A model for resolving the plankton paradox: coexistence in open flows

1. Recent developments in the field of chaotic advection in hydrodynamical/environmental flows encourage us to revisit the population dynamics of competing species in open aquatic systems.
2. We assume that these species are in competition for a common limiting resource in open flows with chaotic advection dynamics. As an illustrative example, we consider a time periodic two-dimensional flow of viscous fluid (water) around a cylindrical obstacle.
3. Individuals accumulate along a fractal set in the wake of the cylinder, which acts as a catalyst for the biological reproduction process. While in homogeneous, well mixed environments only one species could survive this competition, coexistence of competitors is typical in our hydrodynamical system.
4. It is shown that a steady state sets in after sufficiently long times. In this state, the relative density of competitors is determined rather by the fractal nature of the spatial distribution of the advected species, and by their initial conditions, than by their competitive abilities. We argue that two factors, the strong chaotic mixing along a fractal set and the boundary layer around the obstacle, are responsible for the coexistence.     

Large-scale synchrony and inter-annual variability in roach recruitment in the Rhône River: the relative role of climatic factors and density-dependent processes

1. We report patterns of temporal variation in the recruitment of roach ( Rutilus rutilus ). The data consist of the annual abundance of the first 2 year-classes, 0+ and 1+ fish, at four sites in the Rhône River (France) between 1987 and 1997. Over this 11-year period both 0+ and 1+ fish fluctuated strongly.
2. Cross-correlation indicated high spatial synchrony in 0+ dynamics, although correlations among sites in 1+ dynamics were weaker. No clear pattern was apparent in the relationship between the level of synchrony and distance between pairs of sites.
3. The spatial synchrony in 0+ fish could be attributable to large-scale variations in weather, influencing water temperature. Total body length of 0+ roach was correlated with water temperature (expressed in degree-days over 12 °C), and water temperature was the main factor explaining inter-annual variation in 0+ cohort size. Monthly variation in abiotic factors (measured by standard deviation in water temperature and discharge) did not influence 0+ fluctuations. Correlations with June water temperature suggest that year-class strength was mainly determined by abiotic factors during the first few months of life.
4. The absence of spatial synchrony in 1+ fluctuations suggests little correlation between survival and abiotic conditions during the first year of life, other factors influencing survival.
5. Survival in the first year was density-dependent. Intraspecific competition within the 0+ cohort could thus influence the fluctuations in recruitment to older age-classes.
6. The implications of age- or stage-dependent synchrony in temporal variation for species with complex life histories are discussed. Studying spatial synchrony for the different life history stages could enhance our understanding of the population dynamics of spatially structured species.     

Substratum patch selection in the lacustrine mussels Elliptio complanata and Pyganodon grandis grandis

1. Sediment selection was investigated under controlled conditions in two common lake-dwelling species of freshwater mussels (Bivalvia: Unionidae), Elliptio complanata and Pyganodon grandis grandis .
2. Sediment choice was determined in six independent experiments under controlled conditions by distributing mussels randomly or evenly in tanks containing patches of sand and mud, and following their movement among sediment patches in experiments lasting between 30 and 45 days.
3. In all experiments, both species were found most frequently in muddy sediment patches. Movement toward muddy patches occurred rapidly: an average of nearly 80% of Pyganodon grandis grandis were found in mud after 30 days. Elliptio complanata moved rapidly to patches of mud at the start of experiments, but occupation of muddy sediments appeared to decrease after about 30 days.
4. Our results contrast with many field studies that suggest populations of lake-dwelling freshwater mussels infrequently inhabit mud and silt. We therefore postulate that large-scale mussel distribution in lakes is influenced most strongly by factors other than sediment composition.     

