ALLELOPATHIC GROWTH INHIBITION OF NODULATED AND UNNODULATED ALNUS CRISPA SEEDLINGS BY POPULUS BALSAMIFERA

Water extracts of fresh leaves, buds, and leaf litter of Populus balsamifera (balsam poplar) were tested at different dilutions for allelopathic effects on nodulation, nitrogenase activity and growth of nodulated green alder (Alnus crispa) seedlings, and on growth of unnodulated green alder seedlings. All extracts inhibited height growth, root elongation and dry weight increment of nodulated and unnodulated green alder seedlings to some degree during a 2-month experiment. Foliar nitrogen content of both nodulated and unnodulated seedlings was significantly lower in extract treated plants than in controls. Growth inhibition was about 25% less in nodulated than in unnodulated seedlings. The number of nodules per plant in seedlings treated with any balsam poplar extract was only 51% that of control plants. Acetylene reduction by seedlings treated with bud and leaf litter extracts indicated a decrease of 62% compared to controls. Growth inhibition was not mediated by pH or osmotic effects of the extracts. The possible ecological significance is discussed.     

Effect of leaf litter characteristics on leaf conditioning and on consumption by Gammarus pulex

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Flow-substrate interactions create and mediate leaf litter resource patches in streams

1. The roles that streambed geometry, channel morphology, and water velocity play in the retention and subsequent breakdown of leaf litter in small streams were examined by conducting a series of field and laboratory experiments. 2. In the first experiment, conditioned red alder (Alnus rubra Bongard) leaves were released individually in three riffles and three pools in a second‐order stream. The transport distance of each leaf was measured. Several channel and streambed variables were measured at each leaf settlement location and compared with a similar number of measurements taken at regular intervals along streambed transects (‘reference locations’). Channel features (such as water depth) and substrate variables (including stone height, stone height‐to‐width ratio, and relative protrusion) were the most important factors in leaf retention. 3. In the second experiment, the role of settlement location and reach type in determining the rate of leaf litter breakdown was examined by placing individual conditioned red alder leaves in exposed and sheltered locations (on the upper and lower edges of the upstream face of streambed stones, respectively) in riffle and pool habitats. After 10 days, percent mass remaining of each leaf was measured. Generally, leaves broke down faster in pools than in riffles. However, the role of exposure in breakdown rate differed between reach types (exposed pool > sheltered pool > sheltered riffle > exposed riffle). 4. In the third experiment, the importance of substrate geometry on leaf litter retention was examined by individually releasing artificial leaves upstream of a series of substrate models of varying shape. Substrates with high‐angle upstream faces (were vertical or close to vertical), and that had high aspect ratios (were tall relative to their width), retained leaves more effectively. 5. These results show that streambed morphology is an important factor in leaf litter retention and breakdown. Interactions between substrate and flow characteristics lead to the creation of detrital resource patchiness, and may partition leaf litter inputs between riffles and pools in streams at baseflow conditions.     

Nitrogen cycling in dense plantings of hybrid poplar and black alder

Summary Nitrogen cycling was studied during the third growing season in pure and mixed plantings (33×33 cm spacing) of hybrid poplar and black alder in southeastern Canada. After 3 years, hybrid poplar growth and N content of living tissues in a plot and of individual hybrid poplar plants increased with the proportion of black alder in a planting. No differences were detected among N contents of individual alder plants regardless of plot treatment. Black alder allocated a larger portion of its N to roots than hybrid poplar. Symbiotic nitrogen fixation was estimated to account for 80% of the nitrogen in aboveground alder tissues in the pure treatment using natural¹⁵N dilution. N return in leaf litter was estimated to be 70kg ha^–1 in the pure alder treatment and decreased to a minimum of 20 kg ha^–1 in the pure hybrid poplar plots. No difference was detected among treatments for throughfall N content. Nitrogen concentration in roots and leaf litterfall of black alder was higher than hybrid poplar. Significant soil N accretion occurred in mixed plantings containing two alders to one poplar and pure black alder plantings. Nitrogen availability (NO₃–N) increased with the amount of black alder in a plot. Results suggest that the early increase in nitrogen accumulation of hybrid poplar in mixed treatments can be attributed to an increase of total soil N availability resulting from the input of large amounts of N from easily mineralizable alder tissue.     

