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1.
1. In semi‐arid climates, seasonally‐flowing streams provide most of the water required for human use, but knowledge of how water extraction affects ecological processes is limited. Predicted alterations in stream flows associated with the impacts of climate change further emphasize the need to understand these processes. Benthic algae are an important base for stream food webs, but we have little knowledge of how algae survive dry periods or respond to altered flow regimes. 2. We sampled 19 streams within the Grampians National Park, south‐eastern Australia and included four components: a survey of different drought refuges (e.g. permanent pools, dry biofilm on stones and dry leaf packs) and associated algal taxa; a survey of algal regrowth on stones after flows recommenced to determine which refuges contributed to regrowth; reciprocal transplant experiments to determine the relative importance of algal drift and regrowth from dry biofilm in recolonization; direct measurement of algal drift to determine taxonomic composition in relation to benthic assemblage composition. 3. Algae showed little specificity for drought refuges but did depend on them; no species were found that were not present in at least one of the perennial pool, dry biofilm or leaf pack refuges. Perennial pools were most closely correlated with the composition of algal assemblages once flows resumed, but the loss or gain of perennial pools that might arise from stream regulation is unlikely to affect the composition of algal regrowth. However, regulated streams were associated with strong increases in algal density in dry biofilm, including increased densities of Cyanobacteria. 4. A model for algal recolonization in seasonally‐flowing streams identified three pathways for algal recolonization (drift‐dependent, dry biofilm‐dependent and contributions from both), depending on whether streams are diatom‐dominated or dominated by filamentous algae. The model predicted the effects of changes to stream flow regimes on benthic algal recolonization and provides a basis for hypotheses testable in streams elsewhere.  相似文献   

2.
Intermittent water flow regimes characterize streams in many world regions, especially those with arid and semiarid climates. During cease to flow conditions, biofilms on streambed sediments may be exposed to desiccation. Environmental conditions and resource availability change with desiccation and may influence biofilm functioning and whole stream ecosystem processes. Rainfall events during the nonflow phase can rehydrate streambed sediments, but the effect of these pulses on biofilm functioning is unclear. This study aimed to analyze the effects of a rehydration event on biofilm functional diversity during the nonflow period in a subtropical Australian stream. Biofilms from three different stream pools on the same reach; one permanently water-covered and the other two differing in their desiccation time were studied. Biofilms initially differed owing to the time they were exposed to dry conditions but rehydration events significantly increased biofilm functional diversity, producing a “reset” effect on the desiccation exposure, as after that bacterial functioning decreased again because of the new dry conditions. The observed rapid biofilm responses to rehydration during flow intermittency might be essential in sustaining biofilm functional diversity in intermittent streams.  相似文献   

3.
1. Manipulative experiments were carried out in four Hong Kong streams (two shaded, two unshaded) to investigate the impact of grazing by an algivorous fish, Pseudogastromyzon myersi, on benthic algal biomass and assemblage composition. Experiments were conducted and repeated during both the dry and wet seasons to determine whether spate‐induced disturbance modified any grazing effect. Treatments comprised fish exclusion and inclusion via closed and open cages, with a no‐cage treatment used as a control for the cage effect. Treatments were maintained for 4 weeks in each experimental run. 2. Grazing by P. myersi reduced benthic algal biomass and the organic matter content of periphyton in open cages and the no‐cage treatment relative to closed cages. The similarity between open‐cage and no‐cage treatments was evidence that the overall difference among treatments was caused by limiting fish access to closed cages and not merely an artifact of caging. Grazing effects were broadly similar in all streams, but there was a significant statistical interaction between treatments and seasons. 3. Analysis of dry‐season data matched the overall trend in inter‐treatment differences, confirming the effects of grazing by P. myersi on algal biomass and periphyton organic matter. Significant differences in algal assemblage composition between closed‐cage and no‐cage treatments during the dry season reflected reductions in the abundance of erect, stalked diatoms (Gomphonema) and filamentous cyanobacteria (Homeothrix). Removal of these vulnerable overstorey algae by P. myersi resulted in greater abundance of understorey diatoms (Achnanthes and Cocconeis) in the no‐cage treatment in all streams during the dry season. The composition of algal assemblages in open cages was intermediate between the other two treatments. 4. Although fish densities were greater in all streams during the wet season, spate‐induced disturbance obscured grazing effects and there were no significant differences among treatments attributable to fish grazing. Seasonal variation in impacts of P. myersi grazing provides support for the harsh‐benign hypothesis, and confirms that biotic factors are less important controls of stream algal biomass and assemblage structure during periods (i.e. the wet season in Hong Kong) when abiotic disturbances are frequent or intense.  相似文献   

