Specific root length (SRL, m g71 ) is probably the most frequently measured morphological parameter of fine roots. It is believed to characterize economic aspects of the root system and to be indicative of environmental changes. The main objectives of this paper were to review and summarize the published SRL data for different tree species throughout Europe and to assess SRL under varying environmental conditions. Meta-analysis was used to summarize the response of SRL to the following manipulated environmental conditions: fertilization, irrigation, elevated temperature, elevated CO 2 , Al-stress, reduced light, heavy metal stress and physical disturbance of soil. SRL was found to be strongly dependent on the fine root classes, i.e. on the ectomycorrhizal short roots (ECM), and on the roots 50.5 mm, 51 mm, 52 mm and 1 -2 mm in diameter SRL was largest for ECM and decreased with increasing diameter. Changes in soil factors influenced most strongly the SRL of ECM and roots 50.5 mm. The variation in the SRL components, root diameter and root tissue density, and their impact on the SRL value were computed. Meta-analyses showed that SRL decreased significantly under fertilization and Al-stress; it responded negatively to reduced light, elevated temperature and CO 2. We suggest that SRL can be used successfully as an indicator of nutrient availability to trees in experimental conditions.

Fine roots (52 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m 72, and that of spruce and pine 297 g m 72 and 277 g m 72 , respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.

Development of direct and indirect methods for measuring root turnover and the status of knowledge on fine root turnover in forest ecosystems are discussed. While soil and ingrowth cores give estimates of standing root biomass and relative growth, respectively, minirhizotrons provide estimates of median root longevity (turnover time) i.e., the time by which 50% of the roots are dead. Advanced minirhizotron and carbon tracer studies combined with demographic statistical methods and new models hold the promise of improving our fundamental understanding of the factors controlling root turnover. Using minirhizotron data, fine root turnover (y )1 ) can be estimated in two ways: as the ratio of annual root length production to average live root length observed and as the inverse of median root longevity. Fine root production and mortality can be estimated by combining data from minirhizotrons and soil cores, provided that these data are based on roots of the same diameter class (e.g., <1 mm in diameter) and changes in the same time steps. Fluxes of carbon and nutrients via fine root mortality can then be estimated by multiplying the amount of carbon and nutrients in fine root biomass by fine root turnover. It is suggested that the minirhizotron method is suitable for estimating median fine root longevity. In comparison to the minirhizotron method, the radio carbon technique favor larger fine roots that are less dynamics. We need to reconcile and improve both methods to develop a more complete understanding of root turnover.

Applications and limitations of the minirhizotron technique (non-destructive) in relation to two frequently used destructive methods (soil coreing and ingrowth cores) is discussed. Sequential coreing provides data on standing crop but it is difficult to obtain data on root biomass production. Ingrowth cores can provide a quick estimate of relative fine-root growth when root growth is rapid. One limitation of the ingrowth core is that no information on the time of ingrowth and mortality is obtained.The minirhizotron method, in contrast to the destructive methods permits simultaneous calculation of fine-root length production and mortality and turnover. The same fine-root segment in the same soil space can be monitored for its life time, and stored in a database for processing. The methodological difficulties of separating excavated fine roots into living and dead vitality classes are avoided, since it is possible to judge directly the successive ageing of individual roots from the images. It is concluded that the minirhizotron technique is capable of quantifying root dynamics (root-length production, mortality and longevity) and fine-root decomposition. Additionally, by combining soil core data (biomass, root length and nutrient content) and minirhizotron data (length production and mortality), biomass production and nutrient input into the soil via root mortality and decomposition can be estimated.

The effects of soil warming and nitrogen availability on root production, longevity and mortality were studied using minirhizotrons in irrigation (C), fertilized (F), heated (H), and heated-fertilized (HF) plots in a Norway spruce stand in northern Sweden from October 1996 to October 1997. Irrigation was included in all treatment plots. Heating cables were used to maintain the soil temperature in heated plots at 5 1C above that in unheated plots during the growing season. A Kaplan-Meier approach was used to estimate the longevity of fine roots and Cox proportional hazards regression to analyze the effects of the H, F, and HF treatments on the risk of root mortality.The proportion of annual root length production contributed by winter-spring production amounted to 52% and 49% in heated plots and heated-fertilized plots, respectively. The annual root length production in C plots was significantly higher than in other treatments, while the HF treatment gave significantly greater production compared with the F treatment. The risk of mortality (hazard ratio) relative to C plots was higher in H plots (358%) and F plots (191%). The interaction between heating and fertilizing was strongly significant. The increase in the risk of root mortality in combined fertilization and heating (103%) was lower than that in the H or F plots.The results show that nitrogen addition combined with warmer temperatures decreases the risk of root mortality, and fine root production is a function of the length of the growing season. In the future, fertilization combined with the warmer temperatures expected to follow predicted climatic change may increase root production in boreal forests at low fertility sites.