Root distribution of under-planted European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) below the canopy of a mature Norway spruce stand as a function of light

Aboveground and belowground biomass of 15-year-old under-planted European beech seedlings (Fagus sylvatica L.) in Norway spruce stand were studied along a light gradient in three plots, in the northern part of Slovenia. Differences in soil water content, aboveground and fine root biomass distribution were confirmed between studied plots. Light had significant effect on the total biomass, root-shoot ratio (0.388 ± 0.076 under canopy, 0.549 ± 0.042 in the edge, 0.656 ± 0.047 in the open), specific root length (SRL) of fine beech roots (561.9 ± 42.2 under canopy, 664.3 ± 51.2 in the edge, 618.2 ± 72.8 in the open) and specific leaf area in beech, indicating morphological adjustment to shade. However, SRL of beech fine roots indicated no change between plots. The correlation between total aboveground and root biomass and light below the mature stand canopy was higher in the case of diffuse light intensity. Most fine roots of spruce were concentrated in the top (0–20 cm) soil layer. Beech fine roots under canopy and edge conditions were also concentrated in top (0–20 cm) soil layer and exhibited shift downwards to deeper soil horizons in open plot. Root proportion between beech and spruce changed with light toward beech with increasing light intensity for both fine and coarse roots.     

Impact factors on fine root seasonal dynamics in Fraxinus mandshurica plantations

Fine root turnover plays important roles in carbon allocation and nutrient cycling in forest ecosystems. Seasonal dynamics of fine roots is critical for understanding the processes of fine root turnover. From May to October 2002, soil core method was used for estimating the seasonal pattern of fine root (diameter < 1 mm) parameters (biomass, specific root length (SRL) and root length density (RLD)) in a Manchurian ash (Fraxinus mandshurica) plantation located at the Maoershan Experiment Station, Heilongjiang Province, northeast of China. The relationships of fine root biomass, SRL and RLD with available nitrogen in soil, average soil temperature per month in 10 cm depth and soil moisture content were analyzed. Seasonal variation of fine root biomass was significant (P < 0.05). The peak values of fine root biomass were observed both in spring and in autumn, but SRL and RLD were the highest in spring and lowest in autumn. Specific root length and root length density were higher in spring and summer, which means that fine root diameter was thinner. In autumn, both parameters decreased significantly due to secondary incrassation of fine root diameter or the increase of tissue density. Seasonal dynamics of fine roots was associated with available nitrogen in soil, soil temperature in 10 cm depth and moisture content. Fine root biomass has a significant relationship with available NH₄ ⁺-N in soil. Available NO₃ ⁻-N in soil, soil temperature in 10-cm depth and moisture content have a positive correlation with fine root biomass, SRL and RLD, although these correlations are not significant (P > 0.05). But the compound effects of soil available N, soil temperature and soil moisture content are significant to every root parameter. The variations of these three root parameters in different seasons show different physiological and ecological functions in different growing periods. Translated from Scientia Silvae Sinicae, 2006, 42(9): 7–12 [译自: 林业科学]     

Estimating the production and mortality of fine roots in a Japanese cedar (Cryptomeria japonica D. Don) plantation using a minirhizotron technique

Fine roots are a key component of forested ecosystems, but available information is still limited. This study examined the production and mortality of fine roots less than 1 mm in diameter in a Japanese cedar (Cryptomeria japonica D. Don) plantation located on the Kanto Plain in central Japan. We used a minirhizotron technique in combination with soil coring, and collected data for 1 year (May 2002–May 2003). Fine root production and mortality were determined from changes in the lengths of individual fine roots on minirhizotron tubes. Both fine root production and mortality rates were greater in the upper soil than in lower soil levels. Both rates were seasonal, with higher values in summer than in winter; this trend was more pronounced in upper soil levels. These results suggest that environmental conditions, such as temperature or soil properties, affect the production and mortality rates of fine roots. Fine root production and mortality occurred simultaneously, and their rates were similar, which may have led to unclear seasonal changes in fine root standing crop estimates. Soil coring indicated that the fine root biomass of this stand was about 120 g m⁻², of which 40% was from Japanese cedar. The estimated rates of dry matter production and mortality of total fine roots, including understory plants, were both approximately 300 g m⁻² year⁻¹.     