4.
  1. In some regions, climate change is increasing the variability of rainfall and the frequency of extreme events such as drought. Consequently, non-flow periods have grown in length and frequency, both in temporary and in formerly permanent streams. Water abstraction for human use may further prolong these dry periods.
  2. We analysed the resistance and resilience of biofilms from permanent and temporary streams to non-flow conditions. This was achieved by exposing cobbles (collected from permanent and temporary streams) with intact biofilm to 31 days of non-flow, followed by 20 days of stream flow in artificial stream channels. Biofilm resistance and resilience were assessed at a structural (algal biomass, pigment composition, and algae and cyanobacteria composition) and functional level (photosynthetic efficiency and community metabolism).
  3. Algal taxa in biofilms from permanent and temporary streams differed throughout the experiment. Biofilms from permanent streams were less resistant to non-flow than those from temporary streams at structural level. Permanent stream biofilms also presented lower resilience at a structural level, but responded similarly to temporary stream biofilms at a functional level.
  4. Our investigation shows how the non-flow period disturbed permanent stream biofilms, and suggests that temporary stream biofilms will have greater adaptive capacity as hydroperiod becomes shorter due to climate change.
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5.
6.
The objective of this study was to measure the photo‐acclimation response of stream algae inhabiting thin biofilms across a range of light treatments comparing both growth rates and net primary productivity (NPP). Algae were grown on clay tiles incubated in artificial stream channels where light levels were manipulated by layering a neutral‐density shade cloth over each channel. We measured NPP and algal growth rates in the early stages of community development and compared assemblages that were either acclimated or unacclimated to a given light treatment. Algal growth rates did acclimate to light treatment, with saturation occurring at light levels that were substantially lower for the acclimated communities. Growth efficiency calculated from algal biovolume increased by a factor of 3. However, algal NPP showed a weaker photo‐acclimation response, with only a 30% increase in photosynthetic efficiency. Our results indicate that diatom‐dominated periphyton in thin biofilms are probably not light limited in many shaded streams with respect to growth rates, but are light limited with respect to NPP.  相似文献   

7.
1. Nutrient diffusing substrata were used to determine the effect of added inorganic nitrogen (N) and phosphorus (P) on the development of epilithic and epixylic biofilms in 10 North American streams. Four treatments of diffusing substrata were used: Control (agar only), N addition (0.5 m NaNO3), P addition (0.5 m KH2PO4), and N + P combined (0.5 m NaNO3 + 0.5 m KH2PO4). Agar surfaces were covered with glass fibre filters (for epilithon) or discs of untreated white oak wood veneer (for epixylon). 2. We found that if algae showed significant response to nutrient addition, N limitation (either N alone or N with P) was the most frequent response both on GF/F filters and on wood. Despite the low dissolved nutrient concentrations in our study streams, more than a third of the streams did not show any response to N or P addition. In fact, P was never the sole limiting nutrient for algal biofilms in this study. 3. Nutrient addition influenced algal colonisation of inorganic versus organic substrata in different ways. The presence of other biofilm constituents (e.g. fungi or bacteria) may influence whether algal biomass on wood increased in response to nutrient addition. Algae on organic and inorganic substrata responded similarly to nutrient addition in only one stream. 4. Fungal biomass on wood was nutrient limited in six of 10 study streams. N limitation of fungal biomass (with or without secondary P limitation) was most frequent, but P limitation did occur in two streams. 5. Our results show that biomass responses to nutrient addition by the heterotrophic and autotrophic components of the epixylic biofilm were different, though both experienced the same stream nutrient conditions. For algae and fungi growing on wood, limiting nutrients were rarely similar. Only three of nine streams showed the same biomass response to nutrient addition, including two that showed no significant change in biomass despite added nutrients.  相似文献   

