Waldemar Ziegler

Thirst beats hunger - declining hydration during drought prevents carbon starvation in Norway spruce saplings.

Drought-induced tree mortality results from an interaction of several mechanisms. Plant water and carbon relations are interdependent and assessments of their individual contributions are difficult. Because drought always affects both plant hydration and carbon assimilation, it is challenging to disentangle their concomitant effects on carbon balance and carbon translocation. Here, we report results of a manipulation experiment specifically designed to separate drought effects on carbon and water relations from those on carbon translocation. In a glasshouse experiment, we manipulated the carbon balance of Norway spruce saplings exposed to either drought or carbon starvation (CO2 withdrawal), or both treatments, and compared the dynamics of carbon exchange, allocation and storage in different tissues. Drought killed trees much faster than did carbon starvation. Storage C pools were not depleted at death for droughted trees as they were for starved, well-watered trees. Hence drought has a significant detrimental effect on a plants ability to utilize stored carbon. Unless they can be transported to where they are needed, sufficient carbon reserves alone will not assure survival of a drought except under specific conditions, such as moderate drought, or in species that maintain plant water relations required for carbon re-mobilization.

Lethal drought leads to reduction in nonstructural carbohydrates in Norway spruce tree roots but not in the canopy

Summary Heat waves and droughts are expected to increase in frequency and severity in many regions with future climate change, threatening the survival of a number of forest ecosystems. However, our understanding of the physiological processes and mechanisms underlying drought-induced tree mortality is incomplete. Here, we present results on the physiological response of young Norway spruce trees exposed to lethal drought stress. We applied three levels of drought treatment (control, drying–rewetting, complete drought) and monitored relevant physiological functions and processes of carbon and water relations at high temporal resolution until tree death occurred. Only trees subjected to continuous drought died in our experiment. Trees subjected to drying–rewetting cycles consistently recovered in their ability to transport water, indicating that these trees do not suffer permanent damage to the hydraulic system. In all cases, drought reduced carbon assimilation, caused changes in carbon allocation and appeared to have severely reduced phloem functioning and carbon translocation. Structural growth was sacrificed for carbon investment in maintenance respiration and osmoprotection. Severe drought caused trees to rely on stored carbon reserves but, in contrast to above-ground tissues, only root carbon pools were strongly reduced when trees died. Our results indicate that drought-induced changes in carbon allocation, use and transport differ between above- and below-ground tissues in trees. While root death may have been caused by carbon depletion, this was definitely not the case in above-ground tissues. Our findings indicate that mortality mechanisms are not defined at the organism level but rather within tree compartments.

Fire and site type effects on the long-term carbon and nitrogen balance in pristine Siberian Scots pine forests

Effects of fire and site type on carbon (C) and nitrogen (N) balances were determined by following the change of total and component C and N pools along four chronosequences of fire-prone Siberian Scots pine ecosystems. These differed in the mean return interval of surface fires (unburned – moderately burned, 40 years – heavily burned, 25 years) and site quality (lichen versus Vaccinium site type). Of the Vaccinium site type (higher site quality) only a moderately burned chronosequence was studied. A total of 22 even-aged stands were investigated with stand ages ranging from 2 to 383 years. The C balance was dominated by the opposing dynamics of coarse woody debris (CWD) and biomass and could be divided into three phases: (1) Young stands (up to 40 years)acted as a net source for C of 6-10 mol C m-2 year-1 because the previous generation CWD pool originating from stand-replacing crown fires decayed much faster than biomass increased. During this period the C pool in the unburned lichen type chronosequence decreased from 807 to 480 mol C m-2. (2) Middle aged stands (40-100 years) being in a stage of maximum biomass accumulation were a net sink of 8-10 mol C m-2 year-1. (3)Maturestands (100 to > 350 years) continued to sequester C at a lower rate (0.8-2.5mol C m-2 year-1). Differences in the rates of C sequestration during the two later phases could be explained by the complex interaction between surface fire regime and site type. Recurrent surface fires resulted in enhanced mortality and regularly redistributed C from the living to the CWD pool thereby lowering the rate of C sequestration. Site quality determined the potential to recover from disturbance by fire events. Differences in site type did not correlate with soil and total ecosystem N pool size. However, the N status of needles as well as the N pool of physiologically active tissue was highest in the stands of the Vaccinium type. The “woody” C pool (biomass + CWD) was sensitive to differences in surface fire regime and site type. It was lowest in the heavily burned lichen type chronosequence (297 ± 108 mol C m-2), intermediate in the unburned and moderately burned lichen type chronosequence (571 ± 179 mol C m-2) and highest in the moderately burned Vaccinium type chronosequence (810 ± 334 mol C m-2). In contrast, the total soil C pool (organic plus mineral layer down to a depth of 25 cm) was independent of stand age, surface fire regimeand site type and fluctuated around a value of 250 mol C m-2. The organic layer C pool oscillated in response to recurring surface fires and its C pool was dependent on time since fire increasing at a rate of about 1.5 mol C m-2 year-during the first 40 years and then reaching a plateau of 170 mol C m-2. The total ecosystem N pool was 7.4 ± 1.5 mol N m-2 on average of which only 25 % were stored in biomass or coarse woody debris. Total ecosystem N was independent of stand age, surface fire regime and site type. No correlation was found between total ecosystem C and N pools. Average total ecosystem C:N ratio was 114 ± 35 mol C mol N-1. A conceptual model illustrating how changes in the regime of stand-replacing crown fires and recurrent surface fires and changes in site quality interact in determining the long-term C balance in Siberian Scots pine forests is presented.

Advection and resulting CO2 exchange uncertainty in a tall forest in central Germany.

