Non-symbiotic nitrogen fixation in forest ecosystems

Ecological Research - Since the first attempts to increase forest growth using nitrogen (N) fixation, biological N fixation has been intensively studied. Numerous studies that focused on symbiotic...     

Non-symbiotic nitrogen fixing bacteria in tropical soils

Summary Several aspects of heterotrophic, free-living nitrogen-fixing bacteria specific to tropical environments are presented. WhileAzotobacter chroococcum andA. vinelandii seem to occur throughout the world,A. paspali, Derxia gummosa andBeijerinckia spp. have been found mainly in the tropics. Derxia occurred more frequently than previously supposed and was stimulated by moist soil conditions, particularly after flooding.A. paspali growth in soil even under experimental conditions, was strictly limited to the rhizosphere of a few varieties ofPaspalum notatum and the association took almost a year to become established. This work was supported by the Brazilian National Research Council.     

Climate and wildfires in the North American boreal forest

The area burned in the North American boreal forest is controlled by the frequency of mid-tropospheric blocking highs that cause rapid fuel drying. Climate controls the area burned through changing the dynamics of large-scale teleconnection patterns (Pacific Decadal Oscillation/El Niño Southern Oscillation and Arctic Oscillation, PDO/ENSO and AO) that control the frequency of blocking highs over the continent at different time scales. Changes in these teleconnections may be caused by the current global warming. Thus, an increase in temperature alone need not be associated with an increase in area burned in the North American boreal forest. Since the end of the Little Ice Age, the climate has been unusually moist and variable: large fire years have occurred in unusual years, fire frequency has decreased and fire–climate relationships have occurred at interannual to decadal time scales. Prolonged and severe droughts were common in the past and were partly associated with changes in the PDO/ENSO system. Under these conditions, large fire years become common, fire frequency increases and fire–climate relationships occur at decadal to centennial time scales. A suggested return to the drier climate regimes of the past would imply major changes in the temporal dynamics of fire–climate relationships and in area burned, a reduction in the mean age of the forest, and changes in species composition of the North American boreal forest.     

Amino acid abundance and proteolytic potential in North American soils

Studies of nitrogen (N) cycling have traditionally focused on N mineralization as the primary process limiting plant assimilation of N. Recent evidence has shown that plants may assimilate amino acids (AAs) directly, circumventing the mineralization pathway. However, the general abundance of soil AAs and their relative importance in plant N uptake remains unclear in most ecosystems. We compared the concentrations and potential production rates of AAs and NH₄ ⁺, as well as the edaphic factors that influence AA dynamics, in 84 soils across the United States. Across all sites, NH₄ ⁺ and AA-N comprised similar proportions of the total bioavailable N pool (~20%), with NO₃ ⁻ being the dominant form of extractable N everywhere but in tundra and boreal forest soils. Potential rates of AA production were at least comparable to those of NH₄ ⁺ production in all ecosystems, particularly in semi-arid grasslands, where AA production rates were six times greater than for NH₄ ⁺ (P < 0.01). Potential rates of proteolytic enzyme activity were greatest in bacteria-dominated soils with low NH₄ ⁺ concentrations, including many grassland soils. Based on research performed under standardized laboratory conditions, our continental-scale analyses suggest that soil AA and NH₄ ⁺ concentrations are similar in most soils and that AAs may contribute to plant and microbial N demand in most ecosystems, particularly in ecosystems with N-poor soils.     

Biomass and carbon storage of the North American deciduous forest

Field measures of tree and shrub dimensions were used with established biomass equations in a stratified, two-stage cluster sampling design to estimate above-ground ovendry woody biomass and carbon storage of the eastern deciduous forest of North America. Biomass averaged 8.1 ± 1.4 (95% C.I.) kg/m² and totaled 18.1 ± 3.1 (95% C.I.) gigatons. Carbon storage averaged 3.6 ± 0.6 (95% C.I.) kg/m² and totaled 8.1 ± 1.4 (95% C.I.) gigatons. These values are lower than previous estimates commonly used in the analysis of the global carbon budget which range from 17.1 to 23.1 kg/m² for biomass and 7.7 to 10.4 kg/m² for carbon storage. These new estimates for the deciduous forest, together with earlier work in the boreal forest begin to reveal a pattern of overestimation of global carbon storage by vegetation in analyses of the global carbon budget. We discuss reasons for the differences between the new and earlier estimates, as well as implications for our understanding of the global carbon cycle.     