8.
Abstract Relationships between biofilm structural components (algal and bacterial biomass) and the activities of some extracellular enzymes that contribute to the ability to degrade organic matter) were explored for six Atlantic and three Mediterranean streams and rivers. The biofilms in these fluvial systems accounted for a wide range of bacterial and algal biomass and colonized the most common benthic habitats. Ratio of bacteria/algae biomass was lower in Atlantic than in Mediterranean streams, but enzymatic activities (β-glucosidase, β-xylosidase, phosphatase) were in general greater in the Mediterranean stream biofilms. Climatic characteristics (especially temperature) may explain the differences in enzymatic activities between biofilms of similar structure but different flow regime. The ratio β-xylosidase: β-glucosidase was similar (around 0.5) for all streams and substrata considered, showing that there is a general higher utilization of cellobiosic than xylobiosic molecules in fluvial systems. In general, highly heterotrophic biofilms showed lower extracellular enzymatic activities than more autotrophic biofilms. Maximum enzymatic activity is achieved when the algal biomass is two- to threefold higher than the bacterial biomass. The relevance of algal biomass on the heterotrophic ability of biofilms may be related to the physical proximity between the two, but also to the high proportion of polymeric carbohydrates included in algal exudates and lysis products, whose use is enzyme-mediated. Received: January 2000; Accepted: April 2000; Online Publication: 18 July 2000  相似文献   

9.
Food webs in the rhithral zone rely mainly on allochthonous carbon from the riparian vegetation. However, autochthonous carbon might be more important in open canopy streams. In streams, most of the microbial activity occurs in biofilms, associated with the streambed. We followed the autochthonous carbon transfer toward bacteria and grazing protozoa within a stream biofilm food web. Biofilms that developed in a second-order stream (Thuringia, Germany) were incubated in flow channels under climate-controlled conditions. Six-week-old biofilms received either 13C- or 12C-labeled CO?, and uptake into phospholipid fatty acids was followed. The dissolved inorganic carbon of the flow channel water became immediately labeled. In biofilms grown under 8-h light/16-h dark conditions, more than 50% of the labeled carbon was incorporated in biofilm algae, mainly filamentous cyanobacteria, pennate diatoms, and nonfilamentous green algae. A mean of 29% of the labeled carbon reached protozoan grazer. The testate amoeba Pseudodifflugia horrida was highly abundant in biofilms and seemed to be the most important grazer on biofilm bacteria and algae. Hence, stream biofilms dominated by cyanobacteria and algae seem to play an important role in the uptake of CO? and transfer of autochthonous carbon through the microbial food web.  相似文献   

10.
Cascading effects of predators can affect ecosystem properties by changing plant biomass, distribution and assemblage composition. Using data from field surveys and whole‐stream experiments we tested the hypothesis that predatory trout change assemblage composition of benthic algae in high‐elevation streams mediated by grazer behavior. Field surveys revealed that the taxonomic composition of algal assemblages differed significantly between streams that contained trout and those that were fishless; but comparisons of palatable versus unpalatable algal taxa between fish and fishless streams were equivocal because of high natural variability. Therefore, we tested for a behavioral (non‐consumptive) trophic cascade experimentally by adding brook trout chemical cues to six naturally fishless streams for 25 days and compared responses of grazers and algae to six reference streams without fish cues added. Algal response variables included rates of change in the abundance of three physiognomic categories, from most palatable (attached erect and prostrate diatoms) to least palatable (non‐diatoms), as determined from food selectivity analyses of the most common grazers (mayflies and caddisflies). Fish cues did not affect the mean densities or changes in densities of total grazers or any individual grazer species. However, in streams where fish cues were added, rates of accrual of attached erect diatoms, which was the preferred algal type for the grazer most vulnerable to trout predation (Baetis), were higher and their densities increased significantly faster with increasing densities of this grazer species than in reference streams. Results of his experiment support the hypothesis that predator induced suppression of grazer foraging behavior, rather than cascading effects of top predators on grazer density, may contribute to variation in the composition of algal assemblages among streams by allowing proliferation of most palatable algal species.  相似文献   

