Seasonal dynamics of physical and biological processes in the central California Current System: A modeling study

A 3-D physical and biological model is used to study the seasonal dynamics of physical and biological processes in the central California Current System. Comparisons of model results with remote sensing and in situ observations along CalCOFI Line 67 indicate our model can capture the spatial variations of key variables (temperature, nutrients, chlorophyll, and so on) on annual mean and seasonal cycle. In the coastal upwelling system, it is the alongshore wind stress that upwells high nutrients to surface from 60 m and stimulates enhanced plankton biomass and productivity in the upwelling season. As a result, coastal species peak in the late upwelling period (May–July), and oceanic species reach the annual maxima in the oceanic period (August–October). The annual maximum occurs in the late upwelling period for new production and in the oceanic period for regenerated production. From the late upwelling period to the oceanic period, stratification is intensified while coastal upwelling becomes weaker. Correspondingly, the coastal ecosystem retreats from ～300 to ～100 km offshore with significant decline in chlorophyll and primary production, and the oceanic ecosystem moves onshore. During this transition, the decline in phytoplankton biomass is due to the grazing pressure by mesozooplankton in the 0–150 km domain, but is regulated by low growth rates in the 150–500 km offshore domain. Meanwhile, the growth rates of phytoplankton increase in the coastal waters due to deeper light penetration, while the decrease in offshore growth rates is caused by lower nitrate concentrations.     

Ammonium,nitrate and phytoplankton interactions in a freshwater tidal estuarine zone: potential effects of cultural eutrophication

Nitrate and ammonium are the most important nitrogen sources for phytoplankton growth. Differential utilization of inorganic nitrogenous compounds by phytoplankton has been observed and may have significant impacts on primary productivity at local scales. We used enrichment experiments with natural phytoplankton populations from the freshwater tidal zone of the Guadiana estuary, a coastal ecosystem increasingly subjected to anthropogenic influences, to study the effects of nitrate and ammonium on N-consumption and phytoplankton growth. In addition, we used combined additions of nitrate and ammonium to understand the inhibitory effect of ammonium over nitrate uptake. Ammonium concentrations in the freshwater tidal reaches of the Guadiana estuary throughout the sampling period were too low to exert an inhibitory effect on nitrate uptake or a toxic effect on phytoplankton growth. Nitrate was clearly the main nitrogen source for phytoplankton at the study site. Overall, nitrate seemed to become limiting at concentrations lower than 20 μM and N-limitation was particularly significant during summer. A trend of decreasing nitrate uptake with increasing ammonium concentrations and uptake suggested an overall preference for ammonium. However, preference for ammonium was group-specific, and it was observed mainly in green algae and cyanobacteria. In fact, cyanobacteria relied only on ammonium as their N-source. On the contrary, diatoms preferred nitrate, and did not respond to ammonium additions. The increasing eutrophication in the Guadiana estuary and particularly increased inputs of nitrogen as ammonium due to urban waste effluents may result in a shift in phytoplankton community composition, towards a dominance of cyanobacteria and green algae.     

鄱阳湖浮游植物时空变化特征及其对极端洪枯事件的响应

鄱阳湖作为中国最大的淡水湖泊,其水生态健康状态始终是人们关注的热点。近些年,鄱阳湖极端洪旱灾害频发,浮游植物生长受极端洪旱的影响发生了明显变化。为分析浮游植物时空变化特征、探究环境因子对鄱阳湖浮游植物影响机制以及极端洪枯事件对浮游植物的影响,利用结构方程模型(SEM)构建浮游植物与环境因子的影响路径模型,定量分析环境因子对浮游植物的影响程度。结果表明,鄱阳湖浮游植物以蓝藻、绿藻为主且有明显的季节特征,在7月丰水期浮游植物密度达到最高;由结构方程模型(SEM)可知,影响浮游植物密度最关键的因子为物理因子(水温>pH>透明度>溶解氧),其次为营养物质(总氮>硝态氮>总磷>磷酸盐),浮游植物对高温、高营养和高pH较偏好。2020年极端洪水和2022年极端高温干旱,浮游植物密度主要受水温、溶解氧、透明度等物理因子的影响;在影响较小的营养物质中,主要的限制性因素分别为磷和氮。与正常年份相比,极端洪水年鄱阳湖受入湖来水及降雨的增多,湖区水量急剧增加,“稀释”作用超过水温和透明度对浮游植物生长的促进作用造成浮游植物密度和生物量有所下降;在营养物质中,磷成为主要限制性因素。而极端高温干旱年受入湖来水的减少及湖水的快速蒸发,“浓缩”作用超过水温和透明度对浮游植物生长的抑制作用造成浮游植物密度和生物量显著增加,同时,营养物质对浮游植物的作用更加凸显。研究结果表明极端洪枯事件导致鄱阳湖浮游植物变化明显,确定其对浮游植物的影响机制,可以为极端洪枯事件下浮游植物监测、管理提供一定的理论基础。     

