Inter-annual variability of NDVI in response to long-term warming and fertilization in wet sedge and tussock tundra

This study explores the relationship between the normalized difference vegetation index (NDVI) and aboveground plant biomass for tussock tundra vegetation and compares it to a previously established NDVI–biomass relationship for wet sedge tundra vegetation. In addition, we explore inter-annual variation in NDVI in both these contrasting vegetation communities. All measurements were taken across long-term experimental treatments in wet sedge and tussock tundra communities at the Toolik Lake Long Term Ecological Research (LTER) site, in northern Alaska. Over 15 years (for wet sedge tundra) and 14 years (for tussock tundra), N and P were applied in factorial experiments (N, P and N+P), air temperature was increased using greenhouses with and without N+P fertilizer, and light intensity was reduced by 50% using shade cloth. during the peak growing seasons of 2001, 2002, and 2003, NDVI measurements were made in both the wet sedge and tussock tundra experimental treatment plots, creating a 3-year time series of inter-annual variation in NDVI. We found that: (1) across all tussock experimental tundra treatments, NDVI is correlated with aboveground plant biomass (r ²=0.59); (2) NDVI–biomass relationships for tussock and wet sedge tundra communities are community specific, and; (3) NDVI values for tussock tundra communities are typically, but not always, greater than for wet sedge tundra communities across all experimental treatments. We suggest that differences between the response of wet sedge and tussock tundra communities in the same experimental treatments result from the contrasting degree of heterogeneity in species and functional types that characterize each of these Arctic tundra vegetation communities.     

Production and export of dissolved C in arctic tundra mesocosms: the roles of vegetation and water flow

To better understand carbon (C) cycling in arctic tundra we measureddissolved C production and export rates in mesocosms of three tundra vegetationtypes: tussock, inter-tussock and wet sedge. Three flushing frequencies wereused to simulate storm events and determine potential mass export of dissolved Cunder increased soil water flow scenarios. Dissolved C production and exportrates differed between vegetation types (inter-tussock < tussock < wetsedge). In the absence of flushing, dissolved organic C (DOC) dominatedproduction in tussock and inter-tussock soils but was consumed in wet sedgesoils (8.3, 32.7, and –0.4 g C g soil^–1day^–1). Soil water dissolved C concentrations declined over time when flushedat high and medium frequencies but were variable at low flushing frequency.Total yield of dissolved C and DOC increased with increased flushing frequency.The ratio of DOC to dissolved inorganic C exported dropped with increasedflushing under tussock but not inter-tussock or wet sedge vegetation. Massexport per liter of water added declined as flushing frequency increased intussock and inter-tussock mesocosms. Export and production of dissolved C werestrongly correlated with above ground biomass, but not with photosynthetic ratesor below ground biomass. DOC quality was examined by measuring production ofToolik Lake bacteria fed mesocosm soil water. When normalized for DOCconcentration, wet sedge soil water supported significantly higher bacterialproduction. Our results indicate that arctic tundra soils have high potentialsfor dissolved C export, that water flow and vegetation type mainly controldissolved C export, and that responses of aquatic microbes to terrestrial inputsdepend on the vegetation type in the watershed.     

Soil fertility and fine root dynamics in response to 4 years of nutrient (N,P, K) fertilization in a lowland tropical moist forest,Panama