11.
Tropical stream food webs are thought to be based primarily on terrestrial resources (leaf litter) in small forested headwater streams and algal resources in larger, wider streams. In tropical island streams, the dominant consumers are often omnivorous freshwater shrimps that consume algae, leaf litter, insects, and other shrimps. We used stable isotope analysis to examine (1) the relative importance of terrestrial and algal‐based food resources to shrimps and other consumers and determine (2) if the relative importance of these food resources changed along the stream continuum. We examined δ15N and δ13C signatures of leaves, algae, macrophytes, biofilm, insects, snails, fishes, and shrimps at three sites (300, 90, and 10 m elev.) along the Río Espíritu Santo, which drains the Caribbean National Forest, Puerto Rico. Isotope signatures of basal resources were distinct at all sites. Results of two‐source δ13C mixing models suggest that shrimps relied more on algal‐based carbon resources than terrestrially derived resources at all three sites along the continuum. This study supports other recent findings in tropical streams, demonstrating that algal‐based resources are very important to stream consumers, even in small forested headwater streams. This study also demonstrates the importance of doing assimilation‐based analysis (i.e., stable isotope or trophic basis of production) when studying food webs.  相似文献   

12.
1. Mediterranean climate regions are characterised by long summer droughts that usually involve flow intermittency in low‐ to mid‐order streams. Flow intermittency implies flow cessation, drying and subsequent rewetting of the streambed, and affects both autotrophic and heterotrophic processes. The balance between these processes, as well as the balance in the use of carbon (C), nitrogen (N) and phosphorus (P) may change because of the ongoing increase in stream flow intermittency caused by global change in many regions. It is therefore crucial to understand better the consequences of this phenomenon. 2. Our two initial hypotheses were (i) that flow intermittency would impact more on autotrophic than on heterotrophic processes in stream biofilms owing to the higher water dependence of autotrophs, as well as differences in the water storage capacity of the stream biofilm compartments where autotrophic and heterotrophic processes mainly occur (surface cobbles versus hyporheic sediments) and (ii) that the C‐N‐P use by biofilms would change during the dry period (terrestrial phase) owing to the extreme water stress conditions. These hypotheses were tested by analysing the functional response of the main stream biofilms (epilithic, epipsammic and hyporheic) during flow cessation, desiccation and rewetting in a Mediterranean forested stream. The autotrophic response was characterised through changes in the photon yield, whereas the heterotrophic response was characterised by changes in the extracellular enzyme activities. 3. Streambed desiccation had clear effects on the functioning of stream biofilms. Autotrophic biomass decreased by 80% with streambed desiccation, but recovered rapidly after flow resumption. Heterotrophs were more resistant to water stress, especially in the epipsammic and hyporheic biofilms where bacterial cell density decreased only by 20%. 4. Extracellular enzyme activities remained relatively high, and the balance in the C‐N‐P use by biofilms changed during the dry period. The C and P breakdown capacities were maintained during dry conditions, especially in the epipsammic and hyporheic biofilms, but the degradation of N compounds sharply decreased. Elemental molar ratios (C:N and C:P) of the different biofilms also changed with streambed desiccation. C:P ratios increased from 80 to 300, while the C:N ratios increased from 10 to 16. 5. Given the contrasting responses of autotrophic and heterotrophic processes in the different biofilms, our results suggest that the current increase in flow intermittency extent is likely to increase the relative importance of heterotrophic processes in stream ecosystems, as well as the relative contribution of the hyporheic biofilm to C‐N‐P use. Our results further suggest that the longer streams remain dry, the more the biofilm stoichiometry will change.  相似文献   

13.
14.
Trophic cascades are extensively documented in nature, but they are also known to vary widely in strength and frequency across ecosystems. Therefore, much effort has gone into understanding which ecological factors generate variation in cascade strength. To identify which factors covary with the strength of cascades in streams, we performed a concurrent experiment across 17 streams throughout the Sierra Nevada Mountains. We eliminated top consumers from experimental substrates using electrical exclusions and compared the strength of indirect effects of consumers on the biomass of primary producers relative to control patches. In each stream we 1) classified the dominant invertebrate herbivores according to life‐history traits that influence their susceptibility to predators, 2) determined the abundance and diversity of algae and herbivores, and 3) measured production‐to‐biomass ratios (P:B) of the stream biofilm. This allowed us to assess three common predictions about factors thought to influence the strength of trophic cascades: cascade strength 1) is weaker in systems dominated by herbivores with greater ability to evade or defend against predators, 2) is stronger in systems characterized by low species diversity, and 3) increases with increasing producer P:B. When averaged across all streams, the indirect effect of predators increased the biomass of periphyton by a mean 60%. However, impacts of predators on algae varied widely, ranging from effects that exacerbated algal loss to herbivores, to strong cascades that increased algal biomass by 4.35 times. Cascade strength was not related to herbivore traits or species diversity, but decreased significantly with increasing algal diversity and biofilm P:B in a stream. Partial regression analyses suggested that the relationship between cascade strength and algal diversity was spurious, and that the only significant covariate after statistically controlling for cross‐correlations was algal P:B. Our study contributes to the ongoing debate about why trophic cascade strength varies in nature and is useful because it eliminates factors that have no potential to explain variation in cascades within these stream ecosystems.  相似文献   