Costs, benefits and characteristics of mixotrophy in marine oligotrichs

1. Oligotrich ciliates are an important part of most marine plankton communities. Mixotrophic (chloroplast-sequestering) oligotrichs, a common component of marine oligotrich communities, obtain fixed carbon from both photosynthesis as well as the ingestion of particulate food. Mixotrophy, in general, is often considered an adaptation permitting exploitation of food-poor environments. We examined the hypothesis that, among oligotrichs, mixotrophs may be at a disadvantage relative to heterotrophs in food-rich conditions in a nutrient-enrichment experiment. We compared growth responses of mixotrophic and heterotrophic oligotrichs in natural communities from the N.W. Mediterranean Sea in microcosms with daily nutrient additions resulting in increases in nanoflagellates and Synechococcus populations. The results indicated that both mixotrophic and heterotrophic oligotrichs respond to prey increases with rapid growth (μ=1.2 d−1).
2. To examine the hypothesis that the proportion of mixotrophic to heterotrophic oligotrichs changes with the trophic status of a system, increasing with oligotrophy, we examined data from a variety of marine systems. Across systems ranging in chlorophyll concentration from about 0.1 to 40 μg L−1, oligotrich cell concentrations are correlated with chlorophyll concentrations, and mixotrophs are a consistent component of oligotrich communities, averaging about 30% of oligotrich cell numbers.
3. We discuss the costs, benefits and possible uses of mixotrophy in marine oligotrichs and suggest that mixotrophy in marine oligotrichs is not closely linked to the exploitation of food-poor environments, but probably serves a variety of purposes.     

Silica and diatom growth in Lough Neagh: the importance of internal recycling

1. Lough Neagh is a large eutrophic lake with a high dissolved silica (SiO2) concentration arising from the basaltic rock in parts of the catchment and the normal winter maximum concentration is over 8 mg L−1. Based on frequent observations between 1974 and 1997, the annual cycles of SiO2 input, uptake and release are explored.
2. Large spring blooms of the planktonic diatoms Aulacoseira subarctica Haworth and Stephanodiscus astraea occur every year and are terminated by SiO2 depletion. Although there are periods when one diatom species has dominated the other, over the period as a whole, the two diatoms appear to be in a stable coexistence.
3. A model of the effect of river inputs on lake concentration shows that without SiO2 release from the sediment, the maximum spring dissolved SiO2 concentration (and hence diatom crop) would be only about one-third of that actually observed.
4. It is concluded that within-lake processes play a large role and are potentially more variable than catchment processes in determining the available SiO2 in Lough Neagh.
5. The role of benthic animals, especially Chironomus anthracinus, in the sediment SiO2 release process is discussed. Field data suggest the SiO2 release rate is highly sensitive to temperature, but this could be partly caused by an interaction between temperature and animal activity.     

Mixotrophy in planktonic protists: an overview

1. An overview is provided of the role of mixotrophic protists in plankton communities. Consideration of the importance of phagotrophy in the evolution of photosynthetic eucaryotes suggests that mixotrophy as a nutritional strategy can arise rather readily.
2. Mixotrophic protists actually present a spectrum of nutritional strategies. However, recognition of distinct groups of mixotrophs based on nutritional behaviour facilitates consideration of their functional role and of competitive interactions with other types of planktonic protists.
3. Consideration of the costs and benefits of mixotrophy as a nutritional strategy allows the development of several empirical predictions about the probable outcome of resource competition between mixotrophs and obligate phototrophs or phagotrophs. Existing results from laboratory and field experiments allow some of these predictions to be evaluated.
4. These results indicate that, under specified conditions, mixotrophs should represent an important link in the flux of materials through planktonic food webs. However, quantifying these fluxes remains a challenge for the future.     

Assessing the performance of volunteers in monitoring streams

1. Citizens are concerned about the quality of water resources and many participate in monitoring activities, though doubts remain about the quality of the data volunteers collect. We trained volunteers to collect benthic macroinvertebrates using professional protocols. Of the seven stream sites sampled by volunteer crews, six sites were also sampled by professional crews.
2. In the laboratory, volunteers used morphological features to identify as many different taxa as possible within the major insect orders; their identification was approximately to family. Volunteers calculated five metrics: total taxon richness, richness of three key groups (Ephemeroptera, Plecoptera and Trichoptera), and percentage dominance of the three most abundant taxa. All metrics were strongly correlated with (a) the percentage of urbanized area in the catchment and (b) the metrics derived from a more complete taxonomic identification by a professional scientist. Taxon richness metrics declined with urban development, while percent dominance increased.
3. An overall summary multimetric index was used to compare the field and laboratory procedures of volunteers and professionals. Using an ANOVA model, we detected no significant difference between field samples collected by volunteers and professionals. The variance of index values associated with differences between crews was zero. The ability of the index to detect significant differences among sites (statistical power) improved by only 13% for assessments based on professional laboratory identification instead of volunteer laboratory identification.
4. Citizen volunteers, when properly trained, can collect reliable data and make stream assessments that are comparable to those made by professionals. Data collected by volunteers can supplement information used by government agencies to manage and protect rivers and streams.     

