基于全局灵敏度分析的浒苔生长影响参数研究
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国家自然科学基金资助项目(41206111)


Factors influencing Ulva prolifera growth revealed by model based on global sensitivity analysis
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    摘要:

    近年来,以浒苔为主的绿潮灾害频发。构建了浒苔生长模型,以定量分析浒苔生长过程,探索浒苔爆发机理。参数不确定性是模型不确定性的主要来源,以参数灵敏度分析为基础的参数优化有利于提高模型精度,采用Morris方法对模型涉及的主要参数进行了全局灵敏度分析,以研究浒苔生长的主要影响参数。不同于其他有关大型绿藻的模型模拟,模型同时考虑了幼体浒苔和成熟浒苔的生物量变化,并修正了营养盐限制函数以及温度计算函数。全局性的参数灵敏度分析结果表明,最适温度(Topt)、光合作用最适光强(Is)、最大发芽率(Gmax)、浒苔生长所需的氮含量的半饱和系数(kqn)、最大氮摄取率(Vmaxn)这5个参数在浒苔生长模型中具有较大灵敏性。其中,Topt影响最大,IsVmaxn其次,说明浒苔生长主要受温度光照和氮含量限制。相较于局部灵敏度分析仅关注单个参数变化、依赖于初值选取等缺陷,全局灵敏度分析同时从各个参数的取值范围上分析参数对模型结果的影响,能揭示参数之间相互作用的影响。此外,灵敏度较大的参数往往和其他参数之间存在较大相关性。

    Abstract:

    Recently, green tides dominated by Ulva prolifera have had a major impact on coastal ecosystems of China, as a result of the algae's high surface area to volume ratio, high rate of nutrient uptake, low nutrient half-saturation coefficient, and restriction of other algae. Here, a U. prolifera growth model was developed to analyze the growth process and key limiting factors of U. prolifera. To date, ecological modeling related to green macroalgae has mainly concentrated on variability in adult plant biomass; in this study, biomass of early life stages and adult plants were considered separately to clarify the growth pattern of U. prolifera. In addition, nutrient limitation and temperature functions in the growth model were adapted to the environment of coastal China. The model was established with STELLA, and the simulation results reveal the growth pattern of U. prolifera. Parameter uncertainty is the basis of model uncertainty and thus should be assessed; parameter optimization based on sensitivity analysis could improve the precision of the growth model. Sensitivity analysis is also an important tool to improve marine ecological models. In this study, global sensitivity analysis of the Morris method based on statistical sampling was implemented in the U. prolifera growth model. Compared with local sensitivity analysis methods, global sensitivity analysis has the advantage of assessing correlations among parameters and of analyzing the sensitivity of all parameters simultaneously. Compared with other global sensitivity analysis method, the Morris method is efficient. Nevertheless, adequate sampling repetition must be performed. Parameters are sampled from the entire defined domain and are ranked according to the mean and standard deviation of elementary effects to assess their global sensitivity and qualitative correlation. Thus, parameter optimization strategies can be established to improve the precision of high-ranking sensitive parameters while insensitive parameters are used as empirical values. Here, the parameters were sampled 10,000 times to reduce random error, and the mean and standard deviation of elementary effects were ranked with radar graphs. The sensitivity analysis results showed that optimum temperature for growth (Topt), optimum light intensity for photosynthesis (Is), maximum germination rate (Gmax), nitrogen half-saturation constant for growth (kqn), and maximum nitrogen uptake rate (Vmaxn) were sensitive in the growth model, which meant that U. prolifera is mainly limited by temperature, light intensity, and nitrogen. The precision of these five parameters should be improved by further parameter optimization; maximum growth rate (μmax) and reproduction rate-biomass lost by sporulation (Reprod_rate) were not sensitive and could be kept as empirical values. Local and global sensitivity analyses were compared, which revealed that global sensitivity analysis was much reliable because the sensitivity results were not affected by the initial parameter values. Correlation analysis showed that the sensitive parameters were correlated with other parameters.

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刘永志,沈程程,石洪华,郭振.基于全局灵敏度分析的浒苔生长影响参数研究.生态学报,2016,36(13):4178~4186

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