Root distribution of Fagus sylvatica in a chronosequence in western France

The distribution of fine (<2 mm diameter) and small roots (2–20 mm diameter) was investigated in a chronosequence consisting of 9-year-old, 26-year-old, 82-year-old and 146-year-old European beech (Fagus sylvatica) stands. A combination of trench wall observations and destructive root sampling was used to establish whether root distribution and total biomass of fine and small roots varied with stand age. Root density decreased with soil depth in all stands, and variability appeared to be highest in subsoil horizons, especially where compacted soil layers occurred. Roots clustered in patches in the top 0–50 cm of the soil or were present as root channels at greater depths. Cluster number, cluster size and number of root channels were comparable in all stands, and high values of soil exploitation occurred throughout the entire chronosequence. Overall fine root biomass at depths of 0–120 cm ranged from 7.4 Mg ha⁻¹ to 9.8 Mg ha⁻¹, being highest in the two youngest stands. Small root biomass ranged from 3.6 Mg ha⁻¹ to 13.3 Mg ha⁻¹. Use of trench wall observations combined with destructive root samples reduced the variability of these estimates. These records showed that variability in fine root distribution depended more on soil depth and edaphic conditions than on stand age, and suggest that trench wall studies provide a useful tool to improve estimates of fine root biomass.     

Fine-root dynamics of coffee in association with two shade trees in Costa Rica

Fine-root dynamics (diameter < 2.0 mm) were studied on-farm in associations of Coffea arabica with Eucalyptus deglupta or Terminalia ivorensis and in a pseudo-chronosequence of C. arabica-E. deglupta associations (two, three, four and five years old). Coffee plants were submitted to two fertilisation types. Cores were taken in the 0–40 cm soil profile two years after out-planting and subsequently in the following year in depth layers 0–10 and 10–20 cm, during and at the end of the rainy season, and during the dry season. Fine root density of coffee and timber shade trees was greater in the coffee fertilisation strip as compared to unfertilised areas close to the plants or in the inter-rows. Coffee fine roots were more evenly distributed in the topsoil (0–20 cm) whereas tree fine roots were mostly found in the first 10 cm. Although the two tree species had approximately the same fine root length density, lower coffee / tree fine root length density ratios in T. ivorensis suggest that this shade tree is potentially a stronger competitor with coffee than E. deglupta. Coffee and tree fine root length density for 0–10 cm measured during the rainy season increased progressively from two to five-year-aged associations and coffee fine root length density increased relatively more than E. deglupta fine root length density in the four and five-year-aged plantations suggesting that contrary to expectations, coffee fine roots were displacing tree fine roots.     

Long‐term effects of liming on the fine‐root standing crop of Picea abies and Pinus sylvestris in relation to chemical changes in the soil

The long‐term effects of lime application on fine roots of Norway spruce, Picea abies (L.) Karst, and Scots pine, Pinus sylvestris (L.), have been studied in five experimental forest stands subjected to different lime applications 5 to 18 years before the present study was undertaken. The effects of liming does not seem to significantly influence fine‐root development in forest stands in the long term. The only response to liming in measured root variables was a tendency to increased specific root length (SRL = fine‐root length/fine‐root dry weight, m/g). A correlation between increased SRL, decreased root biomass and increased stem volume growth was indicated. Changes in water extractable amounts of mineral elements—P, K, Ca, Mg, Mn, S, Al and Fe‐in bulk soil and rhizosphere soil from the mineral soil layers were studied in a control area and an area treated with 3830 kg CaCO₃ ha^‐1. Few significant differences were found between treatments, and then mainly in the case of Ca.     

Root development across a chronosequence in a Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We investigated the biomass, vertical distribution, and specific root length (SRL) of fine and small roots in a chronosequence of Japanese cedar (Cryptomeria japonica D. Don) plantations in Nara Prefecture, central Japan. Roots were collected from soil blocks up to 50 cm in depth in five plantations of differing age: 4, 15, 30, 41, and 88 years old. Fine-root biomass reached a maximum (639 g m₋₂) in the 15-year-old stand before canopy closure, decreased in the 30-year-old stand (422 g m₋₂), and thereafter was stable. Except in the 30-year-old stand, fine-root biomass increased in deeper soil layers as stand age increased, and the depth at which the cumulative biomass of fine roots reached 90% exhibited a good allometric relationship with mean stem diameter. Both root-length density (root length per unit soil volume) and SRL decreased with soil depth in all stands, indicating that plants mainly acquire water and nutrients from shallow soils. The highest SRL was observed in the 4-year-old stand, but the relationship between SRL and stand age was unclear in older stands. The SRL in surface soils seemed to decrease with increases in root-length density, suggesting that branching of the fine-root system during development is related to density-dependent processes rather than age.     