The question of how tropical trees cope with infertile soils has been challenging to address, in part, because fine root dynamics must be studied in situ. We used annual fertilization with nitrogen (N as urea, 12.5 g N m^?2 year^?1), phosphorus (P as superphosphate, 5 g P m^?2 year^?1) and potassium (K as KCl, 5 g K m^?2 year^?1) within 38 ha of old‐growth lowland tropical moist forest in Panama and examined fine root dynamics with minirhizotron images. We expected that added P, above all, would (i) decrease fine root biomass but, (ii) have no impact on fine root turnover. Soil in the study area was moderately acidic (pH = 5.28), had moderate concentrations of exchangeable base cations (13.4 cmol kg^?1), low concentrations of Bray‐extractable phosphate (PO₄ = 2.2 mg kg^?1), and modest concentrations of KCl‐extractable nitrate (NO₃ = 5.0 mg kg^?1) and KCl‐extractable ammonium (NH₄ = 15.5 mg kg^?1). Added N increased concentrations of KCl‐extractable NO₃ and acidified the soil by one pH unit. Added P increased concentrations of Bray‐extractable PO₄ and P in the labile fraction. Concentrations of exchangeable K were elevated in K addition plots but reduced by N additions. Fine root dynamics responded to added K rather than added P. After 2 years, added K decreased fine root biomass from 330 to 275 g m^?2. The turnover coefficient of fine roots <1 mm diameter ranged from 2.6 to 4.4 per year, and the largest values occurred in plots with added K. This study supported the view that biomass and dynamics of fine roots respond to soil nutrient availability in species‐rich, lowland tropical moist forest. However, K rather than P elicited root responses. Fine roots smaller than 1 mm have a short lifetime (<140 days), and control of fine root production by nutrient availability in tropical forests deserves more study.     

The seasonal dynamics of amino acids and other nutrients in Alaskan Arctic tundra soils

Past research strongly indicates the importance of amino acids in the N economy of the Arctic tundra, but little is known about the seasonal dynamics of amino acids in tundra soils. We repeatedly sampled soils from tussock, shrub, and wet sedge tundra communities in the summers of 2000 and 2001 and extracted them with water (H₂O) and potassium sulfate (K₂SO₄) to determine the seasonal dynamics of soil amino acids, ammonium (NH₄⁺), nitrate (NO₃^–), dissolved organic nitrogen (DON), dissolved organic carbon (DOC), and phosphate (PO₄^2–). In the H₂O extractions mean concentrations of total free amino acids (TFAA) were higher than NH₄⁺ in all soils but shrub. TFAA and NH₄⁺ were highest in wet sedge and tussock soils and lowest in shrub soil. The most predominant amino acids were alanine, arginine, glycine, serine, and threonine. None of the highest amino acids were significantly different than NH₄⁺ in any soil but shrub, in which NH₄⁺ was significantly higher than all of the highest individual amino acids. Mean NO₃^– concentrations were not significantly different from mean TFAA and NH₄⁺ concentrations in any soil but tussock, where NO₃^– was significantly higher than NH₄⁺. In all soils amino acid and NH₄⁺ concentrations dropped to barely detectable levels in the middle of July, suggesting intense competition for N at the height of the growing season. In all soils but tussock, amino acid and NH₄⁺ concentrations rebounded in August as the end of the Arctic growing season approached and plant N demand decreased. This pattern suggests that low N concentrations in tundra soils at the height of the growing season are likely the result of an increase in soil N uptake associated with the peak in plant growth, either directly by roots or indirectly by microbes fueled by increased root C inputs in mid-July. As N availability decreased in July, PO₄^2– concentrations in the K₂SO₄ extractions increased dramatically in all soils but shrub, where there was a comparable increase in PO₄^2– later in the growing season. Previous research suggests that these increases in PO₄^2– concentrations are due to the mineralization of organic phosphorus by phosphatase enzymes associated with soil microbes and plant roots, and that they may have been caused by an increase in organic P availability.     

Root distribution,standing crop biomass and belowground productivity in a semidesert in México