不同水分条件下杨树-玉米复合系统凋落物分解特性

为探讨水分变化对农林复合生态系统凋落物分解特性的影响,以河西走廊杨树（Populus）-玉米（Zea mays）凋落物为研究对象,设置正常水分（9200 m³/hm²,对照）,轻度干旱胁迫（减少15%,7800 m³/hm²）,中度干旱胁迫（减少30%,6400 m³/hm²）3种不同水分处理条件,采用分解袋法研究了不同水分条件下杨树叶和玉米秸秆的质量残留率、分解速率和养分含量变化特征。结果表明：（1）随着干旱胁迫的加剧,两种凋落物的质量残留率均增加,而分解速率降低。经过164 d的分解后,杨树叶和玉米秸秆的质量残留率分别为70.43%-77.49%、63.55%-68.29%。分析表明：水分和时间对各类型凋落物的质量残留率均有极显著的影响（P<0.001）,但二者的交互作用不显著（P>0.05）;干旱胁迫显著降低了玉米秸秆的分解速率,但杨树叶的分解速率却只是在中度干旱胁迫下显著降低（P<0.05）。对于不同类型凋落物而言,分解速率表现为玉米秸秆>杨树叶。（2）两种类型凋落物的氮（N）残留率在分解过程中表现为降低的趋势,但随着干旱程度的加大,N的残留率增加,表明水分抑制了N的释放过程。分解164d后,同一类型凋落物不同水分条件下的N残留率均存在显著差异。对于同一水分条件下不同凋落物而言,玉米秸秆的N残留率最低,而杨树叶最高。总的来说,水分降低对干旱区农林复合系统内凋落物的分解和氮元素含量具有显著的抑制作用。     

Decomposition of Eucalyptus globulus leaves and three native leaf species (Alnus glutinosa, Castanea sativa and Quercus faginea) in a Portuguese low order stream

Leaf decomposition of the exotic evergreen Eucalyptus globulus (eucalyptus), and three native deciduous tree species, Alnus glutinosa (alder), Castanea sativa (chestnut) and Quercus faginea (oak), was compared in a second order stream in Central Portugal. Changes in dry weight, nitrogen and polyphenolic compounds and microbial colonization were periodically assessed for three months.Negative exponential curves fit the leaf weight loss with time for all leaf species. Mass loss rate was in the order alder (K = 0.0161) > chestnut (K = 0.0079) > eucalyptus (K = 0.0068) > oak (K = 0.0037). Microbial colonization followed the same pattern as breakdown rates. Evidence of fungal colonization was observed in alder after 3 days in the stream, whereas it took 21 days in oak leaves to have fungal colonization. Fungal diversity was leaf species-dependent and increased with time. In all cases, percent nitrogen per unit leaf weight increased, at least, at the initial stages of decay while soluble polyphenolics (expressed as percentage per unit leaf weight) decreased rapidly in the first month of leaves immersion.Intrinsic factors such as nitrogen and polyphenolic content may explain differences in leaf decomposition. The possible incorporation of eucalyptus litter into secondary production in a reasonable time span is suggested, although community balance and structure might be affected by differences in allochthonous patterns determined by eucalyptus monocultures.     

The Association of the Stream Caddisfly Lasiocephala basalis (Kol.) (Trichoptera: Lepidostomatidae) with Wood

A population of the lepidostomatid caddisfly, Lasiocephala basalis, was studied in a third order stream, the Ohe, in Hesse, Germany. L. basalis showed an intimate association with wood debris in all of its aquatic life stages. Larvae exhibited a significant preference for organic substrates such as woody debris, leaves, and submerged alder roots. Gut content analysis revealed leaf material and wood fragments as the dominant larval food items classifying larvae as facultative xylophagous. Larval dominance on wood during spring corresponded to minimal leaf litter abundance. In winter, during periods of high discharge, the most important larval habitat was the submerged alder roots at the stream margin, thus acting as flow refugia. Most pupal cases were attached to submerged logs and roots. In addition, most of the egg masses were found on these two substrates. The role of L. basalis larvae in wood degradation is discussed.     