Root distribution of Senna siamea grown on a series of derived-savanna-zone soils in Togo, West Africa

Although crucial for assessing the functioning of alley cropping systems, quantitative information related to the hedgerow tree root distribution remains scarce. Soil mapping and destructive soil sampling was used to assess the impact of soil profile features on selected root characteristics of Senna siamea hedgerows, growing in alley cropping systems in three sites (Glidji, Amoutchou, and Sarakawa) representative for the derived savanna of Togo, West Africa. While the soil profiles in Glidji and Sarakawa contained a clay accumulation horizon, the Amoutchou profile was sandy up to 1 m. The number of small roots (diameter < 2 mm), quantified on a soil profile wall, decreased with depth in all sites. For most soil depths, the abundance of small roots tended to be higher near the tree base, e.g., ranging from 5.3 dm⁻² in Amoutchou to 21.4 dm⁻² in Glidji for the 0–20 cm layer, than in the middle of the alley, e.g., ranging from 3.1 dm⁻² in Amoutchou to 13.8 dm⁻² in Glidji for the 0–20 cm layer. Root length density (RLD) of the 0–10 cm and 10–20 cm layers was significantly higher in Glidji than in Amoutchou (P < 0.05) and in Sarakawa (P = 0.08). Differences in RLD between sites were not significant for layers below 30 cm. For each layer, root weight densities (RWD) were similar in all sites, e.g., ranging from 0.44 mg cm⁻³ in Amoutchou to 0.64 mg cm⁻³ in Glidji in the 0–10 cm layer, indicating that the roots in the Glidji topsoil had a smaller overall diameter than in Amoutchou. In Amoutchou, the relative RLD was lower than in Glidji or Sarakawa for the top 40 cm of soil, while the inverse was observed for the layers between 50 and 100 cm deep and this was related to the sandy soil profile in Amoutchou. Another consequence of the sandy profile was the larger tap root diameter below 50 cm in Amoutchou compared to Sarakawa. For all sites, significant (P < 0.001) linear regressions were observedbetween RLD's, RWD's, and the abundance of small roots, although the variation explained by the regression equations was highest for the relationship between RLD and RWD. The potential of the hedgerows to recover nutrients leached beyond the reach of food crops or the safety-net efficiency was evaluated for the tree sites. This revised version was published online in June 2006 with corrections to the Cover Date.     

The vertical distribution of fine roots of five tree species and maize in Morogoro,Tanzania

In order to assess the possibility of root competition in agroforestry, the vertical distribution of fine roots (< 2 mm in diameter) of five tree species in pure two-year-old stands was compared to that of mature maize.Cassia siamea, Eucalyptus tereticornis, Leucaena leucocephala andProsopis chilensis had a rooting pattern similar to that of maize, i.e. a slow decline in fine root mass from 0–100 cm soil depth.Eucalyptus camaldulensis had its roots evenly distributed down to 100 cm. On an average, the fine root biomass of the tree species was roughly twice as that of the maize. We conclude that the studied tree species are likely to compete with maize and other crops with a similar rooting pattern for nutrients and water.     

Effects of the extreme drought in 2003 on soil respiration in a mixed forest

We present a field study on the drought effects on total soil respiration (SR_t) and its components, i.e., “autotrophic” (SR_a: by roots/mycorrhizosphere) and “heterotrophic” respiration (SR_h: by microorganisms and soil fauna in bulk soil), in a mature European beech/Norway spruce forest. SR_a and SR_h were distinguished underneath groups of beech and spruce trees using the root exclusion method. Seasonal courses of SR_a and SR_h were studied from 2002 to 2004, with the summer of 2003 being extraordinarily warm and dry in Central Europe. We (1) analyzed the soil temperature (T _s) and moisture sensitivity of SR_a and SR_h underneath both tree species, and (2) examined whether drought caused differential decline of SR_a between spruce and beech. Throughout the study period, SR_a of beech accounted for 45–55% of SR_t, independent of the soil water regime; in contrast, SR_a was significantly reduced during drought in spruce, and amounted then to only 25% of SR_t. In parallel, fine-root production was decreased during 2003 by a factor of six in spruce (from 750 to 130 mg l⁻¹ a⁻¹), but remained at levels similar to those in 2002 in beech (about 470 mg l⁻¹ a⁻¹). This species-specific root response to drought was related to a stronger decline of SR_a in spruce (by about 70%) compared to beech (by about 50%). The sensitivity of SR_a and SR_h to changing T _s and available soil water was stronger in SR_a than SR_h in spruce, but not so in beech. It is concluded that SR_a determines the effect of prolonged drought on the C efflux from soil to a larger extent in spruce than beech, having potential implications for respective forest types. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands."     

