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1.
SUMMARY. 1. Despite the widely accepted importance of bacteria and fungi in degrading detritus in aquatic ecosystems there is still very little quantitative information on the abundance and dynamics of these microorganisms. Using epifluorescent microscopy, we measured the biomass of bacteria and fungi during decomposition of three types of leaf detritus. Bacterial production was determined from the rate of incorporation of 3H-thymidine into DNA.
2. The transformation of leaf carbon into dissolved organic carbon and fine particulate organic carbon was followed in order to compare the amounts of leaf material that were converted into these 'end-products' of decomposition versus the amount converted into microbial biomass.
3. The amount of microbial carbon in the leaf-detritus complex never exceeded 5.2% of the total carbon, and fungal biomass was always much greater than bacterial biomass. Despite the greater standing stock of fungi, the rapid turnover of bacteria (doubling about once per day) implies that their role in degrading leaf litter or as a food source for detritivores might be as great as for fungi.
4. Removal of microbial biomass from leaf litter may occur as release of fungal spores and consumption or shedding of bacterial biomass. Fungal spores can be a significant part of the fine particulate organic carbon released from leaf detritus and potentially represent an important food resource for filter-feeding organisms.  相似文献   

2.
As leaves enter woodland streams, they are colonized by both fungi and bacteria. To determine the contribution of each of these microbial groups to the decomposition process, comparisons of fungal and bacterial production are needed. Recently, a new method for estimating fungal production based on rates of [(sup14)C]acetate incorporation into ergosterol was described. Bacterial production in environmental samples has been determined from rates of [(sup3)H]leucine incorporation into protein. In this study, we evaluated conditions necessary to use these methods for estimating fungal and bacterial production associated with leaves decomposing in a stream. During incubation of leaf disks with radiolabeled substrates, aeration increased rates of fungal incorporation but decreased bacterial production. Incorporation of both radiolabeled substrates by microorganisms associated with leaf litter was linear over the time periods examined (2 h for bacteria and 4 h for fungi). Incorporation of radiolabeled substrates present at different concentrations indicated that 400 nM leucine and 5 mM acetate maximized uptake for bacteria and fungi, respectively. Growth rates and rates of acetate incorporation into ergosterol followed similar patterns when fungi were grown on leaf disks in the laboratory. Three species of stream fungi exhibited similar ratios of rates of biomass increase to rates of acetate incorporation into ergosterol, with a mean of 19.3 (mu)g of biomass per nmol of acetate incorporated. Both bacterial and fungal production increased exponentially with increasing temperature. In the stream that we examined, fungal carbon production was 11 to 26 times greater than bacterial carbon production on leaves colonized for 21 days.  相似文献   

3.
1.  1. It has been accepted that aquatic hyphomycetes colonising submerged leaves increase the nutritional value of leaf detritus and suggested that fungal biomass plays a greater role in the growth of shredders than leaf tissue itself. However, it is not clear what proportion of the nutritional needs of shredders is met by fungal biomass.
2.  We fed Pycnopsyche gentilis larvae with tulip poplar ( Liriodendron tulipifera ) leaf discs colonised by the aquatic hyphomycete, Anguillospora filiformis , which had been radiolabelled to quantify the contribution of fungal carbon to the growth of the shredder at different larval developmental stages. Instantaneous growth rates of larvae on this diet were also estimated.
3.  When provided with fungal-colonised leaves (14–16% fungal biomass), the third and the fifth instar larvae of P. gentilis grew at the rates of 0.061 and 0.034 day−1, respectively, but on a diet of sterile leaves, both larval instars lost weight. The incorporation rates of fungal carbon were 31.6 μg C mg−1 AFDM day−1, accounting for 100% of the daily growth rate of the third instar larvae and 8.6 μg C mg−1 AFDM day−1, accounting for 50% of the daily growth rate of the fifth instar larvae.
4.  These results suggest that leaf material colonised by A. filiformis is a high quality food resource for P. gentilis larvae, and that fungal biomass can contribute significantly to the growth of these larvae. Differences in feeding behaviour and digestive physiology may explain the significantly greater assimilation of fungal biomass by the earlier instar than the final instar. To satisfy their nutritional needs the fifth instar larvae would have to assimilate detrital mass that may have been modified by fungal exoenzymes.  相似文献   