Variation in growth rates of the zebra mussel, Dreissena polymorpha, within Lake Wawasee

1. Field experiments conducted in Lake Wawasee in 1995 and 1996 measured the response of shell growth of Dreissena polymorpha to environmental gradients.
2. Shell growth decreased with initial shell length in four mussel size classes ranging between 8 and 22 mm, and decreased with depth, with mussels in shallow water (<4 m) having growth rates nearly twice those of mussels in deeper water (4–7 m).
3. Growth occurred early in the spring–summer period (May–June) with relatively little shell added later in the summer (July–September), and varied significantly among sites within Lake Wawasee, but not between the 2 years of this study.
4. Rank order of sites was consistent for both years implying that environmental conditions responsible for variation in shell growth were stable within Lake Wawasee.
5. Cage design did not have a significant effect on mussel shell growth nor did the distance of growth cages above the bottom (0.5–0.75 m above the bottom versus directly on the bottom).
6. This study demonstrates the sensitivity of adult mussel growth to subtle variation in environmental conditions occurring within and among lakes, with potential consequences for mussel population dynamics and community structure and function.     

The costs of moulting in Daphnia; mineral regulation of carbon budgets

1. Zooplankton differ in their elemental contents. For the essential elements calcium (Ca) and phosphorus (P), Daphnia spp. have particularly high contents, and thus high demands, compared with other cladocera and copepods.
2. There are recent indications that the growth of zooplankton, notably Daphnia , may be directly constrained by the availability of P and Ca.
3. Major pools of P and Ca are associated with the carapace, and the moulting process represents a constant drain of these elements, as well as carbon (C), from the animals.
4. The demands for P and Ca for moulting may pose constraints on the carbon accumulation in the animals, and thus the extensive needs for these elements may translate into a total loss of C transported in the pelagic food web.
5. While the costs of moulting are substantial for the zooplankton, the export of minerals bound in the moult will play only a minor role for the epilimnetic element budgets in most lakes.
     

Dynamics of the exotic Daphnia lumholtzii and native macro-zooplankton in a subtropical chain-of-lakes in Florida, U.S.A.

1. Seasonal dynamics of the exotic Daphnia lumholtzii and native macro-zooplankton species were studied for 2 years in six inter-connected lakes in Florida, U.S.A. The lakes ranged widely in pH, colour and trophic status, and were dominated by copepods.
2. All six lakes contained both D. lumholtzii and the native D. ambigua , but the two species did not overlap in time. D. ambigua was dominant in autumn–spring, coinciding with lower water temperature, higher transparency and lower nutrient and chlorophyll a (Chl a ) concentrations than in summer, when D. lumholtzii was dominant.
3. Based on the field observations, temperature optima were 24 °C for D. ambigua and 29 °C for D. lumholtzii , suggesting that temperature plays a role in determining dominance among the daphnids of subtropical Florida lakes.
4.  D. lumholtzii has not displaced native cladocerans but occupies a 'vacant' seasonal niche, unexploited due to the inability of native taxa to tolerate high temperature. Furthermore, D. lumholtzii did not significantly alter algal–zooplankton interactions. There was evidence of top–down control by grazing, but it was primarily attributable to the native taxa.     

Dwelling at the oxycline: does increased stratification provide a predation refugium for the Lake Victoria sardine Rastrineobola argentea?

1. We tested the hypothesis that increased hypoxia in Lake Victoria provides a refugium for the cyprinid dagaa ( Rastrineobola argentea ) against hypoxia-sensitive Nile perch ( Lates niloticus ).
2. Hypoxia in the main habitat of dagaa was rare during 1979–80, but lasted 3–5 months in 1987 and 1988, during which time adult dagaa spent the day just above the oxycline instead of near the bottom.
3. Dissolved oxygen concentrations in the layer just above the oxycline were not critical for Nile perch, but a mass kill of this species following a sudden upwelling of hypoxic water suggested that oxycline-dwelling is risky.
4. Our data suggest no difference between dagaa and Nile perch in the level at which dissolved oxygen starts to limit their vertical distribution in the water column, but dagaa seems to be even more sensitive to extreme hypoxia (<1–2 mg O2 L−1) than Nile perch.
5. We argue that oxycline-dwelling dagaa are not seeking a predation refugium but that they are limited by low oxygen levels in reaching their feeding areas near the bottom.     

