Effects of burial on leaf litter quality,microbial conditioning and palatability to three shredder taxa

Summary 1. Heterotrophic microorganisms are crucial for mineralising leaf litter and rendering it more palatable to leaf‐shredding invertebrates. A substantial part of leaf litter entering running waters may be buried in the streambed and thus be exposed to the constraining conditions prevailing in the hyporheic zone. The fate of this buried organic matter and particularly the role of microbial conditioning in this habitat remain largely unexplored. 2. The aim of this study was to determine how the location of leaf litter within the streambed (i.e. at the surface or buried), as well as the leaf litter burial history, may affect the leaf‐associated aquatic hyphomycete communities and therefore leaf consumption by invertebrate detritivores. We tested the hypotheses that (i) burial of leaf litter would result in lower decomposition rates associated with changes in microbial assemblages compared with leaf litter at the surface and (ii) altered microbial conditioning of buried leaf litter would lead to decreased quality and palatability to their consumers, translating into lower growth rates of detritivores. 3. These hypotheses were tested experimentally in a second‐order stream where leaf‐associated microbial communities, as well as leaf litter decomposition rates, elemental composition and toughness, were compared across controlled treatments differing by their location within the streambed. We examined the effects of the diverse conditioning treatments on decaying leaf palatability to consumers through feeding trials on three shredder taxa including a freshwater amphipod, of which we also determined the growth rate. 4. Microbial leaf litter decomposition, fungal biomass and sporulation rates were reduced when leaf litter was buried in the hyporheic zone. While the total species richness of fungal assemblages was similar among treatments, the composition of fungal assemblages was affected by leaf litter burial in sediment. 5. Leaf litter burial markedly affected the food quality (especially P content) of leaf material, probably due to the changes in microbial conditioning. Leaf litter palatability to shredders was highest for leaves exposed at the sediment surface and tended to be negatively related to leaf litter toughness and C/P ratio. In addition, burial of leaf litter led to lower amphipod growth rates, which were positively correlated with leaf litter P content. 6. These results emphasise the importance of leaf colonisation by aquatic fungi in the hyporheic zone of headwater streams, where fungal conditioning of leaf litter appears particularly critical for nutrient and energy transfer to higher trophic levels.     

Interactive effects of pH and leaf litter on a shredder, the scirtid beetle, Helodes pulchella, inhabiting tree-holes

1. Scirtid beetles (Coleoptera: Scirtidae) are common inhabitants of water-filled treeholes and interact with other detritivores in a processing chain commensalism. The strength of the commensalism is determined by resource quantity and the organisms involved have different tolerances to low pH. To determine the effects and interactions of resource quantity and pH on one of these leaf-shredding scirtid beetles, Helodes pulchella (Guerin), leaf litter and pH were varied independently in experimental microcosms which mimic treehole habitats.
2. Helodes pulchella growth was affected by both resources and pH in two similar experiments conducted in different years. Scirtid larvae grew more slowly in low resource treatments and also grew more slowly in low pH treatments. Scirtid survival was not affected by pH, but was lower in high resource microcosms in the 1997 experiment. Consumption of leaf litter was much higher in high resource microcosms, even though not all leaf litter was consumed in low leaf litter treatments.
3. Thus, water chemistry and leaf litter quantity could have strong effects on resource availability through effects on these beetle facilitators, in turn affecting other insects living in these detritus-based communities.     

Fungal alteration of the elemental composition of leaf litter affects shredder feeding activity

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The effect of sunlight on leaf litter quality reduces growth of the shredder Klapopteryx kuscheli

• 1 The direct effect of sunlight on the conditioning, breakdown and incorporation of leaf litter in stream food webs has not yet been considered. The aim here was to evaluate the effects of light intensity on the colonization of leaf litter by microorganisms and its resulting quality as food for the stonefly shredder Klapopteryx kuscheli.
• 2 Leaf litter was conditioned for 2 months in an open reach of a second‐order stream in litter bags either exposed to or shaded from direct sunlight. Subsequently, we performed laboratory experiments to test larval consumption, growth, growth efficiency and feeding preference fed on both leaf litter treatments.
• 3 Leaf litter in the unshaded treatment had three times more chlorophyll‐a (Chl‐a) than that in the shaded treatment, 50% lower fungal biomass and similar bacterial abundance. Although larvae did not prefer either food and fed at the same rate on both leaf litter treatments, they grew twice as fast on the shade‐conditioned leaves and attained a two‐fold higher growth efficiency.
• 4 Sunlight can have significant effects on detritus‐based food webs. Riparian modification induced by human activities in forested catchments increases the potential for sunlight to influence detritus dynamics.