杨树和桤木落叶混合分解对土壤微生物生物量的影响

通过室内培养,研究了杨树和江南桤木落叶混合分解过程中两种落叶的混合比例及落叶添加方式对土壤微生物生物量的影响.结果表明:落叶混合比例对土壤微生物生物量碳(MBC)、氮(MBN)均有显著的影响.培养30 d,江南桤木落叶比例为50％以上的土壤MBC和MBN显著高于纯杨树落叶处理及对照;75 d时,江南桤木落叶比例≥40％的土壤MBC和≥30％的土壤MBN均显著高于纯杨树落叶处理及对照;135 d后,江南桤木落叶比例≥20％的土壤MBC和≥40％的土壤MBN均显著高于纯杨树落叶处理及对照.不同混合比例的土壤MBC/MBN无显著差异,总体呈前期增长后期下降的变化趋势.杨树和江南桤木落叶混合分解对土壤MBC和MBN有显著的协同促进作用.在整个培养过程中,落叶添加方式对土壤MBC、MBN和MBC/MBN无显著影响.     

Influence of leaf litter type on the chemical evolution of a soil parent material (sandstone)

The influence of leaves of Quercus suber L. and Eucalyptus globulus Labill. and needles of Pinus pinaster Ait. on a sandstone substrate was assessed through lysimetric studies during a ten-year period at a site in Central Portugal. The decomposition rate of Q. suber leaf litter was similar to that of E. globulus and higher than that of P. pinaster needle litter. The proportion of nitrogen released from the Q. suber leaf litter was higher than that lost from the other organic species. Such a release was proportional to the initial nitrogen content in the substrates. The concentrations of both NH₄-N and NO₃-N were much higher in leachates collected under Q. suber leaf litter than in those collected under the other organic substrates. A similar trend was found in the leachates collected under the mineral substrate influenced by the studied organic substrates. The leachate concentrations of mineral N (especially NO₃-N) were higher from the mineral substrate under Q. suber leaf litter than from this organic substrate itself. The mineral substrate under leaf litter of E. globulus or needle litter of P. pinaster showed an increase in exchangeable base cations and pH values, and a decrease in extractable Al. Conversely, in the substrate with Q. suber leaf litter there was only a slight increase in exchangeable base cations and pH values, and a decrease in extractable Al. These results combined with those obtained in soils under E. globulus plantations indicate that changes found in these soils are due to soil and forest management practices rather than to the decomposition process of the respective of leaf litter.     

Impacts of invasive riparian knotweed on litter decomposition,aquatic fungi,and macroinvertebrates

Bohemian knotweed (Polygonum × bohemicum), the hybrid between Japanese and giant knotweed, is the most common invasive knotweed species in western North America and the most difficult to control. Invasive knotweed congeners spread aggressively along streams and establish dense monotypic stands, reducing riparian plant species diversity. Allochthonous organic matter inputs from riparian plants are an important source of energy and nutrients for organisms in small streams. However, little information exists concerning the influence of knotweed on stream processes. This study examines the quality of Bohemian knotweed leaves compared to native red alder and black cottonwood leaves, along with leaf-associated fungal biomass accumulation, macroinvertebrate communities, and decay rates from three forested streams in western Washington State. Senesced knotweed leaves were lower in nitrogen and phosphorus, and higher in cellulose, fiber, and lignin content than alder leaves, but were more similar to cottonwood leaves. Fungal biomass differed among species and changed over time. Macroinvertebrate shredders collected from leaf packs after 31 days were proportionately more abundant on alder leaves than knotweed and cottonwood. Decay rates were not significantly different among leaf species, but during the first 31 days alder broke down faster than knotweed. After 56 days, all of the leaf packs were mostly decomposed. Overall, these findings do not show major discrepancies between leaf species except those related to initial litter structural and chemical quality. However, changes in the timing and quantity of litter inputs are also important factors to be considered in understanding the impact of invasive knotweed on stream ecosystem processes.     