Modeling the response of primary production and sedimentation to variable nitrate loading in the Mississippi River plume

Increases in nitrate loading to the Mississippi River watershed during the last 50 years are considered responsible for the increase in hypoxic zone size in Louisiana-Texas shelf bottom waters. There is currently a national mandate to decrease the size of the hypoxic zone to 5000 km² by 2015, mostly by a 30% reduction in annual nitrogen discharge into the Gulf of Mexico. We developed an ecosystem model for the Mississippi River plume to investigate the response of organic matter production and sedimentation to variable nitrate loading. The nitrogen-based model consisted of nine compartments (nitrate, ammonium, labile dissolved organic nitrogen, bacteria, small phytoplankton, diatoms, micro- and mesozooplankton, and detritus), and was developed for the spring season, when sedimentation of organic matter from plume surface waters is considered important in the development of shelf hypoxia. The model was forced by physical parameters specified along the river-ocean salinity gradient, including residence time, light attenuation by dissolved and particulate matter, mixed layer depth, and dilution. The model was developed using measurements of biological biomasses and nutrient concentrations across the salinity gradient, and model validation was performed with an independent dataset of primary production measurements for different riverine NO₃ loads. Based on simulations over the range of observed springtime NO₃ loads, small phytoplankton contributed on average 80% to primary production for intermediate to high salinities (>15), and the main contributors to modeled sedimentation at these salinities were diatom sinking, microzooplankton egestion, and small phytoplankton mortality. We investigated the impact of limiting factors on the relationship between NO₃ loading and ecosystem rates. Model results showed that primary production was primarily limited by physical dilution of NO₃, followed by abiotic light attenuation, light attenuation due to mixing, and diatom sinking. Sedimentation was mainly limited by the first three of these factors. Neither zooplankton grazing or plume residence times acted as limiting factors of ecosystem rates. Regarding nutrient reductions to the watershed, simulations showed that about half of the percent decrease in NO₃ load was reflected in decreased plume sedimentation. For example, a 30% decrease in NO₃ load resulted in a 19% decrease in average plume primary production and a 14% decrease in sedimentation. Finally, our model results indicated that the fraction of primary production exported from surface waters is highly variable with salinity (7–87%), a finding which has important implications for predictive models of hypoxic zone size that assume a constant value for this ratio.     

Calibration of a coupled biological-physical model for prediction in a coastal inlet

We describe a numerical forecast system designed for prediction of physical and biological dynamics of a coastal inlet. It is based on a coastal ocean observatory that was located at Lunenburg Bay, Nova Scotia, Canada. Biological, chemical, optical, and physical measurements were collected from instrumented moorings, weekly sampling and detailed surveys from 2002 through 2007. Here we present a framework for calibration and evaluation of an ecosystem model using data from the summer of 2007. A three-dimensional hydrodynamic model was coupled to a simple biological (Nutrients-Phytoplankton-Detritus) model; a simple model was used so results could be compared directly to observed biological and chemical variables using skill scores as a first step toward data-assimilation modeling. As a complement to this analysis, variability of model output, e.g., the nutrient limitation term, was examined to understand the modeled biological response to the simulated physical environment. Skill scores based on variances in observed and simulated time-series of biological components were also investigated. Coastal upwelling/downwelling simulated through this model has been found to increase modeled biological activity in the bay. Also model skill in reproducing the observed patterns in nutrients and phytoplankton has been increased due to the restoring conditions for biology set up at the open ocean boundaries of the bay.     