Plant nitrogen and phosphorus limitation in 98 North American grassland soils

The availability of nutrients is a critical determinant of ecological dynamics in grasslands, but the relationships between soil resource availability and nutrient limitation across ecosystems are not clear. To better understand how soil nutrient availability determines nutrient limitation in vegetation, we grew the same species of grass (Schizachyrium scoparium) in 98 North American grassland soils and fertilized them factorially with nitrogen (N) and phosphorus (P). On average adding N, P, and the two nutrients together increased biomass relative to unfertilized plants by 81%, 22%, and 131%, respectively. Plants grown on low-P soils were not primarily limited by P. Instead, these plants were colimited by N and P, while plants grown on high-P soils were primarily limited by N and only secondarily limited by P. Limitation was not predicted by total soil N. The preponderance of colimitation between N and P on low-P soils suggests that low P availability alters the N cycle to constrain supplies to plants such that N and P are made available in proportion to their demand by plants.     

Rhizobitoxine production and symbiotic compatibility of Bradyrhizobium from Asian and North American lineages of amphicarpaea.

Reciprocal inoculations with Bradyrhizobium sp. isolates from the North American legume Amphicarpaea bracteata (L.) Fern. (Phaseoleae-Glycininae) and from a Japanese population of its close relative Amphicarpaea edgeworthii (Benth.) var. japonica were performed to analyze relative symbiotic compatibility. Amphicarpaea edgeworthii plants formed few or no nodules with any North American bradyrhizobial strains isolated from A. bracteata, but all A. bracteata lineages formed effective nitrogen-fixing nodules with Japanese Bradyrhizobium isolates from A. edgeworthii. However, one group of A. bracteata plants (lineage Ia) when inoculated with Japanese bradyrhizobia developed a striking leaf chlorosis similar to that known to be caused by rhizobitoxine. The beta-cystathionase inhibition assay demonstrated that significant amounts of rhizobitoxine were present in nodules formed by these Japanese bradyrhizobia. No North American bradyrhizobial isolate from A. bracteata induced chlorosis on any plants, and the beta-cystathionase assay failed to detect rhizobitoxine in nodules formed by these isolates. The role of rhizobitoxine in A. edgeworthii nodulation development was tested by inoculating plants with a Bradyrhizobium elkanii rhizobitoxine-producing strain, USDA 61, and two mutant derivatives, RX17E and RX18E, which are unable to synthesize rhizobitoxine. Amphicarpaea edgeworthii inoculated with wild-type USDA 61 developed >150 nodules per plant, while plants inoculated with RX17E and RX18E developed fewer than 10 nodules per plant. Thus, efficient nodule development in A. edgeworthii appears to be highly dependent on rhizobitoxine production by Bradyrhizobium strains.     