长白山白桦(Betula platyphlla)纯林和白桦山杨 (Populus davidiana)混交林凋落物的分解

采用交互分解实验,研究长白山白桦叶片和白桦、山杨与水曲柳混合叶片在白桦纯林和白桦山杨混交林内的分解过程。两年的分解实验结果表明,两种类型叶片均存在一个快速分解阶段和一个慢速分解阶段,森林类型和凋落物类型对凋落物分解率的影响在快速分解阶段不显著而在慢速分解阶段显著;混交林内的环境促进了凋落物分解和养分元素释放;在同一林型内,底物质量高的混合叶片其分解率和养分元素释放率均大于底物质量低的白桦叶片;凋落物的底物质量在一定程度上可以抵消森林类型对凋落物分解的影响;白桦山杨混交林混合叶片分解速率和养分元素释放率要显著大于白桦纯林内的白桦叶片,说明白桦山杨混交林的物质循环速度和养分元素供应能力要显著大于白桦纯林。     

Consumption of leaf detritus by a stream shredder: Influence of tree species and nutrient status

Four species of riparian vegetation (alder, birch, willow and poplar) were fertilized with nitrogen, phosphorus, nitrogen + phosphorus, or no fertilizer (control). The resulting leaf detritus (leached but not microbially colonized) was offered to a stream shredder, Hydatophylax variabilis (Trichoptera: Limnephilidae). In one experiment, shredder consumption of leaf detritus from different nutrient treatments (within tree species) was compared, and in a second experiment, consumption of different tree species (within nutrient treatments) was compared. Larvae preferred leaf detritus from nitrogen + phosphorus treatments (except in poplar where nitrogen treatment was preferred). Alder was preferred over other tree species for all treatments. Chemical and physical analyses of leaf litter showed differences between tree species and nutrient treatments in nutrient content, tannins and leaf toughness. Leaf consumption by larvae was positively associated with nitrogen content and negatively associated with condensed tannin content. Species composition and nutrient status of riparian vegetation may strongly influence detrital food webs in streams.     

Comparisons of the Colonisation by Invertebrates of Three Species of Wood,Alder Leaves,and Plastic “Leaves” in a Temperate Stream

Small woody debris in streams is abundant, and may be a food source or may provide a substrate on which other food sources such as biofilm may develop, both of which may be significant to invertebrates in times of food scarcity. We examined patterns of invertebrate colonisation of small woody debris (veneers of red alder, Douglas‐fir, and western red cedar), red alder leaves, and plastic (as an inert substrate to mimic leaves). Invertebrate colonisation was high on alder leaves, but low on wood substrates and plastic, controlling for the available surface area. Detritivorous invertebrates had significantly higher colonisation rates of alder leaves versus the other four substrates, whereas predators and collectors did not (consistent with their use of these as substrates and not food). All wood decreased in mass by <15% and leaves by ∼50% over the 75 days of the experiment. For all taxa tested, there was no significant difference in their colonisation of the wood veneers versus the plastic sheets. These results suggest that wood was not directly used by these invertebrates as a food source, or that there could be similar biofilm development on the surfaces of these substrates. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Leaf litter quality drives the feeding by invertebrate shredders in tropical streams

Amazon and Cerrado‐forested streams show natural fluctuations in leaf litter quantity along the time and space, suggesting a change on litter quality input. These natural fluctuations of leaf litter have repercussion on the organic matter cycling and consequently effects on leaf decomposition in forested streams. The effects of the quantity of leaf litter with contrasting traits on consumption by larvae of shredder insects from biomes with different organic matter dynamics have still been an understudied question. The Trichoptera Phylloicus spp. is a typical shredder in tropical headwater streams and keep an important role in leaf litter decomposition. Here, we assessed the consumption by shredder Phylloicus spp., from Amazonia and Cerrado biomes, on higher (Maprounea guianensis) and lower quality leaves (Inga laurina) in different proportions and quantities. Experiments were performed concomitantly in microcosms approaches, simulating Cerrado and Amazonian streams. Higher leaf consumption occurred in Cerrado microcosms. Litter quantity influenced negatively leaf consumption by shredders in Cerrado, in opposition to Amazonia, where consumption was not affected by leaf quantity. In both sites, we observed higher consumption by shredders in treatment with only M. guianensis and no difference between other treatments with mixture of leaves. In treatment with litter of I. laurina, we noted the use of substrate for case building (due to the higher leaf toughness), affecting the fragmentation process. Therefore, our results indicate that leaf litter quality drives the preference of consumption by Phylloicus larvae in Cerrado and Amazonia streams.     

