Breakdown of four leaf litter species and associated fauna in a Basque Country forested stream

Leaves of 4 species (Quercus robur, Castanea sativa, Corylus avellana and Pinus pinaster) were incubated in a forested headwater stream using bags (1 mm and 5 mm net size) and trays (5 mm net size). The type of treatment influenced litter breakdown rates. Differences in loss rates were noted between Pinus and the deciduous species, and between Quercus and Corylus. Numbers of invertebrates per sample and per gram leaf AFDW generally increased with exposure time. The taxa colonizing the four leaf species were largely identical but colonization was greatly dependent on type of treatment. Taxa richness (Hmax) increased with time. Differences between Hmax and specific diversity (H) were high, due to the importance of Chironomids. Collectors and scrapers, among other functional groups, exhibited the highest densities. Differences in loss rates between fine and coarse treatments were related to differences in shredder biomass, corroborating the importance of this functional group in litter processing.     

Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

Light,nutrients and the fatty acid composition of stream periphyton

1. While the balance of light and nutrients is known to influence the food quality of herbivores by altering algal phosphorus and nitrogen content, the combined effects of light and nutrients on fatty acid synthesis in freshwater periphyton are relatively unknown. In this study, we manipulated light and phosphorus concentration in large, flow‐through experimental streams to examine their effects on both elemental stoichiometry and fatty acid content in periphyton. 2. Two levels of phosphorus (4 and 80 μg L^?1) and three of light (17, 40, 110 μmol photons m^?2 s^?1) were applied in a factorial design in two separate experiments. Diatoms dominated periphyton communities in both experiments, comprising >95% of algal biovolume. Periphyton growth in the streams was simultaneously affected by both resources, even at low rates of supply. 3. Periphyton C/P and C/N ratios increased with light augmentation and decreased with phosphorus enrichment, and consistent with the light : nutrient hypothesis (LNH). Light effects were strongest in streams with low phosphorus concentrations. 4. Periphyton fatty acids reflected the dominance of diatoms : palmitic (16 : 0), palmitoleic (16 : 1ω7) and eicosapentanoic (20 : 5ω3) were the principal saturated (SAFA), monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA), respectively. Linoleic (18 : 2ω6) and linolenic (18 : 3ω3) acids, characteristic of chlorophytes and cyanophytes, were rare, comprising <2% of total fatty acids. 5. Periphyton fatty acid profiles were highly sensitive to light and phosphorus. The proportion of fatty acids comprised by SAFA and MUFA increased with light augmentation and decreased with phosphorus enrichment, whereas PUFA decreased with light and increased with phosphorus. Light effects on fatty acid composition were strongest in phosphorus‐poor streams. PUFA declined with increasing light/phosphorus ratios in the streams, whereas ‘energy’ fatty acids (16 : 0 and 16 : 1) increased. The ratio of SAFA/PUFA was strongly and positively correlated with C/P and C/N ratios. SAFA and MUFA, normalised to dry mass, increased two‐ to threefold with increasing light, while PUFA normalised to dry mass was not significantly affected by light. 6. Similarities in the responses of fatty acids and elemental stoichiometry to light and phosphorus treatments suggested that they were influenced by a common mechanism. Both components of food quality appeared to be sensitive to light‐regulated rates of carbon fixation which, when coupled with insufficient supplies of phosphorus, caused diatom cells to store surplus carbon in SAFA, MUFA and other carbon‐rich compounds that diluted both essential fatty acids and mineral nutrients.     

Breakdown of tussock grass in streams along a gradient of agricultural development in New Zealand