Chemical composition of Coscinodiscus wailesii and the implication for nutrient ratios in a coastal water, Seto Inland Sea, Japan

Chemical compositions of Coscinodiscus wailesii were determined for four samples of natural cells. Results revealed that the cellular Si:N ratio of C. wailesii cell was 2.4:1.0-5.2:1.0. The impacts of C. wailesii on surrounding coastal water were evaluated from hydrographic observations, in which C. wailesii cell density, nutrients concentrations and temperature were monitored from November 2001 to February 2005 in Harima-Nada, the Seto Inland Sea, Japan. In low temperature periods, from October to December, two peaks of C. wailesii cell density were observed and nutrient concentrations were depleted. The draw-down ratio of Si(OH)(4) and DIN (Si(OH)(4):DIN ratio) in the water column were similar to the cellular Si:N ratio of C. wailesii cells, which have high Si contents. In addition the effects of different Si(OH)(4):DIN ratio were determined for in situ bottle incubation experiments. In the culture experiments, picoplankton (0.2-2.0 microm) which consisted of small flagellates became dominant under low Si(OH)(4):DIN ratios. These results suggested that the sizes distribution shifted to small size and the phytoplankton community was changed to small flagellates after the C. wailesii bloom. These changes would influence predators at higher trophic levels in its coastal ecosystem.     

Elevated ammonium concentrations from wastewater discharge depress primary productivity in the Sacramento River and the Northern San Francisco Estuary

Primary production in the Northern San Francisco Estuary (SFE) has been declining despite heavy loading of anthropogenic nutrients. The inorganic nitrogen (N) loading comes primarily from municipal wastewater treatment plant (WTP) discharge as ammonium (NH(4)). This study investigated the consequences for river and estuarine phytoplankton of the daily discharge of 15 metric tons NH(4)-N into the Sacramento River that feeds the SFE. Consistent patterns of nutrients and phytoplankton responses were observed during two 150-km transects made in spring 2009. Phytoplankton N productivity shifted from NO(3) use upstream of the WTP to productivity based entirely upon NH(4) downstream. Phytoplankton NH(4) uptake declined downstream of the WTP as NH(4) concentrations increased, suggesting NH(4) inhibition. The reduced total N uptake downstream of the WTP was accompanied by a 60% decline in primary production. These findings indicate that increased anthropogenic NH(4) may decrease estuarine primary production and increase export of NH(4) to the coastal ocean.     

Variations of phytoplankton community structure related to water quality trends in a tropical karstic coastal zone

Phytoplankton community structure in coastal areas is a result of various environmental factors such as nutrients, light, grazing, temperature, and salinity. The Yucatan Peninsula is a karstic tropical region that is strongly influenced by submerged groundwater discharge (SGD) into the coastal zone. Phytoplankton community structure and its relationship with regional and local water quality variables were studied in four ports of the northwestern Yucatan Peninsula. Water quality was strongly related to SGD, and variations in phytoplankton community structure were related to local nutrient loading and hydrographic conditions, turbulence, and human impacts. Our study provides an ecological baseline for the Yucatan Peninsula and serves as a basis for establishing monitoring programs to predict changes at sites with high hydrological variation and in developing an early alert system for harmful toxic algal blooms.     

Phytoplankton biomass size structure and its regulation in the Southern Yellow Sea (China): Seasonal variability

Phytoplankton size structure plays a significant role in controlling the carbon flux of marine pelagic ecosystems. The mesoscale distribution and seasonal variation of total and size-fractionated phytoplankton biomass in surface waters, as measured by chlorophyll a (Chl a), was studied in the Southern Yellow Sea using data from four cruises during 2006–2007. The distribution of Chl a showed a high degree of spatial and temporal variation in the study area. Chl a concentrations were relatively high in the summer and autumn, with a mean of 1.42 and 1.27 mg m⁻³, respectively. Conversely, in the winter and spring, the average Chl a levels were only 0.98 and 0.99 mg m⁻³. Total Chl a showed a clear decreasing gradient from coastal areas to the open sea in the summer, autumn and winter cruises. Patches of high Chl a were observed in the central part of the Southern Yellow Sea in the spring due to the onset of the phytoplankton bloom. The eutrophic coastal waters contributed at least 68% of the total phytoplankton biomass in the surface layer. Picophytoplankton showed a consistent and absolute dominance in the central region of the Southern Yellow Sea (>40%) in all of the cruises, while the proportion of microphytoplankton was the highest in coastal waters. The relative proportions of pico- and nanophytoplankton decreased with total biomass, whereas the proportion of the micro-fraction increased with total biomass. Relationships between phytoplankton biomass and environmental factors were also analysed. The results showed that the onset of the spring bloom was highly dependent on water column stability. Phytoplankton growth was limited by nutrient availability in the summer due to the strong thermocline. The combined effects of P-limitation and vertical mixing in the autumn restrained the further increase of phytoplankton biomass in the surface layer. The low phytoplankton biomass in winter was caused by vertical dispersion due to intense mixing. Compared with the availability of nutrients, temperature did not seem to cause direct effects on phytoplankton biomass and its size structure. Although interactions of many different environmental factors affected phytoplankton distributions, hydrodynamic conditions seemed to be the dominant factor. Phytoplankton size structure was determined mainly by the size-differential capacity in acquiring resource. Short time scale events, such as the spring bloom and the extension of Yangtze River plume, can have substantial influences, both on the total Chl a concentration and on the size structure of the phytoplankton.     