     

两种相手蟹对不同红树植物叶片取食的偏好性

大型底栖动物是红树林生态系统中重要的消费者,在红树林的营养物质转化和能量传递过程中具有重要作用。通过设置室内模拟饲喂实验,研究了华南沿海红树林大型底栖动物的常见种--无齿相手蟹(Sesarma dehaani)和双齿相手蟹(S.bidens)对外来红树植物无瓣海桑(Sonneratia apetala)和乡土红树植物秋茄(Kandelia obovata)、桐花树(Aegiceras corniculatum)凋落叶片的取食特性。结果表明,随着淋溶作用的进行,各树种叶片中单宁含量和C/N均逐渐降低,两个数值的变化幅度以秋茄叶片最大,分别达(45.85±7.74)g·kg-1和11.24%,无瓣海桑为(45.15±11.14)g·kg-1和2.43%,桐花树则为(18.75±15.15)g·kg-1和-1.36%。随着叶片中单宁含量和C/N的降低,两种相手蟹的取食量相应地呈逐渐上升的趋势,双齿相手蟹从3.097g到10.079g和无齿相手蟹则从2.480g到8.723g。两种相手蟹种间的取食特性差异不显著(P0.05),但对不同树种的取食上却存在显著差异,其中对桐花树的取食偏好性显著差于无瓣海桑和秋茄(P0.01),这主要与桐花树叶片的高单宁含量和低营养价值有关。研究结论说明大面积引种外来物种无瓣海桑将可能对华南沿海红树林生态系统的食物链结构产生潜在影响,并为红树林的生态恢复和科学造林提供了参考。     

Higher temperature reduces the effects of litter quality on decomposition by aquatic fungi

1. We investigated the effects of riparian plant diversity (species number and identity) and temperature on microbially mediated leaf decomposition by assessing fungal biodiversity, fungal reproduction and leaf mass loss. 2. Leaves of five riparian plant species were first immersed in a stream to allow microbial colonisation and were then exposed, alone or in all possible combinations, at 16 or 24 °C in laboratory microcosms. 3. Fungal biodiversity was reduced by temperature but was not affected by litter diversity. Temperature altered fungal community composition with species of warmer climate, such as Lunulospora curvula, becoming dominant. 4. Fungal reproduction was affected by litter diversity, but not by temperature. Fungal reproduction in leaf mixtures did not differ or was lower than that expected from the weighted sum of fungal sporulation on individual leaf species. At the higher temperature, the negative effect of litter diversity on fungal reproduction decreased with the number of leaf species. 5. Leaf mass loss was affected by the identity of leaf mixtures (i.e. litter quality), but not by leaf species number. This was mainly explained by the negative correlation between leaf decomposition and initial lignin concentration of leaves. 6. At 24 °C, the negative effects of lignin on microbially mediated leaf decomposition diminished, suggesting that higher temperatures may weaken the effects of litter quality on plant litter decomposition in streams. 7. The reduction in the negative effects of lignin at the higher temperature resulted in an increased microbially mediated litter decomposition, which may favour invertebrate‐mediated litter decomposition leading to a depletion of litter stocks in streams.     

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

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Effectors of biotrophic fungi and oomycetes: pathogenicity factors and triggers of host resistance

Many biotrophic fungal and oomycete pathogens share a common infection process involving the formation of haustoria, which penetrate host cell walls and form a close association with plant membranes. Recent studies have identified a class of pathogenicity effector proteins from these pathogens that is transferred into host cells from haustoria during infection. This insight stemmed from the identification of avirulence (Avr) proteins from these pathogens that are recognized by intracellular host resistance (R) proteins. Oomycete effectors contain a conserved translocation motif that directs their uptake into host cells independently of the pathogen, and is shared with the human malaria pathogen. Genome sequence information indicates that oomycetes may express several hundred such host-translocated effectors. Elucidating the transport mechanism of fungal and oomycete effectors and their roles in disease offers new opportunities to understand how these pathogens are able to manipulate host cells to establish a parasitic relationship and to develop new disease-control measures.     