Factors influencing macroinvertebrate colonization of seasonal wetlands: responses to emergent plant cover

1. We conducted field experiments to examine factors influencing macroinvertebrate colonization of seasonally flooded marshes. Few macroinvertebrate species were found aestivating in soils within non-flooded wetlands indicating that most taxa colonize these marshes from other flooded habitats.
2. We manipulated amounts of salt grass ( Distichlis spicata ) to examine how emergent plant cover affects aerial colonization by macroinvertebrates. Areas mowed 3 weeks before flooding had low plant cover, areas mowed 5 and 9 weeks before flooding had medium and high plant cover, respectively, and non-mowed control areas had the most plant cover. Macroinvertebrate numbers and biomass were generally higher in mowed treatment areas than in control areas, but overall diversity was generally higher in high plant cover and control areas than in low plant cover areas.
3. Mosquitoes (Culicidae), brine flies (Ephydridae) and hover flies (Syrphidae) were positively correlated with amount of plant cover, and waterboatmen (Corixidae), midges (Chironomidae) and water scavenger beetles (Hydrophilidae) were negatively correlated with plant cover. Species assemblages changed seasonally among treatment areas because these taxa colonize wetlands at different times in the year.
4. These results demonstrate that invertebrate communities may be different within plant stands with heterogeneous amounts of emergent cover, and management practices that alter the structure of wetland vegetation can influence macroinvertebrate communities colonizing seasonal marshes.     

Epilithic algal communities and their relationship to environmental variables in lakes of the English Lake District

1. The relationship between epilithic algal communities and 17 environmental variables from 17 oligo- to eutrophic lakes in the English Lake District was explored using canonical correspondence analysis (CCA). Total phosphorus (TP) and calcium (Ca) concentration were the most important variables accounting for species distribution.
2. Weighted-averaging regression and calibration models with tolerance downweighting and classical deshrinking were developed to infer TP, Ca, dissolved inorganic carbon (DIC) and conductivity from the relative abundance of 138 epilithic algal taxa.
3. The ranges of the environmental variables covered by the models are 0.8-49.2 μg L−1 for TP, 2.2–13.0 mg L−1 for Ca, 0.5–8.6 mg L−1 for DIC and 38–124 μS cm−1 for conductivity. Within these limits the models can be used to infer chemical properties of lakes from epilithic communities in the English Lake District.
4. The major advantages of using transfer functions based on epilithic communities are the low logistic requirements and the integrative character of algal samples compared to direct point measurements of chemical parameters of the lakes.     

Effects of aquatic plant management on stream metabolism and oxygen balance in streams

1. In unshaded, nutrient-rich streams, prolific growth of stream macrophytes often results in flows that over-top the banks and in high primary production and respiration that may result in extreme diel variations in dissolved oxygen. Consequently, water protection authorities commonly remove macrophytes periodically.
2. We investigated the effect of plant removal on stream metabolism and oxygen balance in two Swiss streams with a high macrophyte biomass. We monitored the concentration of dissolved oxygen before and after macrophytes were removed by cutting and dredging, and calculated rates of gross primary production and ecosystem respiration by means of diel oxygen curves.
3. The removal of plants, which had reached a dry biomass of 320–420 g m−2 immediately before plant removal, had a different impact on stream metabolism in the two streams. In the first (plants removed in May), neither primary production nor ecosystem respiration were significantly affected. In the second (plants removed in late July), gross primary production and ecosystem respiration were reduced by about 70%. In this latter stream gross primary production increased in the first 2 weeks after plant removal but never recovered to pre-disturbance levels.
4. The removal of plants coincided with only a moderate increase in nocturnal oxygen concentration (+1 mg L−1). This, and the rapid partial recovery of stream metabolism in the second stream, suggests that an increase in the oxygen concentration after plant cutting is transient in unshaded, nutrient-rich streams.     

Tests of autotrophic picoplankton as early indicators of nutrient enrichment in an ultra-oligotrophic lake

1. The potential of picocyanobacteria as early indicators of changes in nutrient loading and trophic status was examined in an ultra-oligotrophic lake.
2. The study consisted of in situ, short-term, nutrient-addition bioassays and surveys of picocyanobacterial variables and other indices of trophic state at six sites selected to include a range of localized nutrient loadings to the lake from its catchment. The bioassays and surveys were conducted six times throughout 1 year.
3. Experimental additions of small amounts of ammonium-N and phosphate-P did not stimulate picocyanobacterial growth, and phosphate additions often reduced picocyanobacterial growth rates.
4. A correlation analysis using pooled data from all sites and sampling dates showed that the abundance of aggregated picocyanobacteria was strongly correlated with nutrient concentration and ratios. Other variables, including the concentration of single-cell picocyanobacteria, chlorophyll a and primary production, were poorly correlated with nutrients.
5. The results show that picocyanobacteria in oligotrophic lakes are sensitive to extremely small changes in nutrient availability and that they can respond in complex ways. The mechanisms by which they respond to such changes require further study before they can be used as bio-indicators of nutrient enrichment.     