Potential losses by advection were estimated at Hainich Forest, Thuringia, Germany, where the tower is located at a gentle slope. Three approaches were used: (1) comparing nighttime eddy covariance fluxes to an independent value of total ecosystem respiration by bottom-up modeling of the underlying processes, (2) direct measurements of a horizontal CO2 gradient and horizontal wind speed at 2 m height in order to calculate horizontal advection, and (3) direct measurements of a vertical CO2 gradient and a three-dimensional wind profile in order to calculate vertical advection. In the first approach, nighttime eddy covariance measurements were compared to independent values of total ecosystem respiration by means of bottom-up modeling of the underlying biological processes. Turbulent fluxes and storage term were normalized to the fluxes calculated by the bottom-up model. Below a u(*) threshold of 0.6 m/s the normalized turbulent fluxes decreased with decreasing u(*), but the flux to the storage increased only up to values less than 20% of the modeled flux at low turbulence. Horizontal advection was measured by a horizontal CO2 gradient over a distance of 130 m combined with horizontal wind speed measurements. Horizontal advection occurred at most of the evenings independently of friction velocity above the canopy. Nevertheless, horizontal advection was higher when u(*) was low. The peaks of horizontal advection correlated with changes in temperature. A full mass balance including turbulent fluxes, storage, and horizontal and vertical advection resulted in an increase of spikes and scatter but seemed to generally improve the results from the flux measurements. The comparison of flux data with independent bottom-up modeling results as well as the direct measurements resulted in strong indications that katabatic flows along the hill slope during evening and night reduces the measured apparent ecosystem respiration rate. In addition, anabatic flows may occur during the morning. We conclude that direct measurements of horizontal and vertical advection are highly necessary at sites located even on gentle hill slopes.

High temperature causes negative whole-plant carbon balance under mild drought

Theoretically, progressive drought can force trees into negative carbon (C) balance by reducing stomatal conductance to prevent water loss, which also decreases C assimilation. At higher temperatures, negative C balance should be initiated at higher soil moisture because of increased respiratory demand and earlier stomatal closure. Few data are available on how these theoretical relationships integrate over the whole plant. We exposed Thuja occidentalis to progressive drought under three temperature conditions (15, 25, and 35°C), and measured C and water fluxes using a whole-tree chamber design. High transpiration rates at higher temperatures led to a rapid decline in soil moisture. During the progressive drought, soil moisture-driven changes in photosynthesis had a greater impact on the whole-plant C balance than respiration. The soil moisture content at which whole-plant C balance became negative increased with temperature, mainly as a result of higher respiration rates and an earlier onset of stomatal closure under a warmer condition. Our results suggest that the effect of drought on whole-plant C balance is highly temperature-dependent. High temperature causes a negative C balance even under mild drought and may increase the risk of C starvation.

A new mass conservation approach to the study of CO2 advection in an alpine forest

A new method is proposed for the computation of CO2 Net Ecosystem Exchange (NEE) and its components in a forest ecosystem. Advective flux is estimated by taking into account the air mass conservation principle. For this purpose, wind and dry air density values on the surface of the control volume are first corrected and then the advective flux is estimated on the surface of the control volume. Turbulent flux is also computed along the surface of the control volume while storage flux is computed inside the volume. Additional characteristics of this method are that incompressibility of the mean flow is not assumed a priori, and that vertical and horizontal advective fluxes are not treated separately, but their sum is estimated directly. The methodology is applied to experimental data collected with a three-dimensional scheme at the alpine site of Renon during the Advex project (July 2005). The advection flux was found to be prevailing positive at night and negative during the day, as was found in previous studies on advection for the same site, but showed a lower scatter in half-hour calculated values. We tested the effect of its summation on turbulent and storage fluxes to produce half-hourly values of NEE. Nighttime NEE values were used in functional relations with soil temperature, daytime values with PPFD. The effect of addition of the advection component was an increase in the values of parameters indicating ecosystem respiration, quantum yield, and photosynthetic capacity. The coefficient of correlation between NEE and environmental drivers increased. (Less)

Adjustment to the light environment in small-statured forbs as a strategy for complementary resource use in mixtures of grassland species

BACKGROUND AND AIMS The biological mechanisms of niche complementarity allowing for a stable coexistence of a large number of species in a plant community are still poorly understood. This study investigated how small-statured forbs use environmental niches in light and CO(2) to explain their persistence in diverse temperate grasslands. METHODS Light and CO(2) profiles and the corresponding leaf characteristics of seven small-statured forbs were measured in monocultures and a multi-species mixture within a biodiversity experiment (Jena Experiment) to assess their adjustment to growth conditions in the canopy. KEY RESULTS Environmental conditions near the ground varied throughout the season with a substantial CO(2) enrichment (>70 µmol mol(-1) at 2 cm, >20 µmol mol(-1) at 10 cm above soil surface) and a decrease in light transmittance (to <5 % deep in the canopy) with large standing biomass (>500 g d. wt m(-2)) in the multi-species assemblage. Leaf morphology, biochemistry and physiology of small-statured forbs adjusted to low light in the mixture compared with the monocultures. However, the net carbon assimilation balance during the period of low light only compensated the costs of maintenance respiration, while CO(2) enrichment near the ground did not allow for additional carbon gain. Close correlations of leaf mass per area with changes in light availability suggested that small-statured forbs are capable of adjusting to exploit seasonal niches with better light supply for growth and to maintain the carbon metabolism for survival if light transmittance is substantially reduced in multi-species assemblages. CONCLUSIONS This study shows that adjustment to a highly dynamic light environment is most important for spatial and seasonal niche separation of small-statured forb species in regularly mown, species-rich grasslands. The utilization of short-period CO(2) enrichment developing in dense vegetation close to the ground hardly improves their carbon balance and contributes little to species segregation along environmental niche axes.