15.
To investigate the effects of nitrate enrichment, phosphate enrichment, and light availability on benthic algae, nutrient-diffusing clay flowerpots were colonized with algae at two sites in a Hawaiian stream during spring and autumn 2002 using a randomized factorial design. The algal assemblage that developed under the experimental conditions was investigated by determining biomass (ash-free dry mass and chlorophyll a concentrations) and composition of the diatom assemblage. In situ pulse amplitude-modulated fluorometry was also used to model photosynthetic rate of the algal assemblage. Algal biomass and maximum photosynthetic rate were significantly higher at the unshaded site than at the shaded site. These parameters were higher at the unshaded site with either nitrate, or to a lesser degree, nitrate plus phosphate enrichment. Analysis of similarity of diatom assemblages showed significant differences between shaded and unshaded sites, as well as between spring and autumn experiments, but not between nutrient treatments. However, several individual species of diatoms responded significantly to nitrate enrichment. These results demonstrate that light availability (shaded vs. unshaded) is the primary limiting factor to algal growth in this stream, with nitrogen as a secondary limiting factor.  相似文献   

16.
Stream bacteria may be influenced by the composition and availability of dissolved organic matter (DOM) and inorganic nutrients, but knowledge about how individual phylogenetic groups in biofilm are affected is still limited. In this study, the influence of DOM and inorganic nutrients on stream biofilm bacteria was examined. Biofilms were developed on artificial substrates (unglazed ceramic tiles) for 21 days in a northeastern Ohio (USA) stream for five consecutive seasons. Then, the developed biofilm assemblages were exposed, in the laboratory, to DOM (glucose, leaf leachate, and algal exudates) and inorganic nutrients (nitrate, phosphate, and nitrate and phosphate in combination) amendments for 6 days. Bacterial numbers in the biofilms were generally higher in response to the DOM treatments than to the inorganic nutrient treatments. There were also apparent seasonal variations in the response patterns of the individual bacterial taxa to the nutrient treatments; an indication that limiting resources to bacteria in stream biofilms may change over time. Overall, in contrast to the other treatments, bacterial abundance was generally highest in response to the low-molecular-weight DOM (i.e., glucose) treatment. These results further suggest that there are interactions among the different bacterial groups in biofilms that are impacted by the associated nutrient dynamics among seasons in stream ecosystems.  相似文献   

17.
1. Temporary streams comprise a large proportion of the total length of most stream networks, and the great majority of arid‐land stream networks, so it is important to understand their contribution to biotic diversity at both local and landscape scales. 2. In late winter 2010, we sampled invertebrate assemblages in 12 reaches of a large arid‐land stream network (including perennial and intermittent headwaters, intermittent middle reaches and perennial rivers) in south‐east Arizona, U.S.A. Intermittent reaches had then been flowing for c. 60 days, following a dry period of more than 450 days. We sampled a subset of the perennial study reaches three more times between 2009 and 2011. Since intermittent reaches were dry during these additional sampling periods, we used assemblage data from two other intermittent streams in the study network (sampled in 2004–05 and 2010) to explore interannual variability in intermittent stream assemblage composition. 3. Invertebrate richness was lowest in intermittent reaches, despite their often being connected to species‐rich perennial reaches. The assemblages of these intermittent reaches were not simply a subset of the species in perennial streams, but rather were dominated by a suite of stoneflies, blackflies and midges with adaptations to intermittency (e.g. egg and/or larval diapause). On average, 86% of individuals in these samples were specialists or exclusive to intermittent streams. Predators were 7–14 times more abundant in perennial than in intermittent reaches. 4. Despite being separated by long distances (12–25 km) and having very different physical characteristics, the assemblages of perennial headwaters and rivers were more similar to one another than to intervening intermittent reaches, emphasising the prime importance of local hydrology in this system. 5. The duration and recurrence intervals of dry periods, and the relative importance of dispersal from perennial refuges, probably influence the magnitude of biological differences between neighbouring perennial and temporary streams. Although perennial headwaters supported the highest diversity of invertebrates, intermittent reaches supported a number of unique or locally rare species and as such contribute to regional species diversity and should be included in conservation planning.  相似文献   