Stomatal behaviour, photosynthesis and transpiration under rising CO2

Definitions of the variables used and the units are given in Table 1

The literature reports enormous variation between species in the extent of stomatal responses to rising CO₂. This paper attempts to provide a framework within which some of this diversity can be explained. We describe the role of stomata in the short-term response of leaf gas exchange to increases in ambient CO₂ concentration by developing the recently proposed stomatal model of Jarvis & Davies (1998 ). In this model stomatal conductance is correlated with the functioning of the photosynthetic system so that the effects of increases in CO₂ on stomata are experienced through changes in the rate of photosynthesis in a simple and mechanistically transparent way. This model also allows us to consider the effects of evaporative demand and soil moisture availability on stomatal responses to photosynthesis and therefore provides a means of considering these additional sources of variation. We emphasize that the relationship between the rate of photosynthesis and the internal CO₂ concentration and also drought will have important effects on the relative gains to be achieved under rising CO₂.  

  Table 1 . Abbreviations     

17.

Plankton population dynamics: food web interactions and abiotic constraints   总被引：1，自引：0，他引：1  

Karl O. Rothhaupt 《Freshwater Biology》2000,45(2):105-109

1. In this introduction, I try to follow some developments in plankton ecology, how they have led to current research topics, and how the contributions in this issue of Freshwater Biology are related to these fields of research.
2. Due to several favourable features, such as small size, short generation time and a relatively homogeneous habitat, planktonic organisms remain ideal subjects for theoretical and experimental population ecology.
3. Important current research topics involve: (1) the control of plankton communities by external abiotic factors; (2) bottom-up (limitation by resources) and top-down (control by predators) effects in the food web; (3) the importance of dormant resting stages and benthic–pelagic coupling in plankton dynamics; (4) costs and benefits of the mixotrophic strategy, i.e. the ability to combine a phototrophic and a phagotrophic mode of nutrition.     

18.

Variation in phytoplankton composition and limnological features in a water–water ecotone of the Lower Paraná Basin (Argentina)     

Irina Izaguirre  Inés O'Farrell  Guillermo Tell 《Freshwater Biology》2001,46(1):63-74

1. Structure and dynamics of phytoplankton were studied at six sites in a reach of the Lower Paraná River floodplain, from the main course to a marginal shallow lake, connected through a stream. This system is defined as a water–water ecotone and thus, the aim of this work is to analyse the spatial changes of phytoplankton along the transitional system.
2. Temporal fluctuations in number of species and phytoplankton density were more pronounced in the river than in the shallow lake. Even though most species were ubiquitous, there was a clear segregation of the most abundant populations across the ecotone. Aulacoseira granulata var. granulata and other centric diatoms decreased towards the lentic system.
3. Transparency varied according to river discharge, being higher in the lake during high water phases, while the opposite behaviour was observed during dry periods. These fluctuations were related to precipitation and resuspension processes.
4. Nitrate concentration diminished towards the lake while soluble reactive phosphorus remained fairly homogeneous across the ecotone.     

19.

Response of a zooplankton community to the addition of unsaturated fatty acids: an enclosure study   总被引：1，自引：0，他引：1  

Maarten Boersma  Claus-Peter Stelzer 《Freshwater Biology》2000,45(2):179-188

1. The effect of the addition of emulsions with different fatty acid composition to a semi-natural zooplankton community was studied in enclosures.
2. The reactions of different taxa in the zooplankton community to the addition of the emulsions were different. The copepods showed almost no reaction, nor did the selective cladocerans ( Bosmina ) or rotifers ( Synchaeta or Polyarthra ). The non-selective filterfeeding cladocerans Daphnia and Ceriodaphnia , and the rotifer Keratella , showed responses to the addition of the emulsions.
3.  Keratella showed the highest density in the enclosures with high amounts of highly unsaturated fatty acids added, whereas both Daphnia and Ceriodaphnia reached the highest numbers in the enclosures where we added emulsions of saturated fatty acids only.
4. Our results suggest that different taxa may be limited by different factors, even though they use similar food sources. Hence, we conclude that it is very difficult to generalize on the limiting factors in aquatic systems.     