In a semidesert community in México (Zapotitlán de las Salinas, Puebla) the vertical distribution of roots and root biomass was estimated at 0–100 cm depth on two sampling dates, November 1995 (wet season) and January 1998 (dry season). Root productivity at 7 to 14.5 cm depth was estimated with the in-growth core technique every two months from March 1996 to February 1998. The relationship between environmental factors and seasonal root productivity was analyzed. Finally, we tested the effect of an irrigation equivalent to 20 mm of rain on root production. Seventy four percent of the total number of roots were found at 0-40 cm depth. Very fine roots (<1 mm diameter) were found throughout the soil profile (0-100 cm). In contrast, fine roots (1-3 mm diameter) were found only from 0–90 cm depth, and coarse roots (>3 mm diameter) from 0–60 cm depth. The root biomass was 971.5 g m^–2 (S.D. = 557.39), the very fine and fine roots representing 62.9% of the total. Total root productivity, as estimated with the ingrowth core technique, was 0.031 Mg ha^–1 over the dry season and 0.315 Mg ha^–1 over the wet season. Only very fine roots were obtained at all sampling dates. Rainfall was significantly correlated with very fine root production. The difference between fine root production in non-watered (0.054 g m^–2) and watered (0.429 g m^–2) treatments was significant. The last value was the same as that predicted for a rain of 20 mm, according to the exponential model describing the relation between the production of very fine roots and rainfall at the site.     

Responses of moist non-acidic arctic tundra to altered environment: productivity, biomass, and species richness

In arctic Alaska, researchers have manipulated air temperature, light availability, and soil nutrient availability in several tundra communities over the past two decades. These communities responded quite differently to the same manipulations, and species responded individualistically within communities and among sites. For example, moist acidic tundra is primarily nitrogen (N)‐limited, whereas wet sedge tundra is primarily phosphorus (P)‐limited, and the magnitude of growth responses varies across sites within communities. Here we report results of four years of manipulated nutrients (N and/or P) and/or air temperature in an understudied, diverse plant community, moist non‐acidic tussock tundra, in northern Alaska. Our goals were to determine which factors limit above‐ground net primary productivity (ANPP) and biomass, how community composition changes may affect ecosystem attributes, and to compare these results with those from other communities to determine their generality. Although relative abundance of functional groups shifted in several treatments, the only significant change in community‐level ANPP and biomass occurred in plots that received both N and P, driven by an increase in graminoid biomass and production resulting from a positive effect of adding N. There was no difference in community biomass among any other treatments; however, some growth forms and individual species did respond. After four years no one species has come to dominate the treatment plots and species richness has not changed. These results are similar to studies in dry heath, wet sedge, and moist acidic tundra where community biomass had the greatest response to both N and P and warming results were more subtle. Unlike in moist acidic tundra where shrub biomass increased markedly with fertilization, our results suggest that in non‐acidic tundra carbon sequestration in plant biomass will not increase substantially under increased soil nutrient conditions because of the lack of overstory shrub species.     

Soil aeration in relation to soil physical properties, nitrogen availability, and root characteristics within an arctic watershed

The seasonal change in soil oxygen availability was determined in several habitats along a topographic moisture gradient in an arctic watershed. The effect of changes in soil aeration on soil chemical and plant properties was examined by comparison of the driest (tussocks) and wettest (wet sedge tundra) sites along this gradient. Spatial variability and seasonal change in soil oxygen availability was closely linked to the hydrologic regime and the thickness of the organic soil horizon. The greatest extension of the aerobic soil layer was found beneath well-drained tussocks, while less than 10% of the unfrozen soil layer is aerated in flooded wet sedge tundra. Intertussock areas and watertracks (channels of water drainage) have intermediate levels of aeration. In tussock tundra, soil oxygen diffusion is restricted in the mineral soil layer below the organic horizon due to reduced pore space. Organic matter constituents and their change with depth were similar beneath tussocks and in wet sedge tundra, indicating that factors other than soil aeration (e.g. low soil temperatures, short growing season) are the primary controls on decomposition in these two arctic tundra systems. NH₄ ⁺, the dominant form of inorganic N, was more available in wet sedge tundra than in tussock tundra. At both sites, extractable and soil solution NO₃ ^- concentrations increased 4 to 8 fold in the second part of the growing season, indicating increased nitrifier activity with improved soil oxygen availability. Although soils thawed as deep as 60 cm, approx. 90% of the root biomass was concentrated within 20 cm of the surface. Despite the anaerobic soil environment in wet sedge tundra, the dominant species there, Eriophorum angustifolium, reached slightly greater rooting depths than E. vaginatum, whose roots grow in the elevated, aerobic portion of tussocks. E. angustifolium had a root porosity of 31%, within the range found for wetland species, while roots of E. vaginatum had a porosity close to 12%. Rhizome porosity were low in both species (11%).     