Interspecific hybridization in plant-associated fungi and oomycetes: a review

Fungi (kingdom Mycota) and oomycetes (kingdom Stramenopila, phylum Oomycota) are crucially important in the nutrient cycles of the world. Their interactions with plants sometimes benefit and sometimes act to the detriment of humans. Many fungi establish ecologically vital mutualisms, such as in mycorrhizal fungi that enhance nutrient acquisition, and endophytes that combat insects and other herbivores. Other fungi and many oomycetes are plant pathogens that devastate natural and agricultural populations of plant species. Studies of fungal and oomycete evolution were extraordinarily difficult until the advent of molecular phylogenetics. Over the past decade, researchers applying these new tools to fungi and oomycetes have made astounding new discoveries, among which is the potential for interspecific hybridization. Consequences of hybridization among pathogens include adaptation to new niches such as new host species, and increased or decreased virulence. Hybrid mutualists may also be better adapted to new hosts and can provide greater or more diverse benefits to host plants.     

Seasonal change of protozoa and micrometazoa in a small pond with leaf litter supply

Protozoa and micrometazoa were observed in a small pond with leaf litter over a 2 year period. The total number of taxa observed in this pond was 83 protozoans and 30 micrometazoans, and the correspondence of each taxon and the habitats of surface water, leaf litter on the pond bottom and sedimented mud was noted. The microfauna in the litter on the pond bottom had a higher diversity than in the water or the sedimented mud. Patterns of seasonal change in the density of organisms — with one or two peaks in a year — were recognized in some taxa of protozoa and micrometazoa. Most peaks of bacterivorous protozoa in the leaf litter appeared in late autumn and spring. This phenomenon is considered to be closely related to the litter decomposition, because reduction of dissolved oxygen and/ or high bacterial density were observed in these two seasons. Nineteen species of protozoa from this pond were cultured in vitro with fallen leaves. From the results of the cultures and the observation of food vacuoles of protozoa in situ, part of the food web of the microfauna in the pond was estimated.     

Lower marine fungi (labyrinthulomycetes) and the decay of mangrove leaf litter

This paper deals with the association of members of the Labyrinthulomycetes (Thraustochytriales and Labyrinthulales) with decaying or decayed leaves at an intertidal mangrove at Morib, Malaysia. Representatives of both orders of these obligately marine unicellular eukaryotes of unresolved taxonomic affinities (Chamberlain & Moss, 1998) were consistently isolated from leaves at all stages of decay from the recently fallen to those in an advanced stage of decay, but not from either green or senescent yellow leaves attached to trees. Baiting experiments using -irradiated leaf discs of Sonneratia and Rhizophora spp. immersed in the aquatic environment of the mangrove, revealed that leaf material was colonised by both labyrinthulids and thraustochytrids within 24 hours of immersion at the test site and these organisms were isolated from the leaf material throughout the 14 day study period. In vitro experiments using axenic cultures of three thraustochytrid genera inoculated onto sterile discs of Sonneratia leaves and incubated for 14 days caused loss of both biomass and structural integrity of the leaf material. Freeze fracture, followed by scanning electron microscopy of leaves inoculated with a thraustochytrid and a strain of Labyrinthula, revealed that penetration of the leaf occured after 4 days and that the thraustochytrid was associated with localised degradation of internal leaf tissues. Cellulase production by an isolate of Schizochytrium aggregatum was detected. The results of all the above investigations are discussed with reference to the role of members of the Labyrinthulomycetes in nutrient cycling in the mangrove.     