20.

高原鼠兔种群生产量生态学的研究Ⅲ.高原鼠兔生命率非密度制约与密度制约的种群动态趋势   总被引：1，自引：0，他引：1  

刘季科  聂海燕 《兽类学报》1992,12(2):139

本文以高原鼠兔(Ochotona curzoniae)自然种群生命表的统计参数为基础,根据非密度制约Leslie模型及具有密度制约反馈的标准Leslie修正模型,分别预测了该种群在1982-2001年间的发展趋势。在菲密度制约条件下,该种群呈指数增长。在密度制约存在肘,种群增长趋于平衡状态,且存滔率密度制约较繁殖率密度制约对种群的作用更大。存活率密度制约与非密度制约的年龄结构均为Leslie分布,繁殖率密度制约作用的种群稳定年龄分布更平均,其平衡状态的种群大小则由模型的参数决定。     

Plankton population dynamics: food web interactions and abiotic constraints

1. In this introduction, I try to follow some developments in plankton ecology, how they have led to current research topics, and how the contributions in this issue of Freshwater Biology are related to these fields of research.
2. Due to several favourable features, such as small size, short generation time and a relatively homogeneous habitat, planktonic organisms remain ideal subjects for theoretical and experimental population ecology.
3. Important current research topics involve: (1) the control of plankton communities by external abiotic factors; (2) bottom-up (limitation by resources) and top-down (control by predators) effects in the food web; (3) the importance of dormant resting stages and benthic–pelagic coupling in plankton dynamics; (4) costs and benefits of the mixotrophic strategy, i.e. the ability to combine a phototrophic and a phagotrophic mode of nutrition.     

Variation in phytoplankton composition and limnological features in a water–water ecotone of the Lower Paraná Basin (Argentina)

1. Structure and dynamics of phytoplankton were studied at six sites in a reach of the Lower Paraná River floodplain, from the main course to a marginal shallow lake, connected through a stream. This system is defined as a water–water ecotone and thus, the aim of this work is to analyse the spatial changes of phytoplankton along the transitional system.
2. Temporal fluctuations in number of species and phytoplankton density were more pronounced in the river than in the shallow lake. Even though most species were ubiquitous, there was a clear segregation of the most abundant populations across the ecotone. Aulacoseira granulata var. granulata and other centric diatoms decreased towards the lentic system.
3. Transparency varied according to river discharge, being higher in the lake during high water phases, while the opposite behaviour was observed during dry periods. These fluctuations were related to precipitation and resuspension processes.
4. Nitrate concentration diminished towards the lake while soluble reactive phosphorus remained fairly homogeneous across the ecotone.     

Response of a zooplankton community to the addition of unsaturated fatty acids: an enclosure study

1. The effect of the addition of emulsions with different fatty acid composition to a semi-natural zooplankton community was studied in enclosures.
2. The reactions of different taxa in the zooplankton community to the addition of the emulsions were different. The copepods showed almost no reaction, nor did the selective cladocerans ( Bosmina ) or rotifers ( Synchaeta or Polyarthra ). The non-selective filterfeeding cladocerans Daphnia and Ceriodaphnia , and the rotifer Keratella , showed responses to the addition of the emulsions.
3.  Keratella showed the highest density in the enclosures with high amounts of highly unsaturated fatty acids added, whereas both Daphnia and Ceriodaphnia reached the highest numbers in the enclosures where we added emulsions of saturated fatty acids only.
4. Our results suggest that different taxa may be limited by different factors, even though they use similar food sources. Hence, we conclude that it is very difficult to generalize on the limiting factors in aquatic systems.     

高原鼠兔种群生产量生态学的研究Ⅲ.高原鼠兔生命率非密度制约与密度制约的种群动态趋势

本文以高原鼠兔(Ochotona curzoniae)自然种群生命表的统计参数为基础,根据非密度制约Leslie模型及具有密度制约反馈的标准Leslie修正模型,分别预测了该种群在1982-2001年间的发展趋势。在菲密度制约条件下,该种群呈指数增长。在密度制约存在肘,种群增长趋于平衡状态,且存滔率密度制约较繁殖率密度制约对种群的作用更大。存活率密度制约与非密度制约的年龄结构均为Leslie分布,繁殖率密度制约作用的种群稳定年龄分布更平均,其平衡状态的种群大小则由模型的参数决定。