Summary 1. We measured the breakdown rate of tussock grass in 12 New Zealand streams in catchments that provided a gradient of agricultural development. We also examined the microbial and invertebrate communities associated with decomposing tussock litter.
2. Pristine streams in the study had low concentrations of dissolved inorganic nitrogen (<10  μ g L⁻¹) and dissolved reactive phosphate (<3  μ g L⁻¹), whereas streams in the most developed catchments had high concentrations of nitrate (>2500  μ g L⁻¹) and phosphate (35  μ g L⁻¹), as well as greater amounts of suspended sediment and fine sediment covering the streambed.
3. Breakdown rate and microbial respiration were significantly related across the sites, and both were positively related to concentrations of nitrate and phosphate. Fungal biomass, measured as ergosterol, was positively related to microbial respiration and was also higher at sites with higher concentrations of nutrients. Total and shredding invertebrates were most abundant at the sites with high nutrient concentrations, but abundance of shredding invertebrates was not significantly related to breakdown rate. Amphipods were the most common shredding invertebrate at most sites, but probably did not contribute greatly to high rates of breakdown in streams in agricultural catchments.
4. With the exception of one site, nutrients from agricultural development appeared to have larger positive effects on litter breakdown than negative effects from sedimentation. Litter breakdown can serve as a functional measure of ecosystem health in streams, but caution should be exercised when a stress, such as land use, can have both positive (nutrients) and negative (sedimentation) effects.     

Leaf litter breakdown in a Mediterranean stream characterised by travertine precipitation

1. Breakdown of four leaf species ( Platanus orientalis , Populus nigra , Salix atrocinerea , Rubus ulmifolius ) was studied in a Mediterranean second-order stream characterised by abundant travertine precipitation, a history of fire in its catchment, and a recently revegetated alluvial corridor.
2. Compared to breakdown rates reported in the literature for congeneric species, breakdown of the four species was slow (k = 0.0024–0.0069 day⁻¹ for the tree species, and 0.0103 and 0.0111 day⁻¹ for Rubus ), in spite of high water temperatures, indicating that the travertine layer that quickly covered submerged leaves impeded decomposer activity and physical fragmentation losses.
3. Breakdown rates nevertheless differed between leaf species in a predictable manner, suggesting that the observed mass loss was largely due to biological processes.
4. The observed tendency towards increasing leaf nitrogen and phosphorus concentrations during breakdown suggests that microorganisms were actively involved in leaf breakdown; however, this interpretation must be viewed with caution because of potentially confounding effects by nutrients contained in the travertine layer.
5. Leaf breakdown of the three indigenous species was faster than that of the exotic species P. orientalis . Due to the recalcitrance of its leaves, the frequent use of Platanus in revegetation schemes following the destruction of indigenous vegetation by fire, exacerbates the negative effect of travertine precipitation on leaf breakdown and, by extension, energy flow in Mediterranean karst streams.     

Safranine fluorescent staining of wood cell walls

Safranine is an azo dye commonly used for plant microscopy, especially as a stain for lignified tissues such as xylem. Safranine fluorescently labels the wood cell wall, producing green/yellow fluorescence in the secondary cell wall and red/orange fluorescence in the middle lamella (ML) region. We examined the fluorescence behavior of safranine under blue light excitation using a variety of wood- and fiber-based samples of known composition to interpret the observed color differentiation of different cell wall types. We also examined the basis for the differences in fluorescence emission using spectral confocal microscopy to examine lignin-rich and cellulose-rich cell walls including reaction wood and decayed wood compared to normal wood. Our results indicate that lignin-rich cell walls, such as the ML of tracheids, the secondary wall of compression wood tracheids, and wood decayed by brown rot, tend to fluoresce red or orange, while cellulose-rich cell walls such as resin canals, wood decayed by white rot, cotton fibers and the G-layer of tension wood fibers, tend to fluoresce green/yellow. This variation in fluorescence emission seems to be due to factors including an emission shift toward red wavelengths combined with dye quenching at shorter wavelengths in regions with high lignin content. Safranine fluorescence provides a useful way to differentiate lignin-rich and cellulose-rich cell walls without counterstaining as required for bright field microscopy.     

Widespread hybridization within mound‐building wood ants in Southern Finland results in cytonuclear mismatches and potential for sex‐specific hybrid breakdown