Organic matter contribution to soil fertility improvement and maintenance in red Alder （Alnus rubra） silvopastoral systems

An investigation was conducted to quantify fine roots and roots nodules over the four seasons in forestry and agroforestry alder (Alnus rubra) stands in North Wales. Soil samples collected in each season were excavated at three sampling points (0.30 m, 0.57 m and 1.00 m distance from the base of each tree) from nine trees of the agroforestry and forestry plots. Result showed that the density of live fine root had significant differences in between seasons and treatments (P ＜ 0.001). The mean weight density of live fine root over the four seasons in agroforestry and forestry was 0.27±0.01 kg·m-3 and 0.54±0.03 kg·m-3, respectively. Weight density of dead root in each system remained constant throughout the year. The mean weight density of dead root was also significantly different (P ＜ 0.01) between forestry and agroforestry systems. Weight density of live and dead root nodule was both constant throughout the year and between the different sampling distances. The mean weight densities of live and dead root nodule over the four seasons were 0.09±0.03 kg·m-3 and 0.05±0.03 kg·m-3 in agroforestry and 0.08±0.02 kg·m-3 and 0.03±0.01 kg·m-3 in the forestry plots, respectively.     

Spatial distribution characteristics of fine roots of Populus euphratica in a desert riparian forest

The soil-plant system is a very important subsystem of the soil-plant-atmosphere continuum (SPAC). The water uptake by plant roots is an important subject in the research on water transport in this SPAC and is also the most active study direction in the fields of ecology, hydrology and environment. The study of the spatial distribution pattern of fine roots of plants is the basis of constructing a water absorption model of plant roots. Our study on the spatial distribution pattern of fine roots of Populus euphratica in a desert riparian forest shows that the density distribution of its root lengths can be expressed horizontally as a parabola. The fine roots are concentrated within the range of 0–350 cm from the tree trunk and their amount accounts for 91.9% of the total root mass within the space of 0–500 cm. In the vertical direction, the density distribution of the fine root lengths shows a negative exponential relation with soil depth. The fine roots are concentrated in the 0–80 cm soil layer, accounting for 96.8% of the total root mass in the 0–140 cm soil layer. __________ Translated from Chinese Journal of Ecology, 2007, 26(1): 1–4 [译自: 生态学杂志]     

Morphological plasticity of cotton roots in response to interspecific competition with pecan in an alleycropping system in the southern United States

A study was conducted in northwest Florida, USA, to investigate root development and morphology of cotton (Gossypium hirsutum L.) under pecan (Carya illinoensis K. Koch) trees in an alleycropping experiment. Root:shoot ratio, root biomass, total root length and root length density were examined under three treatments: (1) barrier (separating belowground interspecific competition by trenching to a depth of 120 cm and installing polyethylene barrier), (2) non-barrier (root systems were free to interact), and (3) monoculture of cotton (without above and belowground interspecific competition with trees). Results indicated that plants in the barrier and non-barrier treatments had lower root:shoot ratios compared to the monoculture treatment. Belowground competition for resources between pecan and cotton in the non-barrier treatment resulted in 25 and 33% reduction of total root length (359 cm) when compared to that of the barrier (477 cm) and monoculture (539 cm) treatments, respectively. The non-barrier plants also exhibited the lowest root length density. Specific root length was highest for the monoculture (179 cm g⁻¹) and lowest for the non-barrier treatment (146 cm g⁻¹) with the barrier treatment being intermediate (165 cm g⁻¹). Interspecific competition with pecan significantly altered root development and morphology of cotton plants. Research in agroforestry should take into account the developmental differences in root systems of the associated crop species so that better models incorporating nutrient and water uptake can be developed.     