Climate and species affect fine root production with long-term fertilization in acidic tussock tundra near Toolik Lake,Alaska

Long-term fertilization of acidic tussock tundra has led to changes in plant species composition, increases in aboveground production and biomass and substantial losses of soil organic carbon (SOC). Root litter is an important input to SOC pools, although little is known about fine root demography in tussock tundra. In this study, we examined the response of fine root production and live standing fine root biomass to short- and long-term fertilization, as changes in fine root demography may contribute to observed declines in SOC. Live standing fine root biomass increased with long-term fertilization, while fine root production declined, reflecting replacement of the annual fine root system of Eriophorum vaginatum, with the long-lived fine roots of Betula nana. Fine root production increased in fertilized plots during an unusually warm growing season, but remained unchanged in control plots, consistent with observations that B. nana shows a positive response to climate warming. Calculations based on a few simple assumptions suggest changes in fine root demography with long-term fertilization and species replacement could account for between 20 and 39% of the observed declines in SOC stocks.     

Factors determining plant species richness in Alaskan arctic tundra

Abstract. We studied the relationship between plant N:P ratio, soil characteristics and species richness in wet sedge and tussock tundra in northern Alaska at seven sites. We also collected data on soil characteristics, above‐ground biomass, species richness and composition. The N:P ratio of the vegetation did not show any relationship with species richness. The N:P ratio of the soil was related with species richness for both vegetation types. Species richness in the tussock tundra was most strongly correlated with soil calcium content and soil pH, with a strong correlation between these two factors. N:P ratio of the soil was also correlated with soil pH. Other factors correlated with species richness were soil moisture and Sphagnum cover. Organic matter content was the factor most strongly correlated with species richness in the wet sedge vegetation. N:P ratio of the soil was strongly correlated with organic matter content. We conclude that N:P ratio in the vegetation is not an important factor determining species richness in arctic tundra and that species richness in arctic tundra is mainly determined by pH and flooding. In tussock tundra the pH, declining with soil age, in combination with Sphagnum growth strongly decreases species richness, while in wet sedge communities flooding over long periods of time creates less favourable conditions for species richness.     

Symbiotic N2-fixation in alpine tundra: ecosystem input and variation in fixation rates among communities

Annual inputs of symbiotic N₂-fixation associated with 3 species of alpine Trifolium were estimated in four alpine communities differing in resource supplies. We hypothesized that fixation rates would vary according to the degree of N, P, and water limitation of production, with the higher rates of fixation in N limited communities (dry meadow, moist meadow) and lower rates in P and water limited communities (wet meadow, fellfield). To estimate N₂-fixation rates, natural abundance of N isotopes (¹⁵N) were measured in field collected Trifolium and reference plants and in Trifolium plants grown in N-free medium in a growth chamber. All three Trifolium species relied on a large proportion of atmospherically-fixed N₂ to meet their N requirements, ranging from 70 to 100%. There were no apparent differences in the proportion of plant N derived from fixation among the communities, but differences in the contribution of the Trifolium species to community cover resulted in a wide range of annual N inputs from fixation, from 127 mg m^–2 year^–1 in wet meadows to 810 mg m^–2 year^–1 in fellfields. Annual spatially integrated input of symbiotic N₂-fixation to Niwot Ridge, Colorado was estimated at 490 mg m^–2 year^–1 (5 kg ha^–1 year^–1), which is relatively high in the context of estimates of net N mineralization and N deposition.     