Hybridization and gene flow between diverging lineages are increasingly recognized as common evolutionary processes, and their consequences can vary from hybrid breakdown to adaptive introgression. We have previously found a population of wood ant hybrids between Formica aquilonia and F. polyctena that shows antagonistic effects of hybridization: females with introgressed alleles show hybrid vigour, whereas males with the same alleles show hybrid breakdown. Here, we investigate whether hybridization is a general phenomenon in this species pair and analyse 647 worker samples from 16 localities in Finland using microsatellite markers and a 1200‐bp mitochondrial sequence. Our results show that 27 sampled nests contained parental‐like gene pools (six putative F. polyctena and 21 putative F. aquilonia) and all remaining nests (69), from nine localities, contained hybrids of varying degrees. Patterns of genetic variation suggest these hybrids arise from several hybridization events or, instead, have backcrossed to the parental gene pools to varying extents. In contrast to expectations, the mitochondrial haplotypes of the parental species were not randomly distributed among the hybrids. Instead, nests that were closer to parental‐like F. aquilonia for nuclear markers preferentially had F. polyctena's mitochondria and vice versa. This systematic pattern suggests there may be underlying selection favouring cytonuclear mismatch and hybridization. We also found a new hybrid locality with strong genetic differences between the sexes similar to those predicted under antagonistic selection on male and female hybrids. Further studies are needed to determine the selective forces that act on male and female genomes in these newly discovered hybrids.     

A closed artificial stream for conducting experiments requiring a controlled species pool

Experimental testing of ecological theory in the field is often difficult because of the number of factors influencing the test system. Adequate control of relevant variables may require the use of laboratory microcosms for the performance of proper experimental tests. Our efforts to study the effects of species invasion sequence on community structure and ecosystem properties of the developing system, requires that the experimental systems be closed. To meet our demanding experimental requirements, we have designed a closed artificial stream that permits careful control over algal species introductions while minimizing the risks of contamination by unwanted species. In assessing the performance of these streams, we examined the variability among experimental replicates in water chemistry parameters and biomass accumulation. Those water chemistry parameters that were not greatly influenced by biological processes in the streams (Ca, Mg, bicarbonate, and sulfate) had average coefficients of variation less than 10%. Those parameters that were clearly influenced by biological processes (P, N, Fe, and Si) had average coefficients of variation ranging from 13.5%–40.4%. Ash free dry mass had an average coefficient of variation of 23.3%, but chlorophyll a was much more variable at 183%. This pattern of variation indicates that the artificial streams performed well in terms of replicability, but some experimental goals may require carefully conceived designs that adequately partition sources of variation in biologically mediated variables to obtain satisfactory results.     

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

1. The breakdown of oak ( Quercus robur L.), chestnut ( Castanea sativa Miller) and eucalypt ( Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Agüera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days.
2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day^–1 for chestnut leaf litter, from 0.0036 to 0.0051 day^–1 for oak, and from 0.0027 to 0.0158 day^–1 for eucalypt.
3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water.
4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site.
5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.     

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

1. The breakdown of oak ( Quercus robur L.), chestnut ( Castanea sativa Miller) and eucalypt ( Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Agüera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days.
2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day^–1 for chestnut leaf litter, from 0.0036 to 0.0051 day^–1 for oak, and from 0.0027 to 0.0158 day^–1 for eucalypt.
3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water.
4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site.
5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.     

Nutrient concentration-stream discharge relationships during storm events in a first-order stream

The relationship between nutrient element concentration and stream discharge during storm events was studied in a nutrient-rich first order stream. Stream concentrations of NO₃-N, phosphate-P, Ca, and Mg were determined during the course of and following thunderstorms. Nutrient element trajectories were constructed for the elements monitored and some recognizable and reproducible patterns in nutrient concentrations emerged. NO₃-N and phosphate-P generally increased in concentration during the early stages of increasing stream discharge resulting in a general clockwise trajectory. Ca and Mg consistently showed decreases in concentration during rising water, but the pattern of the trajectories was less constant. The patterns seen for NO₃-N and phosphate-P suggest surface run-off as their origin while the patterns for Ca and Mg reflect their primary origin in groundwater. The ability to detect these differences from the analysis of nutrient trajectories suggests the use of this technique for determining the source of other elements in streams.     

Structural features of lignin macromolecules extracted with ionic liquid from poplar wood

1-Ethyl-3-methylimidazolium acetate ([Emim][CH?COO]) was used for the extraction of lignin from poplar wood (Populus albaglandulosa), which was called to ionic liquid lignin (ILL) and structural features of ILL were compared with the corresponding milled wood lignin (MWL). Yields of ILL and MWL were 5.8±0.3% and 4.4±0.4%, respectively. The maximum decomposition rate (V(M)) and temperature (T(M)) corresponding to V(M) were 0.25%/ °C and 308.2 °C for ILL and 0.30%/ °C and 381.3 °C for MWL. The amounts of functional groups (OMe and phenolic OH) appeared to be similar for both lignins; approximately 15.5% and 6.7% for ILL and 14.4% and 6.3% for MWL. However, the weight average molecular weight (M(w)) of ILL (6347 Da) was determined to be 2/3-fold of that of MWL (10,002 Da) and polydispersity index (PDI: M(w)/M(n)) suggested that the lignin fragments were more uniform in the ILL (PDI 1.62) than in the MWL (PDI 2.64).     