Elevated temperature may intensify the positive effects of nutrients on microbial decomposition in streams

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Leaf water relations of black alder [Alnus glutinosa (L.) Gaertn.] growing at neighbouring sites with different water regimes

 The tree species black alder [Alnus glutinosa (L.) Gaertn.] typically inhabits wet sites in central Europe but is also successful on well drained soils. To test the physiological adjustment of the species in situ, conductances, transpiration rates and water potentials (Scholander pressure chamber) of black alder leaves were investigated at two neighbouring sites with different water regimes: alder trees at an occasionally water logged alder forest and alder shrubs in a nearby, much drier hedgerow. Additional experiments with alder cuttings in nutrient culture showed that leaf conductances and gas exchange were both strongly influenced by the substrate water potential. In situ however, there was little spatial variability within the different parts of a crown and we found that physiological regulation at leaf level was hardly influenced by different site water regimes or different tree sizes. Diurnal courses of leaf water relations as well as their regulation at the leaf level (e.g. the hyperbolic relationship between conductances and ΔW) were strikingly similar at both sites. Leaf water potential in black alder was shown to be a consequence of immediate transpiration rates, which were high in comparison to other tree species (up to 4 mmol H₂O m^–2 s^–1), rather than the water potentials being a factor that influenced conductance and, therefore, transpiration. The always high leaf conductances and consequent high transpiration rates are interpreted as a strategy to maximise productivity through low stomatal limitation at sites where water supply is usually not limited. However, at the same time this behaviour restricts black alder to sites where at least the deep-going roots can exploit water. Received: 10 September 1998 / Accepted: 12 January 1999     

Variety of responses of plant phenolic concentration to CO2 enrichment

Leaf area index (LAI) of a stand of adult black alder trees(Alnus glutinosa L., Gaertn.) was determined by means of threeindependent methods. (1) The seasonal course of LAI was directlyobtained by counting leaves in situ and adding up their areas,estimated from harvested subsamples of leaves. (2) The seasonalvariation of LAI in the stand was estimated using the Li-CorLAI-2000 PCA in parallel and with this instrument a VegetationArea Index (VAI, projected area of all phyto-elements) was actuallymeasured. (3) Maximum LAI was calculated from leaf litter collectionstaking into account specific leaf area within different layersof the alder crown. Direct LAI estimates (1) and calculationsfrom leaf litter (3) revealed the same figure of maximum LAI(4.8). This LAI was reached in August. The LAI-2000 PCA capturedthe seasonal variation and underestimated, by 11% on average,the LAI obtained directly. Compared with results gained withother broad-leaved tree species the LAI-2000 PCA values foralder were reliable. It is suggested that this is due to thehorizontal homogeneous structure of the main leaf layer. Thisis in the periphery of the crown, where 90% of the light interceptionoccurs. Taking the het-erogeneity into account a satisfactorycompatibility of the three methods applied to the alder standwas achieved. Key words: Alnus glutinosa, leaf area index, in situ counting, LAI-2000 PCA, litter collections     