城市湖泊春季绿藻水华特征及其影响因素——以宁波月湖为例

城市湖泊富营养化问题日趋严峻,以往对水华的研究多集中于大型自然淡水湖库,而对小型城市浅水湖泊的水华动态相对较少.以宁波月湖为研究对象,探讨水华暴发期间浮游植物变化特征及与影响因子之间的关系,以期判别影响城市湖泊水华的主控因子.月湖水华期间营养盐水平处于中富营养至极端富营养之间,此次共检出浮游植物8门61属,藻种组成以绿藻门（51.79%）和硅藻门（21.43%）为主,各点位浮游植物生长主要受水温、光照驱动,经历了隐藻门→硅藻门→绿藻门→蓝藻门的演替.水华种为雷氏衣藻（Chlamydomonas reinhardtii）,总藻密度最高达到1.55×10⁸ cells/L,水华暴发后各点位衣藻属比例升高（最高达到81.10%）,群落结构呈现单一化特征.通过Pearson相关性分析和RDA分析发现衣藻属生长与水温、pH、总磷浓度均呈显著正相关,春季气温回升、天气持续晴好,城市浅水湖泊高营养盐负荷、水体流动性差等特点为带鞭毛的衣藻属提供了适宜的生存条件,在环境条件均适宜的情况下拥有最大生长潜力的衣藻属在营养盐、光照等竞争中生长速率明显优于其他藻种,从而发生绿藻水华.     

Responses of a coastal phytoplankton community to increased nutrient input from the Changjiang (Yangtze) River

Nutrient input from the Changjiang River (Yangtze River) has been increasing dramatically since the 1960s. At the mouth of the Changjiang River, the nitrate concentration has increased about three-fold in 40 years, from 20.5 μmol/L in the 1960s to 59.1 μmol/L in the 1980s and to 80.6 μmol/L in 1990–2004. Phosphate concentration increased by a factor of 30%, from 0.59 μmol/L in the 1980s to 0.77 μmol/L in 1990–2004. The increasing nitrate input has arisen mostly from the mid and lower reaches of the Changjiang River, where the river meets one of the most strongly developed agriculture areas in China. Responses of the coastal phytoplankton community to the increasing nutrient inputs are also seen in the available monitoring data. First, a trend of increasing phytoplankton standing stock from 1984 to 2002 appeared in the Changjiang River estuary and adjacent coastal waters, especially in late spring. Secondly, the proportion of diatoms in the whole phytoplankton community showed a decreasing trend from about 85% in 1984 to about 60% in 2000. Finally, red tides/harmful algal blooms increased dramatically in this area in terms of both number and scale. About 30–80 red tide events were recorded each year from 2000 to 2005 in the East China Sea. The scale of some blooms has been in excess of 10,000 km².     