Microbial leaf degraders in boreal streams: bringing together stochastic and deterministic regulators of community composition

1. Leaves that fall into the water represent a new habitat for microorganisms to colonise in streams, providing an opportunity to study colonisation and the subsequent regulation of community structure. We explored community composition of bacteria and fungi on decomposing alder leaves in nine streams in central Sweden, and describe their relationship with environmental variables. Succession of the microbial community was studied in one of the streams for 118 days. Microbial community composition was examined by denaturing gradient gel electrophoresis on replicate samples of leaves from each stream. 2. During succession in one stream, maximum taxon richness was reached after 34 days for bacteria and 20 days for fungi respectively. Replicate samples within this stream differed between each other earlier in colonisation, while subsequently such variation among replicate communities was low and remained stable for several weeks. Replicate samples taken from all the nine streams after 34 days of succession showed striking similarities in microbial communities within‐streams, although communities differed more strongly between streams. 3. Canonical analysis of microbial communities and environmental variables revealed that water chemistry had a significant influence on community composition. This influence was superimposed on a statistical relationship between the properties of stream catchments and microbial community composition. 4. The catchment regulates microbial communities in two different ways. It harbours the species pool from which the in‐stream microbial community is drawn and it governs stream chemistry and the composition of organic substrates that further shape the communities. We suggest that there is a random element to colonisation early in succession, whereas other factors such as species interactions, stream chemistry and organic substrate properties, result in a more deterministic regulation of communities during later stages.     

Indirect Effects of Antibiotics in the Aquatic Environment: A Laboratory Study on Detritivore Food Selection Behavior

With regard to possible detrimental effects of human and veterinary antibiotics in the aquatic environment, most research in this field assesses direct impacts of pharmaceuticals on vertebrate or invertebrate test organisms. Another related area of concern is the possible development of antibiotic-resistant bacteria by introducing antimicrobials into the aquatic compartment. However, indirect effects of antibacterials on the trophic cascade have rarely been investigated. This study contributes with an example of how indirect effects of antibiotics on leaf litter decay can be measured and to what extent shredder organisms might be affected. Results from food-selection experiments using Gammarus pulex (Amphipoda) demonstrated clear preferences for leaves conditioned in the absence versus those conditioned in the presence of two antibiotics, oxytetracycline and sulfadiazine. Although this result suggested that microbial and fungal colonisation during leaf litter conditioning might be adversely affected in the antibiotic-treated groups, analyses of total carbon and nitrogen content of conditioned leaf discs did not reveal differences among the treatments.     

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

银合欢叶作氮源栽培食用菌研究

以银合欢叶作氮源全部或部分取代基质中米糠或麸皮,栽培毛木耳、凤尾菇、金针菇、黑木耳、香菇和佛罗里达侧耳6种食用菌。结果表明,银合欢叶能促进菌丝生长。栽培中发现,含10%银合欢叶和17%麸皮的基质能使金针菇获得增产,而含10%或20%银合欢叶的基质可使其它5种食用菌产量提高11.4%~39.1%。     

Dynamics of yeast populations recovered from decaying leaves in a nonpolluted stream: a 2-year study on the effects of leaf litter type and decomposition time

Here we report on the results of a survey of the yeast populations occurring on submerged leaves (alder, eucalyptus and oak) in a natural mountain stream, during different phases of their decomposition and through two consecutive years. Leaf litter mass loss, total yeast counts, Shannon-Weiner index (H'), yeast community structure and physiologic abilities were analyzed to evaluate the dynamics of yeast communities during decay. Seventy-two yeast taxa were recorded, and in all litter types, species of basidiomycetous affinity predominated over ascomycetous ones. Discriminant analysis of presence/absence data (yeast species) showed significant differences both among substrate types (P<0.0026) and with decomposition time (P<0.0001). Carbon and nitrogen source utilization by yeast strains also varied with the substrate (P<0.0001) and decomposition time (P<0.0001). Further conclusions were that: (1) all litter types have in common ubiquitous yeast species, such as Cryptococcus albidus, Debaryomyces hansenii and Rhodotorula glutinis, among the common 20 yeast species; (2) only a few species were dominant, and most species were rare, being recorded once or twice throughout decomposition; and (3) the order of yeast appearance, and their substrate assimilation patterns, strongly suggest a succession phenomenon. Finally, explanations for the distribution patterns and variations in yeast communities are discussed.     