Influence of slope position, stand type and rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech (Fagus orientalis Lipsky.) and spruce [Picea orientalis (L.) Link]

Oriental beech (Fagus orientalis Lipsky.) and Oriental spruce [Picea orientalis (L.) Link] are the two most common tree species in northeast Turkey. Their distribution, stand type and understorey species are known to be influenced by topographical landforms. However, little information is available as to how these changes affect litter decomposition rates of these two species. Here, we investigated the effects of slope positions (top 1,800 m, middle 1,500 m and bottom 1,200 m), stand type (pure and mixed stands) and purple-flowered rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech and spruce for 4 years using the litterbag technique in the field. Among these three factors, stand type had the strongest influence on litter decomposition (P < 0.001, F = 58.8), followed by rhododendron (P < 0.001, F = 46.8) and slope position (P < 0.05, F = 11.6). Litter decomposition was highest under mixed beech/spruce forest, followed by pure beech and spruce forest. Beech and spruce litter decomposed much faster in mixed bags (beech–spruce) than they did separately under each stand type. Purple-flowered rhododendron significantly reduced litter decomposition of Oriental beech and spruce. Beech and spruce litter decomposed much slower at top slope position than at either bottom or middle position. Differential litter decomposition of Oriental beech and spruce was mainly due to adverse conditions in spruce forest and the presence of rhododendron on the ground which was associated with lower soil pH. Higher elevations (top slope position) slowed down litter decomposition by changing environmental conditions, most probably by decreasing temperature as also other factors are different (pH, precipitation) and no detailed investigations were made to differentiate these factors. The adverse conditions for litter decomposition in spruce forest can be effectively counteracted by admixture of beech to spruce monoculture and by using the clear-cutting method for controlling rhododendron.     

Rooting patterns and fine root biomass of Pinus pinaster assessed by trench wall and core methods

Variability of fine root (diameter < 2 mm) distribution was investigated in four 55 to 56-year-old Maritime pine (Pinus pinaster) stands using a combination of trench wall observations and destructive sampling. Our objectives were to assess patterns of fine root distribution, to estimate tree fine root biomass and to explore interactions with understorey vegetation in a gradient of relevant site conditions. Results showed that root density decreased with soil depth in all stands, and variability appeared to be highest in litter and subsoil layers especially where compacted soil layers occurred. Roots were clustered in patches in the top 0–50 cm of the soil or were present as root channels at greater depths. Cluster number, cluster size and number of root channels were comparable in all four stands. Overall fine root biomass at depths of 0–120 cm ranged from 2.7 to 7.2 Mg ha⁻¹ and was highest for the two driest stands. The use of trench wall records made it possible to reduce the variability of these estimates. Understorey species represented as much as 90% of the total number of fine roots in the upper layers, and the understorey formed a considerable proportion of the total ecosystem biomass, suggesting that understorey species are likely competitors for nutrients in this ecosystem. Further studies should focus on the interaction of the understorey and pine roots and the ecological significance of clustered roots and nutrient distributions.     

Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand

We investigated fine root biomass and distribution patterns in a species-rich temperate Carpinus–Quercus–Fagus–Tilia forest and searched for experimental evidence of symmetry or asymmetry in belowground competition. We conducted extensive root coring and applied the recently introduced in situ-root growth chamber technique for quantifying fine root growth under experimentally altered intra- and interspecific root neighbourhoods in the intact stand. In 75% of all soil cores, fine roots of more than two tree species were present indicating a broad overlap of the root systems of neighbouring trees. Quercus trees had more than ten times less fine root biomass in relation to aboveground biomass or productivity (stem growth) and a much higher leaf area index/root area index ratio than Carpinus, Fagus and Tilia trees. The root growth chamber experiments indicated a high belowground competitive ability of Fagus in interspecific interactions, but a low one of Quercus. We conclude that (1) interspecific root competition is ubiquitous in this mixed stand, (2) root competition between trees can be clearly asymmetric, and (3) tree species may be ranked according to their belowground competitive ability. Fagus was found to be the most successful species in belowground competition which matches with its superiority in aboveground competition in this forest community.     

Response of root distribution of Haloxylon ammodendron seedlings to irrigation amounts in the hinterlands of the Taklimakan Desert, China

We excavated soil to study root distribution in Haloxylon ammodendron seedlings grown with different amounts of irrigation (35, 24.5 and 14 kg water for each plant each time) in the hinterland of the Taklimakan Desert. The results indicated that: 1) With decreasing irrigation amounts, the root biomass tended to be distributed in deeper soil layers. Underground biomass had a significantly negative logarithmic relationship with soil depth under different irrigation amounts. 2) Maximum horizontal spread of roots was twice that of vertical root spread, and horizontal distribution of root biomass was similar under all irrigation amounts. 3) Vertical distribution of fine roots was nearly consistent with vertical changes in soil moisture, and all had a unimodal curve; but peak values of fine root biomass in different soil layers varied with different irrigation amounts. The smaller the amount of irrigation, the deeper were the fine roots concentrated in soil layers. 4) Root length, root surface area and root volume all exhibited a unimodal curve under different irrigation amounts; the less the irrigation amount, the deeper the peak values appeared in soil layers. 5) Rootshoot ratio and ratio of vertical root depth to plant height both increased as irrigation amounts decreased. __________ Translated from Journal of Plant Ecology (Chinese Version), 2007, 31 (5): 769–776 [译自: 植物生态学报]     