Evaluating minirhizotron estimates of fine root longevity and production in the forest floor of a temperate broadleaf forest

The minirhizotron technique (MR) for in situ measurement of fine root dynamics offers the opportunity to obtain accurate and unbiased estimates of root production in perennial vegetation only if MR tubes do not affect the longevity of fine roots. Assuming fine root biomass is near steady-state, fine root production (g m^–2 yr^–1) can be estimated as the ratio of fine root biomass (g m^–2) to median fine root longevity (yr). This study evaluates the critical question of whether MR access tubes affect the longevity of fine roots, by comparing fine root survivorship obtained using MR with those from a non-intrusive in situ screen method in the forest floor horizons of a northern hardwood forest in New Hampshire, USA. Fine root survivorship was measured in 380 root screens during 1993–1997 and in six horizontal minirhizotron tubes during 1996–1997. No statistically significant difference was found between estimates of survivorship of fine roots (<1 mm dia.) at this site from MR versus from in situ screens, suggesting that MR tubes do not substantially affect fine root longevity in the forest floor of this northern hardwood forest and providing greater confidence in measurements of fine root production using the MR technique. Furthermore, the methodology for estimating fine root production from MR longevity data was evaluated by comparison of fine root longevity and production estimates made using single vs. multiple root cohorts, and using root-number, root-length, and root-mass weighted methods. Our results indicate that fine root-length longevity estimates based on multiple root cohorts throughout the year can be used to approximate fine root biomass production. Using this method, we estimated fine root longevity and production in the forest floor at this site to be 314 days (or 0.86 yr) and 303 g m^–2 yr^–1, respectively. Fine root production in this northern hardwood forest is approximately equivalent to standing biomass and was previously underestimated by root in-growth cores. We conclude that the use of MR to estimate fine root longevity and production as outlined here may result in improved estimates of fine root production in perennial vegetation.     

Carbon-Degrading Enzyme Activities Stimulated by Increased Nutrient Availability in Arctic Tundra Soils

Climate-induced warming of the Arctic tundra is expected to increase nutrient availability to soil microbes, which in turn may accelerate soil organic matter (SOM) decomposition. We increased nutrient availability via fertilization to investigate the microbial response via soil enzyme activities. Specifically, we measured potential activities of seven enzymes at four temperatures in three soil profiles (organic, organic/mineral interface, and mineral) from untreated native soils and from soils which had been fertilized with nitrogen (N) and phosphorus (P) since 1989 (23 years) and 2006 (six years). Fertilized plots within the 1989 site received annual additions of 10 g N⋅m^-2⋅year^-1 and 5 g P⋅m^-2⋅year^-1. Within the 2006 site, two fertilizer regimes were established – one in which plots received 5 g N⋅m^-2⋅year^-1 and 2.5 g P⋅m^-2⋅year^-1 and one in which plots received 10 g N⋅m^-2⋅year^-1 and 5 g P⋅m^-2⋅year^-1. The fertilization treatments increased activities of enzymes hydrolyzing carbon (C)-rich compounds but decreased phosphatase activities, especially in the organic soils. Activities of two enzymes that degrade N-rich compounds were not affected by the fertilization treatments. The fertilization treatments increased ratios of enzyme activities degrading C-rich compounds to those for N-rich compounds or phosphate, which could lead to changes in SOM chemistry over the long term and to losses of soil C. Accelerated SOM decomposition caused by increased nutrient availability could significantly offset predicted increased C fixation via stimulated net primary productivity in Arctic tundra ecosystems.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.     

Net Primary Production and Carbon Stocks for Subarctic Mesic–Dry Tundras with Contrasting Microtopography, Altitude, and Dominant Species