Longitudinal patterns of plant diversity in the North American boreal forest

Spatial patterns of plant diversity in the North American boreal forest were examined according to three plant life forms (woody plants, herbaceous plants, and bryophytes) and two taxonomic levels (species and genus), using sixty 9-ha plots sampled in white spruce (Picea glauca (Moench) Voss) and black spruce (Picea mariana (P. Mill.) B.S.P.) ecosystems along a transcontinental transect from the Pacific coast eastwards to the Atlantic coast. The patterns of inventory diversity (represented by alpha diversity), differentiation diversity (represented by the similarity index, habitat-heterogeneity index, similarity decay rate, and length of the first axis in detrended correspondence analysis), and pattern diversity (represented by the mosaic diversity index) were assessed along the transect in both ecosystem types. At the stand level, central North America had the highest alpha diversity in terms of the number of species or genera, and western North America had a higher alpha diversity than eastern North America. At the continental scale, herbaceous plants had the highest beta diversity in terms of floristic change from the eastern to western North America, bryophytes had the lowest beta diversity, and woody plants were in the middle, regardless of ecosystem type and taxonomic level. Central North America had the lowest mosaic diversity across the boreal transect of North America. The white spruce ecosystems had a higher alpha diversity than the black spruce ecosystems regardless of plant life form, taxonomic level and geographic location. The white spruce ecosystems tended to have more bryophytes, less woody plants, and higher species:genus ratio than the black spruce ecosystems. In general, the white spruce and black spruce ecosystems shared the same patterns in diversity changes at different spatial scales, plant life forms, and taxonomic levels across the transect studied. The existing patterns of plant diversity in the North American boreal forest area resulted from a combination of ecological processes and spatial configuration.     

Observed forest sensitivity to climate implies large changes in 21st century North American forest growth

Predicting long‐term trends in forest growth requires accurate characterisation of how the relationship between forest productivity and climatic stress varies across climatic regimes. Using a network of over two million tree‐ring observations spanning North America and a space‐for‐time substitution methodology, we forecast climate impacts on future forest growth. We explored differing scenarios of increased water‐use efficiency (WUE) due to CO₂‐fertilisation, which we simulated as increased effective precipitation. In our forecasts: (1) climate change negatively impacted forest growth rates in the interior west and positively impacted forest growth along the western, southeastern and northeastern coasts; (2) shifting climate sensitivities offset positive effects of warming on high‐latitude forests, leaving no evidence for continued ‘boreal greening’; and (3) it took a 72% WUE enhancement to compensate for continentally averaged growth declines under RCP 8.5. Our results highlight the importance of locally adapted forest management strategies to handle regional differences in growth responses to climate change.     

The response of mammals to forest fire and timber harvest in the North American boreal forest

1. This paper reviews and compares the effects of forest fire and timber harvest on mammalian abundance and diversity, throughout successional time in the boreal forest of North America. 2. Temporal trends in mammal abundance and diversity are generally similar for both harvested and burned stands, with some differences occurring in the initiation stage (0–10 years post disturbance). 3. Small mammals and ungulates are most abundant immediately post disturbance, and decrease as stands age. Lynxes and hares utilize mid-successional stands, but are rare in young and old stands. Bats, arboreal sciurids and mustelids increase in abundance with stand age, and are most abundant in old growth. 4. Substantial gaps in the data exist for carnivores; the response of these species to fire and harvest requires research, as predator–prey interactions can affect mammal community structure in both early and late successional stages. 5. The lack of explicit treatment of in-stand forest structure post disturbance, in the reviewed literature made comparisons difficult. Where forest structure was considered, the presence of downed woody material, live residual trees and standing dead wood were shown to facilitate convergence of mammal communities to a pre-disturbance state for both disturbance types. 6. Mammalian assemblages differed considerably between successional stages, emphasizing the importance of maintaining stands of each successional stage on the landscape when implementing forest management strategies.     

火对森林土壤的影响

林火是林地上自由蔓延的火,主要有计划火烧(控制火烧)和野火2种类型.火烧的严重程度决定了火灾的效果及其影响的持续时间.火能够通过燃烧有机质和改变粘土矿物来影响土壤结构,导致土壤容重增加和持水能力降低.长期来看,火烧引起土壤有机质减少.火烧之后土壤温度和pH值的升高会促进土壤呼吸,而微生物数量的减少、根系的死亡和可分解物质的减少则减弱土壤呼吸.林火消耗了森林的枯落物层,通过氧化、挥发、灰分颗粒对流、淋溶等途径减少了森林土壤的养分含量.火灾对土壤生物有重要影响,火灾引起的高温可直接杀死土壤微生物,同时通过改变土壤的各种理化性质及林地微环境,间接影响土壤微生物生存和群落的组成.火对土壤动物的影响显著小于对土壤微生物的影响,原因是前者具有更高的灵活性来逃避火灾.中低程度火烧主要通过土壤环境变化间接影响土壤酶,而严重火烧则通过高温直接使其变性.今后应开展跨学科的森林火灾长期研究,加强对火烧后土壤动物、微生物和生态系统之间相互关系的研究,注重计划用火、火灾对土壤有机碳、土壤养分库的稳定、土壤斥水性、水土流失强度和生物多样性影响方面的研究.     