4.
5.
The effect of zinc on leaf decomposition by aquatic fungi was studied in microcosms. Alder leaf disks were precolonized for 15 days at the source of the Este River and exposed to different zinc concentrations during 25 days. Leaf mass loss, fungal biomass (based on ergosterol concentration), fungal production (rates of [1-14C]acetate incorporation into ergosterol), sporulation rates, and species richness of aquatic hyphomycetes were determined. At the source of the Este River decomposition of alder leaves was fast and 50% of the initial mass was lost in 25 days. A total of 18 aquatic hyphomycete species were recorded during 42 days of leaf immersion. Articulospora tetracladia was the dominant species, followed by Lunulospora curvula and two unidentified species with sigmoid conidia. Cluster analysis suggested that zinc concentration and exposure time affected the structure of aquatic hyphomycete assemblages, even though richness had not been severely affected. Both zinc concentration and exposure time significantly affected leaf mass loss, fungal production and sporulation, but not fungal biomass. Zinc exposure reduced leaf mass loss, inhibited fungal production and affected fungal reproduction by either stimulating or inhibiting sporulation rates. The results of this work suggested zinc pollution might depress leaf decomposition in streams due to changes in the structure and activity of aquatic fungi.  相似文献   

6.
SUMMARY 1. Decomposition of red maple ( Acer rubrum ) and rhododendron ( Rhododendron maximum ) leaves and activity of associated microorganisms were compared in two reaches of a headwater stream in Coweeta Hydrologic Laboratory, NC, U.S.A. The downstream reach was enriched with ammonium, nitrate, and phosphate whereas the upstream reach was not altered.
2. Decomposition rate, microbial respiration, fungal and bacterial biomass, and the sporulation rate of aquatic hyphomycetes associated with decomposing leaf material were significantly higher for both leaf types in the nutrient-enriched reach. Species richness and community structure of aquatic hyphomycetes also exhibited considerable changes with an increase in the number of fungal codominants in the nutrient-enriched reach.
3. Fungal biomass was one to two orders of magnitude greater than bacterial biomass in both reaches. Changes in microbial respiration rate corresponded to those in fungal biomass and sporulation, suggesting a primary role of fungi in leaf decomposition.
4. Nutrient enrichment increased microbial activity, the proportion of leaf carbon channelled through the microbial compartment and the decomposition rate of leaf litter.  相似文献   

7.
The quantitative significance of bacterial biomass in the nutrition of detritivores remains equivocal. We have used tritiated thymidine to specifically label stable macromolecules in natural assemblages of sediment-associated and detritus-associated bacteria. This material was presented to the isopod (Lirceus sp.) and incorporation of bacterial biomass measured. The isopod incorporated roughly 1 ng bacterial carbon (mg wet wt.)-1 h-1 from leaf discs and about 6 ng mg-1 h-1 from sediment. Calculation of grazing rate from changes in cell counts yields grazing rates from 2.3–17.9 ng C mg-1 h-1. Even the maximum grazing rate, which is an overestimate of C assimilated, represents only 14.7% of C respired by the isopod.  相似文献   

8.
Decomposition of culms (sheaths and stems) of the emergent macrophyte Phragmites australis (common reed) was followed for 16 months in the litter layer of a brackish tidal marsh along the river Scheldt (the Netherlands). Stems and leaf sheaths were separately analyzed for mass loss, litter-associated fungal biomass (ergosterol), nutrient (N and P), and cell wall polymer concentrations (cellulose and lignin). The role of fungal biomass in litter nutrient dynamics was evaluated by estimating nutrient incorporation within the living fungal mass. After 1 year of standing stem decay, substantial fungal colonization was found. This corresponded to an overall fungal biomass of 49 ± 8.7 mg g−1 dry mass. A vertical pattern of fungal colonization on stems in the canopy is suggested. The litter bag experiment showed that mass loss of stems was negligible during the first 6 months, whereas leaf sheaths lost almost 50% of their initial mass during that time. Exponential breakdown rates were −0.0039 ± 0.0004 and −0.0026 ± 0.0003 day−1 for leaf sheaths and stems, respectively (excluding the initial lag period). In contrast to the stem tissue—which had no fungal colonization—leaf sheaths were heavily colonized by fungi (93 ± 10 mg fungal biomass g−1 dry mass) prior to placement in the litter layer. Once being on the sediment surface, 30% of leaf sheath's associated fungal biomass was lost, but ergosterol concentrations recovered the following months. In the stems, fungal biomass increased steadily after an initial lag period to reach a maximal biomass of about 120 mg fungal biomass g−1 dry mass for both plant parts at the end of the experiment. Fungal colonizers are considered to contain an important fraction of nutrients within the decaying plant matter. Fungal N incorporation was estimated to be 64 ± 13 and 102 ± 15% of total available N pool during decomposition for leaf sheaths and stems, respectively. Fungal P incorporation was estimated to be 37 ± 9 and 52 ± 15% of total available P during decomposition for leaf sheaths and stems, respectively. Furthermore, within the stem tissue, fungi are suggested to be active immobilizers of nutrients from the external environment because fungi were often estimated to contain more than 100% of the original nutrient stock.  相似文献   