Temporal and spatial variations in nutrient stoichiometry and regulation of phytoplankton biomass in Hong Kong waters: Influence of the Pearl River outflow and sewage inputs

In 2001, the Hong Kong government implemented the Harbor Area Treatment Scheme (HATS) under which 70% of the sewage that had been formerly discharged into Victoria Harbor is now collected and sent to Stonecutters Island Sewage Works where it receives chemically enhanced primary treatment (CEPT), and is then discharged into waters west of the Harbor. The relocation of the sewage discharge will possibly change the nutrient dynamics and phytoplankton biomass in this area. Therefore, there is a need to examine the factors that regulate phytoplankton growth in Hong Kong waters in order to understand future impacts. Based on a historic nutrient data set (1986–2001), a comparison of ambient nutrient ratios with the Redfield ratio (N:P:Si = 16:1:16) showed clear spatial variations in the factors that regulate phytoplankton biomass along a west (estuary) to east (coastal/oceanic) transect through Hong Kong waters. Algal biomass was constrained by a combination of low light conditions, a rapid change in salinity, and strong turbulent mixing in western waters throughout the year. Potential stoichiometric Si limitation (up to 94% of the cases in winter) occurred in Victoria Harbor due to the contribution of sewage effluent with high N and P enrichment all year, except for summer when the frequency of stoichiometric Si limitation (48%) was the same as P, owing to the influence of the high Si in the Pearl River discharge. In the eastern waters, potential N limitation and N and P co-limitation occurred in autumn and winter respectively, because of the dominance of coastal/oceanic water with low nutrients and low N:P ratios. In contrast, potential Si limitation occurred in spring and a switch to potential N, P and Si limitation occurred in eastern waters in summer. In southern waters, there was a shift from P limitation (80%) in summer due to the influence of the N-rich Pearl River discharge, to N limitation (68%) in autumn, and to N and P co-limitation in winter due to the dominance of N-poor oceanic water from the oligotrophic South China Sea. Our results show clear temporal and spatial variations in the nutrient stoichiometry which indicates potential regulation of phytoplankton biomass in HK waters due to the combination of the seasonal exchange of the Pearl River discharge and oceanic water, sewage effluent inputs, and strong hydrodynamic mixing from SW monsoon winds in summer and the NE monsoon winds in winter.     

Temporal and spatial variations in nutrient stoichiometry and regulation of phytoplankton biomass in Hong Kong waters: influence of the Pearl River outflow and sewage inputs

In 2001, the Hong Kong government implemented the Harbor Area Treatment Scheme (HATS) under which 70% of the sewage that had been formerly discharged into Victoria Harbor is now collected and sent to Stonecutters Island Sewage Works where it receives chemically enhanced primary treatment (CEPT), and is then discharged into waters west of the Harbor. The relocation of the sewage discharge will possibly change the nutrient dynamics and phytoplankton biomass in this area. Therefore, there is a need to examine the factors that regulate phytoplankton growth in Hong Kong waters in order to understand future impacts. Based on a historic nutrient data set (1986-2001), a comparison of ambient nutrient ratios with the Redfield ratio (N:P:Si=16:1:16) showed clear spatial variations in the factors that regulate phytoplankton biomass along a west (estuary) to east (coastal/oceanic) transect through Hong Kong waters. Algal biomass was constrained by a combination of low light conditions, a rapid change in salinity, and strong turbulent mixing in western waters throughout the year. Potential stoichiometric Si limitation (up to 94% of the cases in winter) occurred in Victoria Harbor due to the contribution of sewage effluent with high N and P enrichment all year, except for summer when the frequency of stoichiometric Si limitation (48%) was the same as P, owing to the influence of the high Si in the Pearl River discharge. In the eastern waters, potential N limitation and N and P co-limitation occurred in autumn and winter respectively, because of the dominance of coastal/oceanic water with low nutrients and low N:P ratios. In contrast, potential Si limitation occurred in spring and a switch to potential N, P and Si limitation occurred in eastern waters in summer. In southern waters, there was a shift from P limitation (80%) in summer due to the influence of the N-rich Pearl River discharge, to N limitation (68%) in autumn, and to N and P co-limitation in winter due to the dominance of N-poor oceanic water from the oligotrophic South China Sea. Our results show clear temporal and spatial variations in the nutrient stoichiometry which indicates potential regulation of phytoplankton biomass in HK waters due to the combination of the seasonal exchange of the Pearl River discharge and oceanic water, sewage effluent inputs, and strong hydrodynamic mixing from SW monsoon winds in summer and the NE monsoon winds in winter.     