Mesic–dry tundras are widespread in the Arctic but detailed assessments of net primary production (NPP) and ecosystem carbon (C) stocks are lacking. We addressed this lack of knowledge by determining the seasonal dynamics of aboveground vascular NPP, annual NPP, and whole-ecosystem C stocks in five mesic–dry tundras in Northern Sweden with contrasting microtopography, altitude, and dominant species. Those measurements were paralleled by the stock assessments of nitrogen (N), the limiting nutrient. The vascular production was determined by harvest or in situ growing units, whereas the nonvascular production was obtained from average species growth rates, previously assessed at the sites. Results showed that aboveground vascular NPP (15–270 g m⁻²), annual NPP (214–282 g m⁻² or 102–137 g C m⁻²) and vegetation biomass (330–2450 g m⁻²) varied greatly among communities. Vegetation dominated by Empetrum hermaphroditum is more productive than Cassiope tetragona vegetation. Although the large majority of the apical NPP occurred in early-mid season (85%), production of stems and evergreen leaves proceeded until about 2 weeks before senescence. Most of the vascular vegetation was belowground (80%), whereas most of the vegetation production occurred aboveground (85%). Ecosystem C and N stocks were 2100–8200 g C m⁻² and 80–330 g N m⁻², respectively, stored mainly in the soil turf and in the fine organic soil. Such stocks are comparable to the C and N stocks of moister tundra types, such as tussock tundra. Author Contributions  Matteo Campioli, Anders Michelsen, Roeland Samson, Raoul Lemeur—conceived and designed study, Matteo Campioli, Anders Michelsen, Andreas Demey, Annemie Vermeulen—performed research, Matteo Campioli—analyzed data, and Matteo Campioli—wrote the paper.     

Plant–herbivore interactions in Alaskan arctic tundra change with soil nutrient availability

Herbivores in nutrient‐limited systems such as arctic tundra have been suggested to play a minor role in controlling plant growth simply because they are relatively few in number. However, theory predicts that as net primary productivity (NPP) increases because of greater inputs of nutrients or energy, herbivores may have greater effects on plant growth. This prediction has not been tested in the context of climate warming in arctic tundra, which may increase soil nutrient availability and thus NPP. We examined a long‐term experiment that excluded small and large mammalian herbivores and increased soil nutrients in two arctic Alaskan tundra communities: dry heath (DH) and moist acidic tussock (MAT). In the ninth year of manipulations, we measured weekly growth of three plant species of three growth forms: tussock‐forming graminoid, rhizomatous graminoid, and dwarf deciduous shrub, in each community. All species grew better when fertilized. In DH, this increase in growth was exaggerated when plants were protected from herbivores, confirming that herbivory had a negative effect on plant growth under increased nutrient conditions, but was unimportant under ambient soil conditions. However, in MAT, the importance of herbivory differed among species with fertilization. The tussock‐forming sedge at MAT, Eriophorum vaginatum, grew better and flowered more when fenced under both ambient and amended nutrients compared to plants exposed to herbivores. This species decreases in abundance in long‐term fertilized plots when mammals are present, and our results suggest that herbivory may be accounting for at least some of that loss, in addition to shifts in competitive relationships. Although we only focused on individual plants here rather than the entire community, our results suggest that under the increased soil nutrient conditions expected with continued climate warming in the Arctic, herbivores may become more important in affecting several abundant tundra plant populations, and should not be ignored.     

Fine root vertical distribution and temporal dynamics in mature stands of two enset (Enset ventricosum Welw Cheesman) clones

Quantification of the role of fine roots in the biological cycle of nutrients necessitates understanding root distribution, estimating root biomass, turnover rate and nutrient concentrations, and the dynamics of these parameters in perennial systems. Temporal dynamics, vertical distribution, annual production and turnover, and nitrogen use of fine roots (≤2 mm in diameter) were studied in mature (5-year-old) stands of two enset (Ensete ventricosum) clones using the in-growth bag technique. Live fine root mass generally decreased with increasing depth across all seasons except the dry period. Except for the dry period, more than 70% of the fine root mass was in the above 0-20 cm depth, and the fine root mass in the upper 0–10 cm depth was significantly higher than in the lowest depth (20–30 cm). Live fine root mass showed a seasonal peak at the end of the major rainy season but fell to its lowest value during the dry or short rainy season. The difference between the peak and low periods were significant (p ≤ 0.05). Fine root nitrogen (N) use showed significant seasonal variation where the mean monthly fine root N use was highest during the major rainy season. There were significant effects on N use due to depths and in-growth periods, but not due to clones. Enset fine root production and turnover ranged from 2,339 to 2,451 kg ha⁻¹ year⁻¹ and from 1.55 to 1.80 year⁻¹, respectively. Root N return, calculated from fine root turnover, was estimated at 64–65 kg ha⁻¹ year⁻¹. Fine root production, vertical distribution and temporal dynamics may be related to moisture variations and nutrient (N) fluxes among seasons and along the soil depth. The study showed that fine root production and turnover can contribute considerably to the carbon and nitrogen economy of mature enset plots.     