9.
The contribution of fungi and bacteria to the decomposition of alder leaves was examined at two reference and two polluted sites in the Ave River (northwestern Portugal). Leaf mass loss, microbial production from incorporation rates of radiolabeled compounds into biomolecules, fungal biomass from ergosterol concentration, sporulation rates, and diversity of aquatic hyphomycetes associated with decomposing leaves were determined. The concentrations of organic nutrients and of inorganic nitrogen and phosphorus in the stream water was elevated and increased at downstream sites. Leaf decomposition rates were high (0.013 day(-1) < k < 0.042 day(-1)), and the highest value was estimated at the most downstream polluted site, where maximum values of microbial production and fungal biomass and sporulation were found. The slowest decomposition occurred at the other polluted site, where, along with the nutrient enrichment, the lowest current velocity and dissolved-oxygen concentration in water were observed. At this site, fungal production, biomass, and sporulation were depressed, suggesting that stimulation of fungal activity by increased nutrient concentrations might be offset by other factors. Although bacterial production was higher at polluted sites, fungi accounted for more than 94% of the total microbial net production. Fungal yield coefficients varied from 10.2 to 13.6%, while those of bacteria were less than 1%. The contribution of fungi to overall leaf carbon loss (29.0 to 38.8%) greatly exceeded that of bacteria (4.2 to 13.9%).  相似文献   

10.
1. We compared microbial biomass (bacteria, fungi, algae) and the activity of extracellular enzymes used in the decomposition of organic matter (OM) among different benthic substrata (leaves, coarse and fine substrata) over one hydrological year in a Mediterranean stream.
2. Microbial heterotrophic biomass (bacteria plus fungi) was generally higher than autotrophic biomass (algae), except during short periods of high light availability in the spring and winter. During these periods, sources of OM shifted towards autochthonous sources derived mainly from algae, which was demonstrated by high algal biomass and peptidase activity in benthic communities.
3. Heterotrophic activity peaked in the autumn. Bacterial and fungal biomass increased with the decomposition of cellulose and hemicellulose compounds from leaf material. Later, lignin decomposition was stimulated in fine (sand, gravel) and coarse (rocks, boulders and cobbles) substrata by the accumulation of fine detritus.
4. The Mediterranean summer drought provoked an earlier leaf fall. The resumption of the water flow caused the weathering of riparian soils and subsequently a large increase in dissolved organic carbon and nitrate, which led to growth of bacteria and fungi.  相似文献   

