A comparative study on nutrient cycling in wet heathland ecosystems

Summary The term relative nutrient requirement is introduced in order to measure and to compare the nutrient losses from different perennial plant populations and the amount of nutrient that they need to absorb to compensate these losses. The relative nutrient requirement (L) is defined as the amount of the growth-limiting nutrient that must be taken up to maintain or replace each unit of biomass during a given time interval (e.g., mgN g^-1 biomass year^-1). It is derived that in a plant community with two competing perennial plant populations, species¹ will become dominant if the relative competition coefficient k ₁₂ (sensu De Wit 1960) exceeds the ratio between the relative nutrient requirements of the two species (L ₁/L ₂), whereas species 2 will become dominant, if k ₁₂ is below this critical ratio. The above-ground litter production was measured inwet heathland communities dominated by Erica tetralix or by Molinia caeruleain order to estimate N and P losses from theaboveground biomass and to calculate the relative N and P requirements of these species. Molinia lost during one year 63% and 34%, respectively, of the amount of N and P present in the above-ground biomass at the end of the growing season. These losses were in Erica 27% and 31%, respectively. The relative N requirements of the two species show the same difference: 7.5 and 2.6 mg N g^-1 yr^-1, respectively, in Molinia and in Erica. Also the relative P requirement of Molinia is higher as well as that of Erica (0.18 versus 0.08 mg P g^-1 yr^-1). The relative competition coefficient of Molinia with respect to Erica (k _me ) is equal to unity under unfertilized conditions but increases with increasing nutrient supply. Under nutrient-poor conditions k _me is below the critical ratio of the relative nutrient requirements of the two species (L _m /L _e =2.9 or 2.3), so that Erica will be the dominant species. After an increase in nutrient availability k _me increases and exceeds this critical limit which results in Molinia replacing Erica. During the last 20 years this replacement of Erica-dominant communities by monocultures of Molinia has been observed in almost all wet heathlands in The Netherlands along with a strong increase in nitrogen availability.     

Nutrient use efficiency in evergreen and deciduous species from heathlands

Summary The nutrient (N, P) use efficiency (NUE: g g^–1 nutrient), measured for the entire plant, of field populations of the evergreen shrubs Erica tetralix (in a wet heathland) and Calluna vulgaris (in a dry heathland) and the deciduous grass Molinia caerulea (both in a wet and a dry heathland) was compared. Erica and Calluna are crowded out by Molinia when nutrient availability increases. NUE was measured as the product of the mean residence time of a unit of nutrient in the population (MRT: yr) and nutrient productivity (A: annual productivity per unit of nutrient in the population, g g^–1 nutrient yr^–1. It was hypothesized that 1) in low-nutrient habitats selection is on features leading to a high MRT, whereas in high-nutrient habitats selection is on features leading to a high A; and that 2) due to evolutionary trade-offs plants cannot combine genotypically determined features which maximize both components of NUE.Both total productivity and litter production of the Molinia populations exceeded that of both evergreens about three-fold. Nitrogen and phosphorus resorption from senescing shoots was much lower in the evergreens compared with Molinia. In a split-root experiment no nutrient resorption from senescing roots was observed. Nutrient concentrations in the litter were equal for all species, except for litter P-concentration of Molinia at the wet site. Both Erica and Calluna had a long mean residence time of both nitrogen and phosphorus and a low nitrogen and phosphorus productivity. The Molinia populations showed a shorter mean residence time of N and P and a higher N- and P-productivity. These patterns resulted in an equal nitrogen use efficiency and an almost equal phosphorus use efficiency for the species under study. However, when only aboveground NUE was considered the Molinia populations had a much higher NUE than the evergreens.The results are consistent with the hypotheses. Thus, the low potential growth rate of species from low-nutrient habitats is probably the consequence of their nutrient conserving strategy rather than a feature on which direct selection takes place in these habitats.     