Traits mediate drought effects on wood carbon fluxes

CO₂ fluxes from wood decomposition represent an important source of carbon from forest ecosystems to the atmosphere, which are determined by both wood traits and climate influencing the metabolic rates of decomposers. Previous studies have quantified the effects of moisture and temperature on wood decomposition, but these effects were not separated from the potential influence of wood traits. Indeed, it is not well understood how traits and climate interact to influence wood CO₂ fluxes. Here, we examined the responses of CO₂ fluxes from dead wood with different traits (angiosperm and gymnosperm) to 0%, 35%, and 70% rainfall reduction across seasonal temperature gradients. Our results showed that drought significantly decreased wood CO₂ fluxes, but its effects varied with both taxonomical group and drought intensity. Drought‐induced reduction in wood CO₂ fluxes was larger in angiosperms than gymnosperms for the 35% rainfall reduction treatment, but there was no significant difference between these groups for the 70% reduction treatment. This is because wood nitrogen density and carbon quality were significantly higher in angiosperms than gymnosperms, yielding a higher moisture sensitivity of wood decomposition. These findings were demonstrated by a significant positive interaction effect between wood nitrogen and moisture on CO₂ fluxes in a structural equation model. Additionally, we ascertained that a constant temperature sensitivity of CO₂ fluxes was independent of wood traits and consistent with previous estimates for extracellular enzyme kinetics. Our results highlight the key role of wood traits in regulating drought responses of wood carbon fluxes. Given that both climate and forest management might extensively modify taxonomic compositions in the future, it is critical for carbon cycle models to account for such interactions between wood traits and climate in driving dynamics of wood decomposition.     

APPLIED ISSUES: Exploring the response of functional indicators of stream health to land‐use gradients

1. Broad‐scale assessment of stream health is often based on correlative relationships between catchment land‐use categories and measurements of stream biota or water chemistry. Few studies have attempted to characterise the response curves that describe how measures of ecosystem function change along gradients of catchment land use, or explored how these responses vary at broad spatial scales. 2. In autumn 2008, we conducted a survey of 84 streams in three bioregions of New Zealand to assess the sensitivity of functional indicators to three land‐use gradients: percentage of native vegetation cover, percentage of impervious cover (IC) and predicted nitrogen (N) concentration. We examined these relationships using general linear models and boosted regression trees to explore monotonic, non‐monotonic and potential threshold components of the response curves. 3. When viewing the responses to individual land‐use gradients, four of five functional indicators were positively correlated with the removal of native vegetation cover and N. In general, weaker and less responsive models were observed for the IC gradient. An analysis of the response to multiple stressors showed δ¹⁵N of primary consumers and gross primary productivity (GPP) to be the most responsive functional indicators to land‐use gradients. The multivariate models identified thresholds for change in the relationship between the functional indicators and all three land‐use gradients. Apparent thresholds were <10%IC, between 40 and 80% loss of native vegetation cover and at 0.5 and 3.2 mg L^?1 N. 4. The strength of regression models and the nature of the response curves suggest that measures of ecosystem function exhibit predictable relationships with land use. Furthermore, the responses of functional indicators varied little among three bioregions. This information provides a strong argument for the inclusion of functional indicators in a holistic assessment of stream health.     

Linking fish trait responses to in‐stream habitat in reconstructed valley‐plugged stream reaches of the Coastal Plain,U.S.A.