11.
Annual production of leaf-decaying fungi in a woodland stream   总被引:6,自引:0,他引:6  
1. Fungi are thought to be important mediators of energy flow in the detritus-based food webs of woodland streams. However, until recently, quantitative methods to assess their contribution have been lacking. Growth rates of leaf-decaying fungi can be estimated from rates of acetate incorporation into ergosterol which, together with estimates of fungal biomass from ergosterol concentrations, enables calculation of fungal production. In this study, I used this method to estimate total production of leaf-decaying fungi over an annual cycle in a small woodland stream, Walker Branch, Tennessee, U.S.A. To calculate fungal biomass and production on an areal basis, I determined the amount of leaf litter occurring in the stream by sampling transects randomly selected in each of ten 10-m sections every 20–50 days. Subsamples of leaves chosen from five of the transects were used to determine ergosterol concentrations and in situ rates of acetate incorporation into ergosterol. 2. Leaf litter, fungal biomass m–2, and fungal production m–2 were highly seasonal. Leaf litter ranged from 249 g m–2 in November to less than 5 g m–2 during the summer. Fungal biomass as percentage of leaf litter ranged from 4.4 to 8.8% during the year, but on an areal basis ranged from 11 to 13 g m–2 during November to January to 0.25 g m–2 in June, primarily due to the seasonal variation in amount of leaf litter present. Fungal growth rates averaged 2.6% day–1 (0.9–7.0% day–1) during the year. Daily production of leaf-decaying fungi ranged from 0.49 g m–2 in November, when the amount of leaf litter was at its maximum, to 0.006 g m–2 during the summer when the amount of leaf litter was low. Annual production of leaf-decaying fungi was 34 g m–2, with an annual production to biomass ratio (P/B) of 8.2. 3. Fungal spore concentrations in the stream were also seasonal and were correlated with amount of leaf litter m–2 and fungal biomass m–2. Spore concentrations varied between one and four spores ml–1 throughout most of the year, but increased to eighteen spores ml–1 shortly after the greatest amount of leaf litter was present in the stream during November.  相似文献   

12.
The relative contributions of fungi and bacteria to carbon flow from submerged decaying plant litter at different levels of inorganic nutrients (N and P) were studied. We estimated leaf mass loss, fungal and bacterial biomass and production, and microbial respiration and constructed partial carbon budgets for red maple leaf disks precolonized in a stream and then incubated in laboratory microcosms at two levels of nutrients. Patterns of carbon flow for leaf disks colonized with the full microbial assemblage were compared with those colonized by bacteria but in which fungi were greatly reduced by placing leaf disks in colonization chambers sealed with membrane filters to exclude aquatic hyphomycete conidia but not bacterial cells. On leaves colonized by the full microbial assemblage, elevated nutrient concentrations stimulated fungi and bacteria to a similar degree. Peak fungal and bacterial biomass increased by factors of 3.9 and 4.0; cumulative production was 3.9 and 5.1 times higher in the high nutrient in comparison with the low nutrient treatment, respectively. Fungi dominated the total microbial biomass (98.4 to 99.8%) and cumulative production (97.3 and 96.5%), and the fungal yield coefficient exceeded that of bacteria by a factor of 36 and 27 in low- and high-nutrient treatments, respectively. Consequently, the dominant role of fungi in leaf decomposition did not change as a result of nutrient manipulation. Carbon budgets indicated that 8% of leaf carbon loss in the low-nutrient treatment and 17% in the high-nutrient treatment were channeled to microbial (essentially fungal) production. Nutrient enrichment had a positive effect on rate of leaf decomposition only in microcosms with full microbial assemblages. In treatments where fungal colonization was reduced, cumulative bacterial production did not change significantly at either nutrient level and leaf decomposition rate was negatively affected (high nutrients), suggesting that bacterial participation in carbon flow from decaying leaf litter is low regardless of the presence of fungi and nutrient availability. Moreover, 1.5 and 2.3 times higher yield coefficients of bacteria in the reduced fungal treatments at low and high nutrients, respectively (percentage of leaf carbon loss channeled to bacterial production), suggest that bacteria are subjected to strong competition with fungi for resources available in leaf litter.  相似文献   

13.
1. We investigated the resistance of aquatic microbial decomposers to Cu and Zn stress and their ability to recover after release from metal exposure, by examining leaf mass loss, fungal reproduction and microbial biomass and diversity.
2. Alder leaves, colonised in a reference stream, were exposed in microcosms to copper (Cu) or zinc (Zn), alone or in mixtures, with metals added together or sequentially (at day 0 or after 10 days). After 20 days, half of the microcosms were released from metals.
3. Leaf mass loss and fungal reproduction were reduced in most metal treatments, and the structure of fungal and bacterial communities was altered as indicated by identification of conidia and DNA fingerprinting based on denaturing gradient gel electrophoresis. Metals reduced the biomass of bacteria, but not that of fungi.
4. After release from metal stress, the structure of fungal communities became similar to that of control, and a recovery of microbial activity seemed to occur as shown by the lack of differences in leaf mass loss, bacterial biomass and fungal reproduction between control and metal treatments.  相似文献   