Nitrogen losses from perennial grass species

Nitrogen losses from plants may occur through a variety of pathways, but so far, most studies have only quantified losses of nutrients by above-ground litter production. We used ¹⁵N pulse labelling to quantify total nitrogen losses from above- and below-ground plant parts. Using this method we were able to include also pathways other than above-ground litter production. To test the hypothesis that species from nutrient-poor habitats lose less nitrogen than species from more fertile soils, six perennial grasses from habitats with a wide range of nutrient availability were investigated: Lolium perenne, Arrhenatherum elatius, Anthoxanthum odoratum, Festuca rubra, F. ovina and Molinia caerulea. The results of an experiment carried out in pots in a green-house at two fertility levels show that statistically significant losses occur through pathways other than above-ground litter production. In the low fertility treatment, most (70%) losses from L. perenne occurred by litter production, but in Ar. elatius, F. rubra, F. ovina and M. caerulea, more than 50% of labelled N losses took place by root turn-over, leaching or exudation from roots. When nutrient supply increased, the ¹⁵N losses in above-ground dead material increased in all species and in Ar. elatius, A. odoratum and F. rubra the ¹⁵N losses via other pathways decreased. Ranked according to decreasing turnover coefficient the sequence of species was: L. perenne, A. odoratum, F. rubra, F. ovina, Ar. elatius, M. caerulea. These results suggest that species adapted to sites with low availability of nutrients lose less nitrogen (including above- and below-ground losses) than species adapted to more fertile soils.     

The effect of increased nutrient availability on vegetation dynamics in wet heathlands

A three year fertilization experiment was conducted in which nitrogen (N series: 20 g N m^–2 yr^–1), phosphorus (P series: 4 g P m^–2 yr^–1) and potassium (K series: 20 g K m^–2 yr^–1) were added to a mixed vegetation of Erica tetralix and Molinia caerulea. At the end of each growing season the percentage cover of each species was determined. At the end of the experiment percentage cover of each species was found to be positively correlated with the harvested biomass. In the unfertilized control series the cover of Erica and Molinia did not change significantly during the experiment. In all fertilized series however, especially in the P series, cover of Erica decreased significantly. The cover of Molinia increased significantly in the P series only.In the fertilized series the biomass of Erica and total biomass per plot did not change significantly compared with the control series. In the P series the biomass of Molinia increased significantly.It is suggested that with increasing phosphorus or nitrogen availability Molinia outcompetes Erica because the former invests more biomass in leaves which in turn permits more carbon to be allocated to the root system, which thereupon leads to a higher nutrient uptake.     

Root production and root turnover in two dominant species of wet heathlands

Summary Root biomass production, root length production and root turnover of Erica tetralix and Molinia caerulea were estimated by sequential core sampling and by observations in permanent minirhizotrons in the field. Root biomass production, estimated by core sampling, was 370 (Erica) and 1080 (Molinia) g m^-2 yr^-1. This was for both species equal to aboveground production. Assuming steady-state conditions for the root system, root biomass turnover rates (yr^-1), estimated by core sampling, were 1.72 (Erica) and 1.27 (Molinia). Root length production of both species, estimated by minirhizotron observations, varied significantly with observation depth. Root length turnover rate (yr^-1) of both species did not vary significantly with observation depth and averaged 0.92 in Erica and 2.28 in Molinia. Reasons are given for the discrepancy between the results of the two types of turnover measurements. The data suggest that the replacement of Erica by Molinia in a wet heathland, which occurs when nutrient availability increases, leads to an increased flow of carbon and nutrients into the soil-system. Therefore, there may be a positive feedback between dominance of Molinia and nutrient availability.     

A comparison of mineral uptake and translocation by above-ground and below-ground root systems ofSalix syringiana

Nutrient uptake and translocation by above-ground adventitious roots and below-ground roots of woodySalix syringiana saplings were studied with gamma spectrometry. Each of four radionuclides (⁷⁵Se,¹³⁸Cs,⁵⁴Mn, and⁶⁵Zn) administered to adventitious and belowground roots were detected in stems and leaves within one month. Nuclides tended to be immobilized in the leaves and branches closest to the adventitious roots that absorbed them, while nuclides absorbed from below-ground sources were distributed more evenly throughout the plant. The capacity of adventitious roots to acquire nutrients from above-ground sources suggests they function as a potential auxiliary pathway of nutrient uptake and might enhance plant nutrient status where below-ground root uptake it hindered by adverse soil conditions.     