Valley‐plug formation is a challenging consequence of stream channelization especially in physiographic regions with highly erodible soils. Upstream channel degradation and incision results in accelerated sediment delivery processes wherein downstream aggradation decreases stream power and creates sand‐clogged channels. Channel reconstruction is now meeting hydrogeomorphic goals related to valley‐plug remediation, yet there exists a need to understand how this practice also facilitates ecological restoration. We evaluated fish trait response to in‐stream habitat conditions in channelized, recently restored, and “least‐disturbed” reference reaches of Coastal Plain streams in West Tennessee. Restored reaches were ecologically similar to channelized reaches, having higher proportions of nest‐guarding omnivores that were correlated with higher percentages of pool habitats and lower wetted width:depth ratios compared to reference reaches. Reference reaches had higher proportions of fast‐water dwelling and specialized insectivores that were correlated with high abundance of large woody debris, high‐wetted width:depth ratios, and low percentages of pool habitats. We conclude that in‐stream habitats in reconstructed channels have yet to promote reach‐scale ecological restoration relative to fish assemblage organization because trait‐habitat associations were not similar to reference conditions. However, our results lend to the development of ecological restoration targets that can be incorporated in future channel reconstruction projects in valley‐plugged, channelized streams of the Coastal Plain.     

Genetic improvement of the chemical composition of Scots pine (Pinus sylvestris L.) juvenile wood for bioenergy production

Chemical composition is one of the key characteristics that determines wood quality and in turn its suitability for different end products and applications. The inclusion of chemical compositional traits in forest tree improvement requires high‐throughput techniques capable of rapid, non‐destructive and cost‐efficient assessment of large‐scale breeding experiments. We tested whether Fourier‐transform infrared (FTIR) spectroscopy, coupled with partial least squares regression, could serve as an alternative to traditional wet chemistry protocols for the determination of the chemical composition of juvenile wood in Scots pine for tree improvement purposes. FTIR spectra were acquired for 1,245 trees selected in two Scots pine (Pinus sylvestris L.) full‐sib progeny tests located in northern Sweden. Predictive models were developed using 70 reference samples with known chemical composition (the proportion of lignin, carbohydrates [cellulose, hemicelluloses and their structural monosaccharides glucose, mannose, xylose, galactose, and arabinose] and extractives). Individual‐tree narrow‐sense heritabilities and additive genetic correlations were estimated for all chemical traits as well as for growth (height and stem diameter) and wood quality traits (density and stiffness). Genetic control of the chemical traits was mostly moderate. Of the major chemical components, highest heritabilities were observed for hemicelluloses (0.43–0.47), intermediate for lignin and extractives (0.30–0.39), and lowest for cellulose (0.20–0.25). Additive genetic correlations among chemical traits were, except for extractives, positive while those between chemical and wood quality traits were negative. In both groups (chemical and wood quality traits), correlations with extractives exhibited opposite signs. Correlations of chemical traits with growth traits were near zero. The best strategy for genetic improvement of Scots pine juvenile wood for bioenergy production is to decrease and stabilize the content of extractives among trees and then focus on increasing the cellulose:lignin ratio.     

The Effect of Macroinvertebrate Exclusion on Leaf Breakdown Rates in a Tropical Headwater Stream1

The role of macroinvertebrates in the process of leaf breakdown is well studied in temperate streams, but less is known about their role in the tropics. We investigated the effect of reducing macroinvertebrate access to leaf material on leaf breakdown rates in a forested headwater stream in the Luquillo Experimental Forest, Puerto Rico. We measured leaf mass loss using fine and coarse mesh bags over 12 weeks for two common riparian species: Cecropia schreberiana (Moraceae) and Dacryodes excelsa (Burseraceae). Coarse mesh allowed freshwater shrimp and other macroinvertebrates to access leaf material, while fine mesh did not. Leaf breakdown rates did not differ between C. schreberiana and D. excelsa in coarse mesh bags (?0.0375/day vs. ?0.0395/day, respectively), but C. schreberiana breakdown was significantly slower than D. excelsa in fine mesh bags (?0.0159/day vs. ?0.0266/day). Additionally, breakdown in fine mesh bags was significantly slower compared to coarse mesh bags for C. schreberiana, but less so for D. excelsa. Breakdown rates for all treatments were fast relative to those in temperate‐zone streams indicating that both macroinvertebrates and macroinvertebrate‐independent processing can strongly influence leaf decomposition in tropical streams. The difference between C. schreberiana and D. excelsa indicates that the effect of macroinvertebrate exclusion can change with leaf type.     

Individual and combined effects of fine sediment and the herbicide glyphosate on benthic macroinvertebrates and stream ecosystem function

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