not affected, yet hydrostatic tensions in shoot xylem increased. When root temperature was increased again, xylem tension reduced, leaf growth recovered rapidly, carbon fluxes from shoot to root increased, and carbohydrate pools were depleted. We hypothesize that the decreased uptake of water in cool roots diminishes the growth potential of the entire plant – especially diurnally, when the growing leaf loses water via transpiration. As a consequence, leaf growth and metabolite concentrations can vary enormously, depending on root‐zone temperature and its heterogeneity inside pots.     

Net primary production and light use efficiency in a mixed coniferous forest in Sweden

Simple light use efficiency (ɛ) models of net primary production (NPP) have recently been given great attention (NPP = ɛ × absorbed photosynthetically active radiation). The underlying relationships have, however, not been much studied on a time step less than a month. In this study daily NPP was estimated as the sum of net ecosystem exchange (NEE) and heterotrophic respiration (R_h) of a mixed pine and spruce forest in Sweden. NEE was measured by eddy correlation technique and R_h was estimated from measurements of forest floor respiration (R_f) and the root share of R_f. The total yearly NPP was on average 810 g C m⁻² year⁻¹ for 3 years and yearly ɛ was between 0.58 and 0.71 g C MJ⁻¹, which is high in comparison with other studies. There was a seasonal trend in ɛ with a relatively constant level of approximately 0.90 g C MJ⁻¹ from April to September Daily NPP did not increase for daily intercepted radiation above 6 MJ m⁻² d⁻¹, indicating that between-years variation in NPP is not directly dependent on total Q_i. The light was most efficiently used at an average daytime temperature of around 15 °C. At daytime vapour pressure deficit above 1400 Pa ɛ was reduced by approximately 50%.     

Leaf growth dynamics of two congener gymnosperm tree species reflect the heterogeneity of light intensities given in their natural ecological niche

Chamaecyparis obtusa var. formosana and Chamaecyparis formosensis are congener gymnosperm tree species native to Taiwan cloud forests; occupying different niches there. While the seedlings of C. formosensis occur predominantly under bright conditions in large forest gaps, seedlings of C. obtusa var. formosana are mainly found below the canopy of mature forests or in small gaps. It is well known that congener species occupying different niches typically differ in several ecophysiological and morphological traits, but the differences in growth dynamics of such species are still totally unclear, as the diurnal growth dynamics of gymnosperm leaves have not been investigated before. Modern methods of digital image sequence processing were used in this study to analyse the leaf growth dynamics of the two species. We found that both species show similar base–tip gradients and pronounced diurnal growth rhythms with maxima in the evening. Differences between the two species concerning their growth dynamics correlated closely with their ecological amplitudes and abundances. Chamaecyparis obtusa var. formosana grew faster than C. formosensis in low light intensity, typical for closed-canopy situations, and reacted quickly by increasing or decreasing growth rate when light intensity changed within a range typically found below small canopy gaps. In contrast to this, C. formosensis grew better in light intensities typical for open vegetation situations, but reacted slower towards changes of light intensity. Based on those results, the hypothesis can be developed that fluctuations of leaf growth dynamics reflect heterogeneities of the light environment within the niche occupied by a given species.     

Herbivore-induced jasmonic acid bursts in leaves of Nicotiana attenuata mediate short-term reductions in root growth

Root growth in Nicotiana attenuata is transiently reduced after application of oral secretions (OS) of Manduca sexta larvae to wounds in leaves. Feeding of M. sexta or OS elicitation is known to result in jasmonic acid (JA) and ethylene bursts, and activates a suite of defence responses. Because both plant hormones are known to strongly reduce root growth, their activation might account for the observed reduction of root growth following herbivory. To test this hypothesis, we measured primary root growth with digital image sequence processing at high temporal resolution in antisense- lipoxygenase 3 (as LOX3 ) and inverted repeat- coronatin-insensitive 1 (ir COI1 ) seedlings which are impaired in JA biosynthesis and perception, respectively, and wild-type (WT) seedlings. Higher root growth rates in ir COI1 compared with WT were observed after OS elicitation. The dynamics of wound-induced root growth reduction coincide with the dynamics of root growth reduction induced by external application of methyl JA. In an experiment with 1-methylcyclopropen (1-MCP), a potent ethylene receptor blocker, no wounding-specific difference between growth of 1-MCP-treated plants and non-treated plants was observed, suggesting that wound-induced endogenous JA and not ethylene mediates the wounding-specific reduction in root growth. Yet, inhibiting the ethylene response by applying 1-MCP led to markedly increased root growth compared with that of control plants, indicating that ethylene normally suppresses plant growth in N. attenuata seedlings.     

Jasmonic acid does not mediate root growth responses to wounding in Arabidopsis thaliana

Jasmonic acid (JA) is a crucial plant defence signalling substance that has recently been shown to mediate herbivory-induced root growth reduction in the ecological model species Nicotiana attenuata . To clarify whether JA-induced reduction of root growth might be a general response increasing plant fitness under biotic stress, a suite of experiments was performed with the model plant Arabidopsis thaliana . JA bursts were elicited in leaves of A. thaliana in different ways. Root growth reduction was neither induced by foliar application of herbivore oral secretions nor by direct application of methyl jasmonate to leaves. Root growth reduction was observed when leaves were infected with the pathogen Pseudomonas syringae pv. tomato, which persistently induces the JA signalling pathway. Yet, high resolution growth analyses of this effect in wild type and JA biosynthesis knock-out mutants showed that it was elicited by the bacterial toxin coronatine that suggests ethylene- but not JA-induced root growth reduction in A. thaliana . Overall, the results demonstrate that the reaction of root growth to herbivore-induced JA signalling differs among species, which is discussed in the context of different ecological defence strategies among species.