18.
1. The apparent absence of a specialist herbivorous grazer guild from many acid streams suggests that algae-grazer linkages in acid-stream food webs are weak or absent. It has been hypothesized that the absence of herbivores is a consequence of the low quality and/or quantity of biofilms in acid streams.
2. We compared the taxonomic composition, biomass and potential nutritional quality of epilithic biofilms from four acid and four circumneutral streams, and examined whether nymphs of a herbivore–detritivore, the stonefly Nemurella pictetii (Plecoptera: Nemouridae), could grow equally well when fed on eight biofilms from four acid and four circumneutral streams.
3. Biofilms from acid and circumneutral streams differed strongly in algal composition, the former having relatively more coccoid green algae but fewer diatoms and filamentous green algae. Diatom floras differed with stream water pH.
4. The quantity (i.e. area-specific chlorophyll content, algal numbers and AFDM) and quality (biomass-specific protein and soluble carbohydrate content) of biofilms differed significantly, both among sites of similar pH, and overall between the groups of acid and circumneutral streams.
5. Nymphs of N. pictetii grew successfully on biofilms for 8–10 weeks up to emergence. However, no systematic differences in growth rate were found between the two groups of acid and circumneutral streams. Differences in the digestibility of benthic algae from different sources, and the adjustment of nymphal feeding rates, are discussed in the light of a lack of a clear relationship between growth and food quality.  相似文献   

19.
20.
1. Of the relatively few studies that have examined consequences of amphibian declines on stream ecosystems, virtually all have focused on changes in algae (or algal‐based food webs) and little is known about the potential effects of tadpoles on leaf decomposition. We compared leaf litter decomposition dynamics in two neotropical streams: one with an intact community of tadpoles (with frogs) and one where tadpoles were absent (frogless) as a result of a fungal pathogen that had driven amphibians locally extinct. The stream with tadpoles contained a diverse assemblage (23 species) of larval anurans, and we identified five species of glass frog (Centrolenidae) tadpoles that were patchily distributed but commonly associated with leaf detritus and organic sediments in pools. The latter reached total densities of 0–318 tadpoles m?2. 2. We experimentally excluded tadpoles from single‐species leaf packs incubated over a 40‐day period in streams with and without frogs. We predicted that decomposition rates would be higher in control (allowing access of tadpoles) treatments in the study stream with frogs than in the frogless stream and, in the stream with frogs, in the control than in the tadpole exclusion treatment. 3. In the stream with frogs, Centrolene prosoblepon and Cochranella albomaculata tadpoles were patchily distributed in leaf packs (0.0–33.3 m?2). In contrast to our predictions, leaf mass loss and temperature‐corrected leaf decomposition rates in control treatments were almost identical in our stream with frogs (41.01% AFDM lost, kdegree day = ?0.028 day?1) and in the frogless stream (41.81% AFDM lost, kdegree day = ?0.027 day?1) and between control and tadpole exclusion treatments within each stream. Similarly, there were no significant differences in leaf pack bacterial biomass, microbial respiration rates or macroinvertebrate abundance between treatments or streams. Invertebrate assemblages on leaf packs were similar between treatments (SIMI = 0.97) and streams (SIMI = 0.95) and were dominated by larval Chironomidae, Simuliidae (Diptera) and larval Anchytarsus spp. (Coleoptera). 4. In contrast to dramatic effects of grazing tadpoles on algal communities observed previously, tadpoles had no major effects on decomposition. While centrolenid tadpoles were common in the stream with frogs, their patchy distribution in both experimental and natural leaf packs suggests that their effects on detrital dynamics and microbes are probably more localised than those of grazing tadpoles on algae.  相似文献   

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