Re-examination of the elemental composition of some Caryophyllaceae on North American ultramafic soils

A paper of Kruckeberg et al. (1993) reported the hyperaccumulation of nickel by Arenaria rubella at an ultramafic site at Olivine Bridge in the state of Washington, USA. Several aspects of the publication led the present authors to doubt the reliability of the data, leading to a reinvestigation of the behaviour of this species at this site. Extensive sampling of two species of the Caryophyllaceae, A. rubella and Cerastium arvense, was then carried out at Olivine Bridge in two different years, together with the collection of soil samples. The plant samples were treated in various ways in an attempt to reduce the potential influence of soil contamination. The results of this work indicate that A. rubella and C. arvense both behave in a normal way in respect of their nickel uptake from ultramafic soil. The elements nickel, iron, cobalt, chromium and magnesium have been found at levels only 1/10–1/100 of those reported in the earlier work. The weaknesses in the earlier study have led to a consideration of several aspects of putative nickel hyperaccumulation, especially as applied to species in the Caryophyllaceae. Particularly important are washing procedures, selection of material for analysis, and the use of various criteria for assessing the likelihood of soil contamination of plant samples. A careful re-examination of several other reported cases of nickel hyperaccumulation by members of the Caryophyllaceae is now recommended.     

Phylogeography of the North American red fox: vicariance in Pleistocene forest refugia

Fossil, archaeological, and morphometric data suggest that indigenous red foxes in North America were derived from vicariance in two disjunct refugia during the last glaciation: one in Beringia and one in the contiguous USA. To test this hypothesis, we conducted a phylogeographical analysis of the North American red fox within its presettlement range. We sequenced portions of the mitochondrial cytochrome b (354 bp) gene and D-loop (342 bp) from 220 historical red fox specimens. Phylogenetic analysis of the cytochrome b gene produced two clades that diverged c . 400 000 years before present ( bp ): a Holarctic and a Nearctic clade. D-loop analyses of the Nearctic clade indicated three distinct subclades (≥ 99% Bayesian posterior probability); two that were more recently derived (rho estimate c . 20 000 bp ) and were restricted to the southwestern mountains and the eastern portion of North America, and one that was older (rho estimate c . 45 000 bp ) and more widespread in North America. Populations that migrated north from the southern refugium following deglaciation were derived from the colonization of North America during or prior to the Illinoian glaciation (300 000–130 000 bp ), whereas populations that migrated south from the northern refugium represent a more recent colonization event during the Wisconsin glaciation (100 000–10 000 bp ). Our findings indicate that Nearctic clade red foxes are phylogenetically distinct from their Holarctic counterparts, and reflect long-term isolation in two disjunct forest refugia during the Pleistocene. The montane lineage, which includes endangered populations, may be ecologically and evolutionarily distinct.     

Effect of drought on Bradyrhizobium japonicum populations in Midwest soils

Background and aims

Bradyrhizobium japonicum and soybean (Glycine max (L.) Merr.) form a symbiotic association which allows for biological nitrogen fixation (BNF) to help meet the nitrogen (N) requirement of soybean plants. Rhizobial inoculants are not always used in soybean production in the Midwestern USA because of high naturalized soil populations, but drought conditions experienced in the region during the 2012 growing season may have led to a decline in numbers resulting in the need for inoculation the following growing season. Therefore, the effect of drought on B. japonicum population size was investigated in this study.

Methods

Drought conditions, 8 weeks long or 4 weeks long preceded (STOP) or followed (START) by 4 weeks of normal watering, were simulated in two contrasting soil types in a greenhouse setting with soybeans as host plants. Drought conditions were monitored by measuring water content. Population size of B. japonicum was quantified using quantitative real-time polymerase chain reaction (qPCR) and most probable number (MPN) methods and compared to population from non-drought treatment.