Lake Kinneret phytoplankton: Stability and variability during twenty years (1970–1989)

The general features of phytoplankton seasonal succession, abundance and distribution in Lake Kinneret, as based on observations from 1970 through 1989, are summarised. Throughout this period of observation, the large, thecate dinoflagellatePeridinium gatunense formed an annual, late winter to early spring bloom resulting in very high standing stock levels. The dominance of these dinoflagellates has a profound impact upon the lake ecosystem. In the summer and fall the phytoplankton assemblage consisted mainly of nanoplanktonic green and blue-green algae and diatoms. Picophytoplankton, mainly picocyanobacteria, were present in low numbers during the dinoflagellate bloom but reached maximum abundance (10⁵ cells · ml^–1) in the epilimnion during the summer and fall. Within a given year, chlorophyll concentrations correlated well with estimates of wet weight biomass, derived from microscope counts. However, interannual averages of chlorophyll did not correlate closely with those for wet weight biomass. Both wet weight biomass and chlorophyll standing stocks fluctuated more than 2 fold from 1970 through 1989 but no extreme, long-term, continuous trend of increase or decrease was observed. Thus, phytoplankton has remained relatively stable although there has been a significant rise in the levels of summer-fall biomass since 1981. The main factor responsible for this may have been increased available phosphorus; the abundance of phytoplankton did not show any clear, long-term relation to that of herbivorous zooplankton.     

Mussel farming as a nutrient reduction measure in the Baltic Sea: consideration of nutrient biogeochemical cycles

Nutrient loads from the land to the sea must be reduced to combat coastal eutrophication. It has been suggested that further mitigation efforts are needed in the brackish Baltic Sea to decrease nutrients, especially in eutrophic coastal areas. Mussel farming is a potential measure to remove nutrients directly from the sea. Mussels consume phytoplankton containing nitrogen (N) and phosphorus (P); when the mussels are harvested these nutrients are removed from the aquatic system. However, sedimentation of organic material in faeces and pseudo-faeces below a mussel farm consumes oxygen and can lead to hypoxic or even anoxic sediments causing an increased sediment release of ammonium and phosphate. Moreover, N losses from denitrification can be reduced due to low oxygen and reduced numbers of bioturbating organisms. To reveal if mussel farming is a cost-effective mitigation measure in the Baltic Sea the potential for enhanced sediment nutrient release must be assessed.     

Intra-annual variability in nutrients in the Godavari estuary,India

Daily variations in nutrients were monitored for 15 months (September 2007–November 2008) in the Godavari estuary, Andhra Pradesh, India, at two fixed locations. River discharge has significant influence on nutrients loading to the estuary, which peaks during June–August (peak discharge period; monsoon) whereas exchanges at the sediment–water interface, groundwater and rainwater contribute significantly during other period. Despite significant amount of nutrients brought by discharge to the study region, phytoplankton biomass, in terms of chlorophyll-a (Chl a), did not increase significantly due to high suspended load and shallow photic depth. Nutrients showed downward gradient towards downstream of the estuary from upstream due to dilution by nutrient poor seawater and biological uptake. The N:P ratios were higher than Redfield ratio in both upstream and downstream of the estuary during no discharge period suggesting PO₄ to be a limiting nutrient for phytoplankton production, at levels <0.10 μmol L⁻¹. On the other hand, Si:N ratios were always more than unity during entire study period at both the stations indicating that Si(OH)₄ is not a limiting nutrient. Our results suggest that suspended matter limits phytoplankton biomass during peak discharge period whereas PO₄ during no discharge period.     

火烧对洞庭湖湿地荻(Miscanthus sacchariflorus)和苔草(Carex brevicuspis)群落土壤性质的影响

火烧作为调控因子,对植物群落结构和生态系统功能具有重要影响,但在湖泊湿地中研究较少.通过野外调查取样与实验室分析,探讨火烧对洞庭湖湿地主要群落类型——荻(Miscanthus sacchariflorus)和苔草(Carex brevicuspis)土壤化学性质的影响.结果表明:火烧后,苔草群落土壤硝态氮含量显著减少64.6%,有机质含量增加26.3%;而荻群落土壤与之相反,硝态氮含量增加186.9%,有机质含量减少22.9%.火烧后,苔草群落的全氮、铵态氮、全碳和全磷含量均显著增加,分别增加了75.4%、36.3%、102.7%和76.9%,而荻群落土壤与对照组间无显著差异.总体上,火烧对荻群落土壤养分影响不大,可作为芦苇场的一种管理方式,但火烧促进苔草群落土壤养分释放,有助于苔草群落提前萌芽和生长,并引起牲畜牧食增加.