The relation between above- and belowground biomass allocation patterns and competitive ability

Summary In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m⁻² yr⁻¹). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.     

羌塘高原降水梯度植物叶片、根系性状变异和生态适应对策

叶片和根系是植物获取资源的最重要的器官,其性状随环境梯度的变化反映了植物光合碳获取和水分与养分的吸收能力及其对环境变化适应的生态对策。羌塘高原降水梯度带高寒草地群落叶片和根系成对性状关系研究不仅能揭示环境梯度对植物性状的塑造作用,也可为理解寒、旱和贫瘠等极端环境下植物的适应策略提供依据。为此,选择3组具有代表性的叶片和根系成对性状:比叶面积(SLA)和比根长(SRL);单位质量叶氮含量(LN_(mass))和单位质量根氮含量(RN_(mass));单位面积叶氮含量(LN_(area))和单位长度根氮含量(RN_(length)),分析不同优势植物地上、地下成对性状变异特征及其与环境因子的关系,探讨植物性状对高寒生态系统水分和养分限制因素的适应策略。研究表明,区域气候和土壤环境导致的叶片性状变异大于根系性状的变异,干旱端的植物既具有高的SRL,又具有高的叶片和根系的养分含量(LN_(mass),LN_(area)和RN_(mass))。SLA-SRL、LN_(mass)-RN_(mass)、LN_(area)-RN_(length)均表现为权衡关系,在干旱端(年降雨量MAP 400 mm)的高寒草原、荒漠草原和极湿润端(MAP 600 mm)的高寒草甸这种权衡关系更为明显,而中间区域(400 MAP 600 mm)的高寒草甸养分和水分限制不是很强烈,叶片和根系性状更多地表现出协同关系。从植物功能类群来看,苔草和禾草类植物叶片和根系成对性状之间具有更强烈的权衡关系。干旱端植物通过增加SRL和叶片、根系养分含量来提高水分和养分的吸收能力,同时通过叶片高的氮含量提高光合碳获取能力,保障了根系生长的物质来源,表现出地上和地下同时投入的策略。干旱端植物保持较高的养分含量是抵御和适应严酷的寒、旱和贫瘠的环境胁迫的重要策略。而在湿润端植物则采取增加SLA,维持地上光合生产力的生态策略。     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

Fire increases aboveground biomass, seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Role of desert annuals in nutrient flow in arid area of Northwestern China: a nutrient reservoir and provider

Previous studies have tested the “vernal dam” hypothesis of spring ephemeral herbs in hardwood forests. The desert annual is a component of the desert ecosystem that takes advantage of water resources and temperature conditions during the rainy season to rapidly complete its life cycle within several months. To understand the role desert annual/ephemeral plants play in nutrient flow, we studied vegetation cover, nitrogen content and litter production of annual plants and litter decomposition rate in plant communities dominated by four shrubs (Haloxlon ammodendron, Hedysarum scoparium, Calligonum mongolicum, and Nitraria tangutorum) and two dominant annuals (Agriophyllum squarrosum and Halogeton arachnoideus Moq) in Minqin, northwestern China. Results indicate that over half of the total vegetation cover was provided by annuals. Annuals also took up a large amount of nitrogen (0.46–3.78 g N m⁻²) along the oasis–desert ecotone. Litter production and nutrient content were higher in areas dominated by annual plants than in areas dominated by shrubs. Furthermore, the litter decomposition rate of the annuals was higher than that of the shrubs, except for the shrub H. ammodendron, although almost all of the litter’s carbon (C) and nitrogen (N) remained after 6 months of decomposition. Without the annuals, more nutrients and rainwater might be lost through leaching or dust transfer caused by the wind erosion. In addition, green twigs of the annuals are the food for some animals, we found some green twigs and litter from annuals left in front of gerbil and rabbit burrows, sometimes even blocking these burrows. Thus, desert summer annuals, like nutrient reservoirs and providers, take up nutrients during the rainy season, providing some animals and microbes with food, and finally release these nutrients after death. Bao-Ming Chen and Gen-Xuan Wang contributed equally to this work.     

