使用SSI模型描述昆虫温发育速率的SAS程序

Sharpe-Schoolfield-Ikemoto(SSI)模型是用来描述温度对昆虫发育速率影响的一个重要数学方程,它既能够很好地拟合温发育速率的数据,又能提供若干个重要温参数,特别是内禀最适发育温度(TΦ)。此温度反映了昆虫生存的最佳温环境。然而,由于SSI模型的参数过多,结构显得较为复杂,给实际的数据拟合带来了困难。一个基于R平台的软件包已经被开发出来,专门用于执行SSI模型的参数拟合。但目前由于国内许多研究者青睐于使用SAS统计软件,所以本文提供了一个基于SAS统计软件的程序,用以快速拟合SSI模型的参数。通过拟合棉铃虫Helicoverpa armigera Hübner蛹期的温发育速率数据,发现此SAS程序具有很好的拟合效果,同时进一步证明了SSI模型对描述温度对昆虫发育速率影响的有效性。     

昆虫发育模型研究进展

<正> 昆虫发育模型描述温度对昆虫发育过程的影响。目前,国内外广泛使用的昆虫发育模型归纳起来有3类:(1)发育速率模型;(2)发育时间分布模型;(3)随机发育模型。下面简要介绍各类模型的特点及应用情况。     

昆虫发育过程中的速率累加效应对其日均发育率的影响

昆虫发育速率V与温度T之间呈S型曲线关系。在确定适合昆虫发育的变温范围的基础上,利用萝卜蚜的试验数据,通过计算机模拟,探讨了“速率累加”过程通过V与T之间的曲线关系在各种变温条件下对昆虫日均发育速率所产生的影响。结果表明,在排除温度波动本身可改变瞬时发育速率的前提下,这种速率累加效应导致在低温区内变温下的发育比在恒温下快,在高温区内则相反,且温度变幅越广差异就越大。文中指出,由于日均发育速率会依温度的变化方式和幅度不同而发生改变,而依据恒温速率曲线可计算出各种变温下的发育进度,故对于预测昆虫在适温区发育进度而言,恒温试验结果比变温试验结果具有更广泛的适用性。     

昆虫种群动态模拟模型

昆虫是动物界中最大的类群,与人类有着密切的利害关系。对昆虫的数量预测与符合经济和生态规律的管理,一直都被国内外列入重点研究课题。种群动态模拟是害虫管理中重要的基础工作。近十年来,关于昆虫种群动态模型的理论和实验研究进展迅速。现分别从单种种群和多种种群两个方面对国内外近些年来昆虫种群动态模拟模型的研究进展进行了概括和总结。单种种群从两个方面阐述:一是最基本的种群动态模拟模型Log istic方程的研究成果,包括方程的修正、参数的拟合与最优捕获策略等;另一个方面是对种群动态模拟常用的矩阵模型的概述,主要介绍不等期年龄组、矩阵维数的变化、矩阵维数与历期的关系、个体之间的发育差异以及发育速率差异等等对昆虫种群动态模型的影响。多种群主要从建模和模型应用两个部分对国内外研究成果进行综述。最后,对种群动态模拟模型研究的发展方向做了深入地讨论,即在原有的数据采集工作的基础上,使用面向对象程序设计语言,把各种要素包括各种物种及各种环境条件抽象成类,用消息传递来表示昆虫种群内个体与个体、昆虫种群与环境之间的相互作用,再结合先进的数学算法,建立一个直观的、操作简单的昆虫种群动态模型库,使模型结构与现实世界有最大的相似性。这样就可以实现昆虫种群动态的可视化、立体化、实时化和精确化的监测及预测。     

用非线性模型估测恒温和变温下棉铃虫蛹的发育率

为了深入分析和探讨昆虫发育与环境温度的关系, 在恒温（15~37℃）和交替变温（12/18~34/40℃）下测定了棉铃虫Helicoverpa armigera蛹的发育历期(d),分别用线性模型和非线性模型（Logan模型﹑Lactin模型和王氏模型）拟合其发育率(1/d)数据。结果表明,这3个非线性模型能更准确地描述发育率与温度之间的曲线关系,判定系数（R2）在0.9878～0.9991之间。对全部观测数据的进一步研究表明,只要有6个分布合适的观测数据,就可以用这些非线性模型获得相当满意的估测效果。如果缺乏高温下的测定数据,用非线性模型预测的昆虫发育率可能失真。分析了蛹在恒温和变温下发育率差异的可能原因,讨论了应用这3个非线性模型预测蛹期发育的优点和缺点,指出用非线性模型取代线性日·度模型进行害虫发生预测和益虫饲养管理的合理性和必要性。     

作物发育温度非线性效应Beta模型的特征分析

Beta模型在反映温度对作物发育非线性效应方面被广泛采用.为使该函数满足在设定的最适温度下函数值取值最大的条件,得到Beta模型的特殊形式.分析认为,该模型满足温度对作物发育影响的三基点规律、较好地反映发育速率对温度变化的响应特征、温度三基点在模型中应比较明确的3个规范性要求,具有较强的变化特征表达能力,可以近似表达二次函数、高斯函数等函数的变化.分析了Beta模型与积温法计算结果的关系.通过对Beta模型一阶和二阶导数的分析,明确了其变化特征,指出已有研究对参数P生物学意义解释的不妥之处,提出作物发育速度对温度的非线性响应是感温性的本质特征的观点,并从新的视角肯定了参数P在一定程度上可以作为作物感温性强弱的度量.     