桂西北喀斯特区原生林与次生林鲜叶和凋落叶化学计量特征

研究喀斯特生态脆弱区植物新鲜叶片与凋落叶的元素化学计量学性状,对该地区森林生态系统的恢复与重建具有重要指导意义。在桂西北喀斯特区分别选取了3个原生林群落与3个次生林群落,研究其建群种植物新鲜叶片和凋落叶的C、N、P元素含量及其生态化学计量特征。结果发现,6个群落建群种新鲜叶片C、N、P含量(其平均含量分别为404.3、22.5、1.75 mg/g)均大于凋落叶(平均含量分别为376.5、19.0、1.35 mg/g),鲜叶C:N、C:P、N:P比值(均值分别为17.8、244.9、13.8)均小于凋落叶(均值分别为19.3、315.3、16.3)。6种植物新鲜叶片N、P含量大于凋落叶,而N:P比小于凋落叶,表明喀斯特区植物对N的再吸收率大于P。3个原生林群落建群种鲜叶与凋落叶的平均C、N含量均大于次生林,而P含量则略小于次生林;原生林鲜叶与凋落叶的C:N比均小于次生林,C:P、N:P则大于次生林,推测次生林相对于原生林有更快的生长速率。原生林鲜叶N:P比为13—15之间,次生林鲜叶N:P比为11—12之间,次生林鲜叶与凋落叶的N:P比均小于原生林,说明原生林凋落物分解相对较慢,原生林能相对多的保留养分以供植物吸收,更能适应喀斯特石生环境。植物鲜叶和凋落叶的C:N与N:P比值均呈极显著正相关,说明叶片养分元素间具有共变的特性;叶片N、P含量呈正相关关系,表明植物N:P比具有相对的稳定性,这是高等陆生植物C-N-P元素计量的普遍规律,体现了植物群落对环境的适应。     

落叶收集法测定叶面积指数的快速取样方法

叶面积指数(LAI)是植被冠层结构的一个重要参数,它不仅是许多生态和气候模型的重要输入变量,而且是生态系统动态变化监测的一个重要指标。LAI可通过各种间接和直接的手段来观测,而间接观测的LAI值常常需要直接的观测数据来校验。落叶收集法是一种广泛使用的直接观测LAI的方法,过去的研究还未发现有涉及落叶收集的取样技术及其观测精度的内容。对长白山和北京地区落叶阔叶林的落叶进行了3a的观测,每年一次性收集落叶样品分析,研究结果表明:①不同层次落叶的含水量差异巨大,且落叶含水量的日变化明显。上下层落叶的含水量绝对值差异高达10%以上,日变化绝对值差异高达20%以上。因此,在野外收集落叶样本时,为减小落叶含水量变化所导致的LAI观测误差,应从上到下直到地面进行取样,且尽可能多地收集落叶样本。②在落叶阔叶人工林和天然林里,无论样地的大小(1hm2或30m×30m样地),无论取样单元的大小(1m2或25m2分辨率),林内的LAI分布很不均匀,LAI介于0-15.5(1m2分辨率的1hm2样地)或者2.6-9.1(25m2分辨率的30m×30m样地)。③要准确测定落叶林的LAI,收集落叶的样地面积越大越好,且尽量选择地势平坦的样地。对于1hm2或者30m×30m大小的样地,可随机布设一个10m×10m的小样地来观测,精度分别可达85%、80%。④10m×10m小样地的LAI观测,可将其分为4个相邻的5m×5m小样进行取样。对每个5m×5m小样,快速的取样方法是:Ⅰ.随机布设6个1m2取样,这样取样可以保证在99%概率水平上,100m2、30m×30m和1hm2样地的LAI观测精度分别为90%、75%、70%左右。Ⅱ.随机布设11个1m2取样。可以保证在99%概率水平上,100m2、30m×30m和1hm2样地的LAI观测精度分别为94%、80%、75%左右。     

Preconditioning of leaves by solar radiation and anoxia affects microbial colonisation and rate of leaf mass loss in an intermittent stream

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