Tropical tree species composition affects the oxidation of dissolved organic matter from litter

Plant species effects on soil nutrient availability are relatively well documented, but the effects of species differences in litter chemistry on soil carbon cycling are less well understood, especially in the species-rich tropics. In many wet tropical forest ecosystems, leaching of dissolved organic matter (DOM) from the litter layer accounts for a significant proportion of litter mass loss during decomposition. Here we investigated how tree species differences in soluble dissolved organic C (DOC) and nutrients affected soil CO₂ fluxes in laboratory incubations. We leached DOM from freshly fallen litter of six canopy tree species collected from a tropical rain forest in Costa Rica and measured C-mineralization. We found significant differences in litter solubility and nutrient availability. Following DOM additions to soil, rates of heterotrophic respiration varied by as much as an order of magnitude between species, and overall differences in total soil CO₂ efflux varied by more than four-fold. Variation in the carbon: phosphorus ratio accounted for 51% of the variation in total CO₂ flux between species. These results suggest that tropical tree species composition may influence soil C storage and mineralization via inter-specific variation in plant litter chemistry.     

Polyphenols in litter from tropical montane forests across a wide range in soil fertility

In nutrient-poor ecosystems high polyphenol concentrations in plant litter have been proposed to influence soil nutrient availability in benefit of the plants. We addressed the question whether litter polyphenol concentrations vary across a soil chronosequence of almost identical geology, climate and plant species composition, but of a wide range in nitrogen (N) and phosphorus (P) availability in the Hawaiian Islands. Concentrations of total phenolics (TPh) and proanthocyanidins (PA) in leaf litter of the dominant tree species Metrosideros polymorpha were higher at the oldest, P-limited site compared to the youngest, N-limited site, with intermediate values at the two relatively fertile sites co-limited by N and P. Polyphenol concentrations in fine root litter differed considerably from those observed in leaf litter and varied differently across the soil age gradient. Long-term fertilization did not significantly alter polyphenol concentrations in Metrosideros litter at either site. Moreover, green leaves and leaf litter of Metrosideros showed similar relative differences among sites when compared between natural populations and plants from the same populations but grown in a common garden. These results suggest that polyphenol concentrations inherently vary among populations of the dominant tree species in Hawaiian montane forests possibly indicating an adaptation to ecosystem properties such as substrate age related differences in soil fertility. The combined above- and below-ground input rate of TPh ranged from 62.4 to 170.8 g/m²/yr and was significantly higher at the P-limited than at the N-limited site. Root-derived polyphenols contributed a much higher absolute and relative amount of phenolic input at the N-limited than at the P-limited site. The differences in amount, quality, and pathways of input might suggest specific interactions with soil processes and nutrient cycling among the Hawaiian rainforests studied here.     

Fine root turnover and litter production of Norway spruce in a long-term temperature and nutrient manipulation experiment

Background and aims

Increased soil temperature and nutrient availability enhance soil biological activity. We studied how these affect fine root growth and survival, i.e. below-ground litter production, in relation to above-ground foliage litter production of Norway spruce (Picea abies (L.) Karst.).

Methods

The treatments, irrigation (I), soil warming + irrigation (WI), fertilization + irrigation (FI) and soil warming + fertilization + irrigation (WFI) were started in 1987 (F, I) and in 1995 (W). The annual production of fine root litter was estimated from minirhizotrons (survival) and soil-cores (biomass) and the annual above-ground litter production from litter traps.