14.
The contribution of fungi and bacteria to the decomposition of alder leaves was examined at two reference and two polluted sites in the Ave River (northwestern Portugal). Leaf mass loss, microbial production from incorporation rates of radiolabeled compounds into biomolecules, fungal biomass from ergosterol concentration, sporulation rates, and diversity of aquatic hyphomycetes associated with decomposing leaves were determined. The concentrations of organic nutrients and of inorganic nitrogen and phosphorus in the stream water was elevated and increased at downstream sites. Leaf decomposition rates were high (0.013 day−1 < k < 0.042 day−1), and the highest value was estimated at the most downstream polluted site, where maximum values of microbial production and fungal biomass and sporulation were found. The slowest decomposition occurred at the other polluted site, where, along with the nutrient enrichment, the lowest current velocity and dissolved-oxygen concentration in water were observed. At this site, fungal production, biomass, and sporulation were depressed, suggesting that stimulation of fungal activity by increased nutrient concentrations might be offset by other factors. Although bacterial production was higher at polluted sites, fungi accounted for more than 94% of the total microbial net production. Fungal yield coefficients varied from 10.2 to 13.6%, while those of bacteria were less than 1%. The contribution of fungi to overall leaf carbon loss (29.0 to 38.8%) greatly exceeded that of bacteria (4.2 to 13.9%).  相似文献   

15.
Abstract. 1. In laboratory nests the ants thoroughly licked leaf fragments by a rasping action of their glossa before inoculating the food fungus. The extent of this licking varied with the substrate and was influenced by the thickness of surface leaf waxes and by the chemical attractiveness of the substrate.
2. Microscopy and wettability studies showed that licking removes the wax layer present on leaves. It also removes or inhibits micro-organisms present.
3. Chemically de-waxed and decontaminated leaves and ant-licked leaves proved the best substrates for artificially culturing the ant fungus.
4. The process of substrate preparation circumvents some of the normal defence mechanisms of green plants against fungal attack. The ant fungus thus utilizes nutritionally richer resources of green leaves normally available only to specialized parasitic fungi, rather than the nutritionally inferior shed leaves normally available to saprophytic fungi.  相似文献   

16.
Decaying macrophytes are an important source of carbon and nutrients in fungal and bacterial communities of northern prairie wetlands. Dead macrophytes do not collapse into the water column immediately after death, and decomposition by fungi and bacteria begins while the plants are standing. The seasonal variations in fungal biomass and production on Scirpus lacustris stems, both above and below water, were measured to assess which environmental factors were dominant in affecting these variations in a typical prairie wetland. Fungal biomass and production were measured from early May to November, just prior to freeze-up. Fungal decomposition began and was greatest in the spring despite low water temperatures. The fungal production, as measured by the incorporation of [1-(14)C]acetate into ergosterol, ranged from 1.8 to 376 microg of C g of ash-free dry mass (AFDM)(-1) day(-1), and the biomass, as estimated by using ergosterol, ranged from nondetectable to 5.8 mg of C g of AFDM(-1). There was no significant difference in biomass or production between aerial and submerged portions of Scirpus stems. The water temperature was correlated with fungal production (r = 0.7, P < 0.005) for aerial stem pieces but not for submerged pieces. However, in laboratory experiments water temperature had a measurable effect on both biomass and production in submerged stem pieces. Changes in fungal biomass and productivity on freshly cut green Scirpus stems decaying in the water either exposed to natural solar radiation or protected from UV radiation were monitored over the summer. There was no significant difference in either fungal biomass (P = 0.76) or production (P = 0.96) between the two light treatments. The fungal biomass and rates of production were within the lower range of the values reported elsewhere, probably as a result of the colder climate and perhaps the lower lability of Scirpus stems compared to the labilities of the leaves and different macrophytes examined in other studies performed at lower latitudes.  相似文献   

17.
1. To assess the impact of metal mixtures on microbial decomposition of leaf litter, we exposed leaves previously immersed in a stream to environmentally realistic concentrations of copper (Cu) and zinc (Zn) (three levels), alone and in all possible combinations. The response of the microbial community was monitored after 10, 25 and 40 days of metal exposure by examining leaf mass loss, fungal and bacterial biomass, fungal reproduction and fungal and bacterial diversity.
2. Analysis of microbial diversity, assessed by denaturing gradient gel electrophoresis and identification of fungal spores, indicated that metal exposure altered the structure of fungal and bacterial communities on decomposing leaves.
3. Exposure to metal mixtures or to the highest Cu concentration significantly reduced leaf decomposition rates and fungal reproduction, but not fungal biomass. Bacterial biomass was strongly inhibited by all metal treatments.
4. The effects of Cu and Zn mixtures on microbial decomposition of leaf litter were mostly additive, because observed effects did not differ from those expected as the sum of single metal effects. However, antagonistic effects on bacterial biomass were found in all metal combinations and on fungal reproduction in metal combinations with the highest Cu concentrations, particularly at longer exposure times.  相似文献   