Leaf area index determination in an alder forest: a comparison of three methods

Leaf area index (LAI) of a stand of adult black alder trees(Alnus glutinosa L., Gaertn.) was determined by means of threeindependent methods. (1) The seasonal course of LAI was directlyobtained by counting leaves in situ and adding up their areas,estimated from harvested subsamples of leaves. (2) The seasonalvariation of LAI in the stand was estimated using the Li-CorLAI-2000 PCA in parallel and with this instrument a VegetationArea Index (VAI, projected area of all phyto-elements) was actuallymeasured. (3) Maximum LAI was calculated from leaf litter collectionstaking into account specific leaf area within different layersof the alder crown. Direct LAI estimates (1) and calculationsfrom leaf litter (3) revealed the same figure of maximum LAI(4.8). This LAI was reached in August. The LAI-2000 PCA capturedthe seasonal variation and underestimated, by 11% on average,the LAI obtained directly. Compared with results gained withother broad-leaved tree species the LAI-2000 PCA values foralder were reliable. It is suggested that this is due to thehorizontal homogeneous structure of the main leaf layer. Thisis in the periphery of the crown, where 90% of the light interceptionoccurs. Taking the het-erogeneity into account a satisfactorycompatibility of the three methods applied to the alder standwas achieved. Key words: Alnus glutinosa, leaf area index, in situ counting, LAI-2000 PCA, litter collections     

Consequences of the colonisation of leaves by fungi and oomycetes for leaf consumption by a gammarid shredder

1. Leaf litter breakdown by shredders in the field is affected by leaf toughness, nutritional value and the presence of secondary compounds such as polyphenols. However, experiments involving the use of single fungal strains have not supported the assumption that leaf parameters determine food selection by shredders perhaps because of a failure to test for high consumption prior to isolation of fungal strains, overrepresentation of hyphomycetes or the potential effects of accompanying bacteria. In this study, we used bacteria‐free, actively growing fungi and oomycetes isolated from conditioned leaf litter for which a shredder had already shown high consumption rates. 2. Black alder (Alnus glutinosa) leaf litter was exposed to the littoral zone of Lake Constance in autumn, and subsamples were analysed for leaf parameters and consumption by Gammarus roeselii under standard conditions at regular intervals. On dates with a high consumption rate of the exposed leaves, 14 single strains of fungi and oomycetes were isolated, freed of bacteria and grown on autoclaved leaves. 3. Six of eight measured leaf parameters of exposed leaves were significantly correlated with Gammarus consumption rates, with high colinearity among leaf parameters hampering the identification of causal relations between leaf parameters and feeding activity. 4. When single strains of fungi and oomycetes were grown on autoclaved leaf litter, toughness of colonised leaves was always lower than in the control and the content of protein, N and P were increased. There were pronounced strain‐specific effects on leaf parameters. Consumption rates also differed significantly, with nine of fourteen isolates consumed at higher rates than controls and none proving to be a deterrent. Protein and polyphenol content were significantly correlated with consumption rates. Oomycete‐colonised leaves were consumed at similar rates but were of lower food quality than fungi‐colonised leaves. 5. We argue that direct strain‐specific attractant or repellent effects of fungi and oomycetes on consumption by G. roeselii are not important. However, we found indirect strain‐specific role operating via effects on leaf parameters.     

Effects of leaf litter and its fungal colonization on the diet of Limnomysis benedeni (Crustacea: Mysida)

The strong invasive freshwater mysid Limnomysis benedeni, a detritivorous–herbivorous feeder, has a preference for small food particles, but also feeds on leaf litter. Here, we tested whether leaf litter consumption by L. benedeni depends on the tree species and leaf conditioning (two types of physical and biological leaf conditioning). At the physical leaf conditioning, L. benedeni was fed with shortly leached or extensively leached leaves of five tree species in laboratory food assays. The mysid consumed shortly leached leaves of Copper Beech, Lombardy Poplar, Common Oak, and especially White Willow, and did not feed on shortly leached Black Alder leaves. The consumption of extensively leached leaves by L. benedeni did not depend on the tree species. Overall, 74% of the variation of the leaf consumption by L. benedeni was explained by the significant interaction of the factors carbon content and polyphenol content of the leaves, caused the feeding strategy of L. benedeni. For the biological leaf conditioning, the mysids consumed to a high degree naturally conditioned leaves, followed by leaves colonized by one of three fungi, but oomycete-colonized leaf litter and autoclaved leaves were consumed at similar low levels. Our results indicate that L. benedeni feeds on different types of conditioned leaves to different extents, and therefore may affect leaf litter degradation in many invaded freshwaters.