Results and conclusions

The number and elongation of fine roots tended to be higher in WI and I compared to the other treatments, which may indicate nutrient shortage. Fine roots in the WFI treatment had the lowest median longevity and from three to fourfold higher below-ground litter production compared to WI, FI or I - higher soil temperature increased the litter input particularly into the mineral soil. Only fertilization increased the above-ground litter production. As warmer and more nutrient-rich soil significantly shortened the fine root lifespan and increased the litter input, the storage of carbon in boreal forest soil may increase in the future.     

Nutrient release from decomposing leaf litter of Banksia ornata,Dark Island heathland,South Australia

Distinct O₁ and O₂ layers, representing annual litter fall, enabled the sequential loss of biomass and nutrients (phosphorus and nitrogen) to be reconstructed in undisturbed litter layers of Banksia ornata in the Dark Island heathland, South Australia. Apart from an initial loss in biomass and nitrogen, the dry weight and nutrient content of the O₁ layer, exposed to the desiccating influence of the atmosphere, remained relatively constant until covered by the following year's leaf fall. Under the blanket of newly fallen leaves, biomass decomposition proceeded continuously through autumn, winter, spring, into the dry summer season. Even though the biomass of the decomposing leaf (O₂) layer decreased continuously, its nutrient content remained relatively constant until the summer season was reached when total decomposition and nutrient loss occurred. During spring, fine rootlets invaded the decomposing litter layer (O₂) and, together with decomposer fungi, bacteria and soil fauna, maintained the total nutrient content of the decomposing leaf at a constant level. By late spring-early summer shoot growth of the dominant heath species was initiated, inducing the mobilization of the nutrients stored in the decomposing litter layer.     

Effects of elevated CO2 on nutrient cycling in a sweetgum plantation

The effects of elevated CO₂ on nutrient cycling and selected belowground processes in the closed-canopy sweetgum plantation were assessed as part of a free-air CO₂ enrichment (FACE) experiment at Oak Ridge, Tennessee. We hypothesized that nitrogen (N) constraints to growth response to elevated CO₂ would be mitigated primarily by reduced tissue concentrations (resulting in increased biomass production per unit uptake) rather than increased uptake. Conversely, we hypothesized that the constraints of other nutrients to growth response to elevated CO₂ would be mitigated primarily by increased uptake because of adequate soil supplies. The first hypothesis was not supported: although elevated CO₂ caused reduced foliar N concentrations, it also resulted in increased uptake and requirement of N, primarily because of greater root turnover. The additional N uptake with elevated CO₂ constituted between 10 and 40% of the estimated soil mineralizeable N pool. The second hypothesis was largely supported: elevated CO₂ had no significant effects on tissue concentrations of P, K, Ca, or Mg and caused significantly increased uptake and requirement of K, Ca, and Mg. Soil exchangeable pools of these nutrients are large and should pose no constraint to continued growth responses. Elevated CO₂ also caused increased microbial biomass, reduced N leaching and increased P leaching from O horizons (measured by resin lysimeters), reduced soil solution NH ₄ ⁺ , SO ₄ ^2– , and Ca²⁺ concentrations, and increased soil solution pH. There were no statistically significant treatment effects on soil nutrient availability as measured by resin capsules, resin stakes, or in situ incubations. Despite significantly lower litterfall N concentrations in the elevated CO₂ treatment, there were no significant treatment effects on translocation or forest floor biomass or nutrient contents. There were also no significant treatment effects on the rate of decomposition of fine roots. In general, the effects of elevated CO₂ on nutrient cycling in this study were not large; future constraints on growth responses imposed by N limitations will depend on changes in N demand, atmospheric N deposition, and soil mineralization rates.     