18.
SUMMARY 1. We used a combination of field surveys and field and laboratory experiments to examine the effects of predatory small-mouthed salamander larvae, Ambystoma barbouri (Kraus and Petranka), on the density and microhabitat use of benthic isopods, Lirceus fontinalis (Rafinesque).
2. A survey of four field stations yielded a significant negative relationship between the densities of salamander larvae and isopods.
3. A field experimental manipulation of salamander density confirmed that salamanders reduce isopod density and isopod use of open micro-habitat. Salamanders did not significantly affect the mean or coefficient of variation in isopod size.
4. A laboratory experiment showed that in the absence of refuges for isopods, salamanders were capable of eating large numbers of isopods; small isopods were particularly vulnerable to salamander predation.  相似文献   

19.
The growth response of the hyphae of mycorrhizal fungi has been determined, both when plant and fungus together and when only the fungus was exposed to a temperature change. Two host plant species, Plantago lanceolata and Holcus lanatus, were grown separately in pots inoculated with the mycorrhizal fungus Glomus mosseae at 20/18 degrees C (day/night); half of the pots were then transferred to 12/10 degrees C. Plant and fungal growth were determined at six sequential destructive harvests. A second experiment investigated the direct effect of temperature on the length of the extra-radical mycelium (ERM) of three mycorrhizal fungal species. Growth boxes were divided in two equal compartments by a 20 micro m mesh, allowing only the ERM and not roots to grow into a fungal compartment, which was either heated (+8 degrees C) or kept at ambient temperature. ERM length (LERM) was determined on five sampling dates. Growth of H. lanatus was little affected by temperature, whereas growth of P. lanceolata increased with temperature, and both specific leaf area (SLA) and specific root length (SRL) increased independently of plant size. Percentage of colonized root (LRC) and LERM were positively correlated with temperature when in symbiosis with P. lanceolata, but differences in LRC were a function of plant biomass. Colonization was very low in H. lanatus roots and there was no significant temperature effect. In the fungal compartment LERM increased over time and was greatest for Glomus mosseae. Heating the fungal compartment significantly increased LERM in two of the three species but did not affect LRC. However, it significantly increased SRL of roots in the plant compartment, suggesting that the fungus plays a regulatory role in the growth dynamics of the symbiosis. These temperature responses have implications for modelling carbon dynamics under global climate change.  相似文献   

20.
We studied the effects of epiphytic and endophytic phyllosphere fungi and pathogenic birch rust fungus infection of mountain birch Betula pubescens ssp czerepanovit trees on the larval performance of leaf beetle Phratora potaris We assessed the effects of epiphytic fungi by growing larvae on leaves from trees with manipulated fungal densities We also monitored larval perfonnance and endophytic fungal densities among tree groups classified by herbivory or rust fungus densities The differences in expenmentally manipulated epiphytic fungal densities did not affect larval relative growth rates (RGR) of the species, instead we found significant tree effects Phratora polaris RGR was higher on trees with high level of herbivory than on trees with low herbivory, nevertheless, endophyte densities between these groupings did not differ In the rust fungus expenment, P polarts performance was lowest on trees with low infection compared to no and high infection trees We also did not find correlations among tree-specific endophyte densities and P polaris performance on high and low herbivory trees and trees classified by rust fungus infection Although antagonism among fungi and induction of tree defences cannot be excluded, we suggest that epiphytic and endophytic fungi of mountain birch have negligible effects on P polaris larval performance under natural conditions, probably due to mountain birch variability and a loose ecological connection between mountain birch and its epi- and endophytes Mountain birch and pathogenic birch rust have a more tightly linked relationship, which may also affect insect herbivores Still, leaf properties may play an important role and the effects will depend on the relative timing of the rust infection, herbivore development and changes in leaf quality  相似文献   

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