Tree roots in a changing world

Globally, forests cover 4 billion hectares or 30% of the Earth's land surface, and 20%–40% of the forest biomass is made up of roots. Roots play a key role for trees: they take up water and nutrients from the soil, store carbon (C) compounds, and provide physical stabilization. Estimations from temperate forests of Central Europe reveal that C storage in trees accounts for about 110 t C ha₋₁, of which 26 t C ha₋₁ is in coarse roots and 1.2 t C ha₋₁ is in fine roots. Compared with soil C, which is about 65 t C ha₋₁ (without roots), the contribution of the root C to the total belowground C pool is about 42%. Flux of C into soils by plant litter (stemwood excluded) compared with the total soil C pool, however, is relatively small (4.4 t C ha₋₁ year₋₁) with the coarse and fine roots each contributing about 20%. Elevated CO₂ concentrations and N depositions lead to increased plant biomass, including that of roots. Recent analysis in experiments with elevated CO₂ concentrations have shown increases of the forest net primary productivity by about 23%, and, in the case of poplars, an increase of the standing root biomass by about 62%. The turnover of fine roots is also positively influenced by elevated CO₂ concentrations and can be increased in poplars by 25%–45%. A recently established international platform for scientists working on woody root processes, COST action E38, allows the exchange of information, ideas, and personnel, and it has the aim to identify knowledge gaps and initiate future collaborations and research activities.     

Root biomass and distribution of a <Emphasis Type="Italic">Picea–Abies</Emphasis> stand and a <Emphasis Type="Italic">Larix–Betula</Emphasis> stand in pumiceous Entisols in Japan

Root biomass and root distribution were studied in Entisols derived from the thick deposition of volcanic pumice on Hokkaido Island, Japan, to examine the effect of soil conditions on tree root development. The soil had a thin (<10 cm) A horizon and thick coarse pumiceous gravel layers with low levels of available nutrients and water. Two stands were studied: a Picea glehnii–Abies sachalinensis stand (PA stand) and a Larix kaempferi–Betula platyphylla var. japonica stand (LB stand). The allometric relationships between diameter at breast height (DBH) and aboveground and belowground biomass of these species were obtained to estimate stand biomass. The belowground biomass was small: 30.6 Mg ha₋₁ for the PA stand and 24.3 Mg ha₋₁ for the LB stand. The trunk/root ratios of study stands were 4.8 for the PA stand and 4.3 for the LB stand, which were higher than those from previous studies in boreal and temperate forests. All species developed shallow root systems, and fine roots were spread densely in the shallow A horizon, suggesting that physical obstruction by the pumiceous layers and their low levels of available water and nutrients restricted downward root elongation. The high trunk/root ratios of the trees may also have resulted from the limited available rooting space in the study sites.     

Biomass of fine and small roots in two Japanese black pine stands of different ages

The biomass and the spatial distribution of fine and small roots were studied in two Japanese black pine (Pinus thunbergii Parl.) stands growing on a sandy soil. More biomass of fine and small roots was found in the 17-year-old than in the 40-year-old stand. There were 62 g m⁻² of fine roots and 56 g m⁻² of small roots in the older stand, which represented mean values of 608 g for fine and 552 g for small roots per tree, respectively. In the younger stand, a total of 85 g m⁻² of fine roots and 66 g m⁻² of small roots were determined, representing a mean of 238 g for fine and 186 g for small roots per tree, respectively. Fine and small root biomasses decreased linearly with a soil depth of 0–50 cm in the older stand. In the younger stand, the fine and small roots developed only up to a depth of 30 cm. Horizontal distributions (with regard to distance from a tree) of both root groups were homogeneous. A positive correlation in the amount of biomass of fine and small roots per m² relative to tree size was found. Fine and small root biomasses increased consistently from April to July in both stands. The results also indicated earlier growth activity of the fine roots than small roots at the beginning of the growing season. The seasonal increases in fine and small root biomasses were slightly higher in the younger stand than the older stand.