Fire in the Brazilian Amazon: 1. Biomass,nutrient pools,and losses in slashed primary forests

Deforestation in the Brazilian Amazon has resulted in the conversion of >230,000 km² of tropical forest, yet little is known on the quantities of biomass consumed or the losses of nutrients from the ecosystem. We quantified the above-ground biomass, nutrient pools and the effects of biomass burning in four slashed primary tropical moist forests in the Brazilian Amazon. Total above-ground biomass (TAGB) ranged from 292 Mg ha^-1 to 436 Mg ha^-1. Coarse wood debris (>20.5 cm diameter) was the dominant fuel component. However, structure of the four sites were variable. Coarse wood debris comprised from 44% to 69% of the TAGB, while the forest floor (litter and rootmat) comprised from 3.7 to 8.0% of the TAGB. Total biomass consumption ranged from 42% to 57%. Fires resulted in the consumption of >99% of the litter and rootmat, yet <50% of the coarse wood debirs. Dramatic losses in C, N, and S were quantified. Lesser quantities of P, K, and Ca were lost by combustion processes. Carbon losses from the ecosystem were 58–112 Mg ha^-1. Nitrogen losses ranged from 817 to 1605 kg ha^-1 and S losses ranged from 92 to 122 kg ha^-1. This represents losses that are as high as 56%, 68%, and 49% of the total above-ground pools of these nutrients, respectively. Losses of P were as high as 20 kg ha^-1 or 32% of the above-ground pool. Losses to the atmosphere arising from primary slash fires were variable among sites due to site differences in concentration, fuel biomass, and fuel structure, climatic fluctuations, and anthropogenic influences. Compared to fires in other forest ecosystems, fires in slashed primary tropical evergreen forests result in among the highest total losses of nutrients ever measured. In addition, the proportion of the total nutrient pool lost from slash fires is higher in this ecosystem compared to other ecosystems due to a higher percentage of nutrients stored in above-ground biomass.     

Plant Biomass,Nutrient Concentration and Nutrient Storage in a Tropical Dry Forest in the South–west of Madagascar

Plant biomass, mineral composition and the amounts of nutrients in the different fractions of the vegetation were determined for a dense dry deciduous forest growing on light red sands in south-western Madagascar. Complete harvesting and soil coring were used to determine the above- and below-ground biomass respectively. The above-ground biomass, weighing 118 t ha⁻¹ (dry matter), was mostly (96%) made up of phanerophytes (woody trees and shrubs >25 cm tall). Dead material (litter and dead wood on the soil surface) represented 13.8 t ha⁻¹. These results fit well into the range of values reported for other tropical ecosystems. The below-ground biomass was 17.8 t ha⁻¹ giving a root/shoot ratio of 0.15. Rooting is superficial. The nutrient concentration in this dry forest on light reddish-brown sands is, as in other dry forests, considerably higher than that usually found for humid forests. Calcium is the most abundant element. The plant biomass Ca/K ratio is much higher than that of humid tropical forests. In spite of its high originality, this Madagascan dry forest has the same behaviour as other dry forests of the world.     

Global change and root function

Global change includes land-use change, elevated CO₂ concentrations, increased temperature and increased rainfall variability. All four aspects by themselves and in combination will influence the role of roots in linking below- and above-ground ecosystem function via organic and inorganic resource flows. Root-mediated ecosystem functions which may be modified by global change include below-ground resource (water, nutrients) capture, creation and exploitation of spatial heterogeneity, buffering of temporal variations in above-ground factors, supply and storage of C and nutrients to the below-ground ecosystem, mobilization of nutrients and C from stored soil reserves, and gas exchange between soil and atmosphere including the emission from soil of greenhouse gases. The theory of a functional equilibrium between root and shoot allocation is used to explore predicted responses to elevated CO₂ in relation to water or nutrient supply as limiting root function. The theory predicts no change in root:shoot allocation where water uptake is the limiting root function, but substantial shifts where nutrient uptake is (or becomes) the limiting function. Root turnover will not likely be influenced by elevated CO₂, but by changes in regularity of water supply. A number of possible mechanisms for root-mediated N mineralization is discussed in the light of climate change factors. Rhizovory (root consumption) may increase under global change as the balance between plant chemical defense and adapted root consuming organisms may be modified during biome shifts in response to climate change. Root-mediated gas exchange allows oxygen to penetrate into soils and methane (CH₄) to escape from wetland soils of tundra ecosystems as well as tropical rice production systems. The effect on net greenhouse gas emissions of biome shifts (fens replacing bogs) as well as of agricultural land management will depend partly on aerenchyma in roots.