Litter and fine-root production in three types of tropical premontane rain forest in SE Venezuela

We investigated the productivity of three premontane primary forest sites in an extremely nutrient-poor environment on the Guyana shield in SE Venezuela. Aboveground litter production (total 5.58; leaves 4.30 Mg ha^–1 year^–1) was low, but comparable to other tropical forests. Due to the low nutrient status, net production of fine roots was among the highest ever reported (11.14 Mg ha^–1 year^–1). Only 20% of fine root stock was alive, 80% consisted of necromass. Similar values were obtained for dead and living root tips. Element concentrations in fine roots (including bio- and necromass), especially Ca, were low, whereas Al concentrations were relatively high.The upper limit of average fine root lifetime was 253 days. We conclude that the high proportion of necromass was mainly caused by slow mineralisation of nutrient-poor fine roots.     

Mean annual temperature influences local fine root proliferation and arbuscular mycorrhizal colonization in a tropical wet forest

Mean annual temperature (MAT) is an influential climate factor affecting the bioavailability of growth‐limiting nutrients nitrogen (N) and phosphorus (P). In tropical montane wet forests, warmer MAT drives higher N bioavailability, while patterns of P availability are inconsistent across MAT. Two important nutrient acquisition strategies, fine root proliferation into bulk soil and root association with arbuscular mycorrhizal fungi, are dependent on C availability to the plant via primary production. The case study presented here tests whether variation in bulk soil N bioavailability across a tropical montane wet forest elevation gradient (5.2°C MAT range) influences (a) morphology fine root proliferation into soil patches with elevated N, P, and N+P relative to background soil and (b) arbuscular mycorrhizal fungal (AMF) colonization of fine roots in patches. We created a fully factorial fertilized root ingrowth core design (N, P, N+P, unfertilized control) representing soil patches with elevated N and P bioavailability relative to background bulk soil. Our results show that percent AMF colonization of roots increased with MAT (r² = .19, p = .004), but did not respond to fertilization treatments. Fine root length (FRL), a proxy for root foraging, increased with MAT in N+P‐fertilized patches only (p = .02), while other fine root morphological parameters did not respond to the gradient or fertilized patches. We conclude that in N‐rich, fine root elongation into areas with elevated N and P declines while AMF abundance increases with MAT. These results indicate a tradeoff between P acquisition strategies occurring with changing N bioavailability, which may be influenced by higher C availability with warmer MAT.     

Tundra plant structure and production in relation to the environment

The alpine and polar climatic limit for growth of woody plants is very much dependent on the mean temperatures of the warmest three or four summer months. Tundra plants with perennating buds close to the ground are sheltered by insulating snow cover. Many tundra plants can grow at temperatures 5–10°C below 0°C and also have low optimum temperatures. Total net production of tundra plants may be as high as 900 g/m²/yr as dry weight in moist and eutrophic low alpine shrub tundra and in antarctic moss mats. The variation in tundra plant production is often observed to be greater between different stands (communities) within one locality than between localities, because of very important variation in soil moisture and nutrients between the stands. On a global scale the biomass of vascular plants increases by an order of magnitude from the climatic severe polar desert to semidesert and again from there to moist shrub tundra. The cryptogam biomass increases only 2–10 fold from polar desert to low arctic shrub tundra. To a certain limit unfavourable climatic conditions are worse to above- than to belowground plant parts. Highest root biomass compared to top (up to 20 times higher) is observed in wet monocotyledonous polar and alpine communities. In polar desert root biomass is small again, as compared to tops and also in lower latitudes and altitudes of temperate regions.Presented at he Eighth International Congress of Biometeorology, 9–14 September 1979, Shefayim, Israel.