Results

Using both quantification methods, the response of B. japonicum to drought treatments was minimal.

Conclusions

Drought conditions 4 to 8 weeks long did not reduce B. japonicum population size to levels which would affect soybean growth and development.     

Parallel evolution of dwarf ecotypes in the forest tree Eucalyptus globulus

Three small populations of a dwarf ecotype of the forest tree Eucalyptus globulus are found on exposed granite headlands in south-eastern Australia. These populations are separated by at least 100 km. Here, we used 12 nuclear microsatellites and a chloroplast DNA marker to investigate the genetic affinities of the dwarf populations to one another and to their nearest populations of tall E. globulus. Cape Tourville was studied in greater detail to assess the processes enabling the maintenance of distinct ecotypes in close geographical proximity. The three dwarf populations were not related to one another and were more closely related to adjacent tall trees than to one another. At Cape Tourville the dwarf and tall ecotypes were significantly differentiated in microsatellites and in chloroplast DNA. The dwarf and tall populations differed in flowering time and no evidence of pollen dispersal from the more extensive tall to the dwarf population was found. The three dwarf populations have evolved in parallel from the local tall ecotypes. This study shows that small marginal populations of eucalypts are capable of developing reproductive isolation from nearby larger populations through differences in flowering time and/or minor spatial separation, making parapatric speciation possible.     

Nitrification in coniferous forest soils

Net nitrification rates tend to be low or negligible in the forest floor of many coniferous forests of North-East Scotland. The most likely process controls are substrate availability, pH, allelopathy, water potential, nutrient status and temperature. These are discussed in relation to field and laboratory studies of net and potential rates of nitrification.Fungi make up by far the largest part of the nitrifier community in the coniferous forest floor. Very little is known about the distribution and activity of autotrophs in these systems, although it is certain that in vitro evidence suggesting autotrophs cannot nitrify at pH levels characteristic of coniferous forest soils is unrealistic.Because of the metabolic diversity of nitrifying fungi, a variety of organic and inorganic nitrification pathways may exist in coniferous forests. The possible involvement of free radicles in fungal nitrification in coniferous forest soils is also suggested.A complete understanding of nitrification in coniferous forest soils can only result from field characterisation of N flux such as through the use of ¹⁵N. This must be combined with ecophysiological characterisation of the organisms involved in order that the complexity of nitrification in coniferous forest soils can be resolved.     

Postfire response of North American boreal forest net primary productivity analyzed with satellite observations

Fire is a major disturbance in the boreal forest, and has been shown to release significant amounts of carbon (C) to the atmosphere through combustion. However, less is known about the effects on ecosystems following fire, which include reduced productivity and changes in decomposition in the decade immediately following the disturbance. In this study, we assessed the impact of fire on net primary productivity (NPP) in the North American boreal forest using a 17‐year record of satellite NDVI observations at 8‐ km spatial resolution together with a light‐use efficiency model. We identified 61 fire scars in the satellite observations using digitized fire burn perimeters from a database of large fires. We studied the postfire response of NPP by analyzing the most impacted pixel within each burned area. NPP decreased in the year following the fire by 60–260 g C m^?2 yr^?1 (30–80%). By comparing pre‐ and postfire observations, we estimated a mean NPP recovery period for boreal forests of about 9 years, with substantial variability among fires. We incorporated this behavior into a carbon cycle model simulation to demonstrate these effects on net ecosystem production. The disturbance resulted in a release of C to the atmosphere during the first 8 years, followed by a small, but long‐lived, sink lasting 150 years. Postfire net emissions were three times as large as from a model run without changing NPP. However, only small differences in the C cycle occurred between runs after 8 years due to the rapid recovery of NPP. We conclude by discussing the effects of fire on the long‐term continental trends in satellite NDVI observed across boreal North America during the 1980s and 1990s.