温度对咖啡豆象实验种群发育和繁殖的影响

为探讨温度对咖啡豆象Araecerus fasciculatus实验种群生长发育和繁殖参数的影响, 以及昆虫生命活动与环境因子之间的关系, 为其种群数量控制提供依据, 本研究在17, 20, 23, 26, 29和32℃下, 研究咖啡豆象各虫态的发育与繁殖; 用线性回归、Logistic模型及“王-兰-丁”模型对温度与咖啡豆象各虫态之间的数学关系进行拟合; 通过最小二乘法计算了咖啡豆象的发育起点温度和有效积温。结果表明: 17~29℃之间, 随着温度的升高, 咖啡豆象幼虫的发育速率逐渐增大, 至32℃, 发育速率稍微降低。温度为27.43℃时, 咖啡豆象幼虫有理论最大发育速率0.493 d^-1。卵、幼虫、蛹和卵-蛹期的发育起点温度分别为9.60, 11.48, 7.60和10.55℃, 有效积温分别为135.51, 523.57, 157.09和828.63日度。温度对咖啡豆象实验种群发育和繁殖有显著影响, 发育起点温度和有效积温较低是该虫危害严重的主要原因之一。本研究拟合数学模型可为该害虫的发生危害预测及综合管理提供技术支撑。     

变温促进昆虫发育的酶学解释

本文关于如何评价变温对昆虫发育速率影响的讨论表明,在比较两点(T_1,T_2)变温与恒温下的发育速率时,相应恒温下的发育速率为R_(con)(T_1,T_2)=(R_(con)(T_1) R_(con)(T_2))/2,而不是T_1、T_2平均值所对应的发育速率R_(con)(T_1,T_2)=R_(con)(T_1 T_2/2)。 在探讨发育速率与温度关系时,认为,(1)发育速率实际上是代谢速率的表现,从本质上说是由昆虫体内无数生化反应所决定的。每步生化反应所需酶不同。(2)不同酶,其活力与温度的关系不同。即使同种昆虫,体内不同种类酶的最大活力温度不同。因此,在不同温度下,代谢链中起限速作用的酶不同。发育速率与温度的关系实质上是代谢链中所有酶与温度关系的综合表现。     

基于支持向量回归的棉铃虫蛹发育历期估测

温度与发育速率关系模拟是昆虫学研究的一个重要内容, 传统基于经验风险最小的非线性参数模型(Logan模型、Lactin模型和王氏模型)存在诸多弊端。本文基于结构风险最小的改进支持向量回归（SVR）研究温度与棉铃虫Helicoverpa armigera蛹发育历期关系。结果表明: 与传统非线性模型相比, SVR模型性能优异; 基于全部92个样本, SVR模型拟合和留一法预测的决定系数R²分别为0.998和0.996, 估测的蛹期三基点温度更可信。从全部样本中依温度均匀选取部分样本实施独立预测, 当训练集为20个样本时, SVR模型独立预测的R²为0.981, 优于传统非线性模型中独立预测最佳的Lactin模型（R²=0.958）; 当训练集进一步减少到12个样本时, SVR模型的R²仅降低到0.964, 而传统非线性模型均已不适用。结果提示SVR模型在小样本情况下较传统非线性模型优势明显, 在昆虫发育历期估测建模中有应用前景。     

桃蚜、萝卜蚜发育速率在恒温和变温下的变化规律研究

本文报道桃蚜(Myzus persicae)萝卜蚜(Lipaphis erysimi)在一系列恒温、变温下的发育速率,结果表明:(1)恒温下发育速率与温度关系遵循有下上限的logistic曲线;(2)除了在恒温下随温度升高死亡率急剧增加、发育速率迅速下降的高温区(桃蚜:>28℃,萝卜蚜:>30℃)外,温度波动不改变发育速率与温度的函数关系;(3)变温下,在上述高温区内的发育速率仍基本按经典的logistic曲线随温度升高而上升,但在一温度下的发育速度又随停留时间延长而逐渐下降;在恒温下只能完成部分虫期发育的低温区,变温下发育速率亦接近经典的logistic曲线。文中根据上述结果提出了一个昆虫发育率在恒温和变温下变化规律的初步模型,并发现文献中有大量的基本符合这一模型例证的。     

Food limitation and insect outbreaks: complex dynamics in plant-herbivore models

1. The population dynamics of many herbivorous insects are characterized by rapid outbreaks, during which the insects severely defoliate their host plants. These outbreaks are separated by periods of low insect density and little defoliation. In many cases, the underlying cause of these outbreaks is unknown. 2. Mechanistic models are an important tool for understanding population outbreaks, but existing consumer-resource models predict that severe defoliation should happen much more often than is seen in nature. 3. We develop new models to describe the population dynamics of plants and insect herbivores. Our models show that outbreaking insects may be resource-limited without inflicting unrealistic levels of defoliation. 4. We tested our models against two different types of field data. The models successfully predict many major features of natural outbreaks. Our results demonstrate that insect outbreaks can be explained by a combination of food limitation in the herbivore and defoliation and intraspecific competition in the host plant.     

Effects of temperature on development, survival and reproduction of insects: experimental design, data analysis and modeling

The developmental response of insects to temperature is important in understanding the ecology of insect life histories. Temperature-dependent phenology models permit examination of the impacts of temperature on the geographical distributions, population dynamics and management of insects. The measurement of insect developmental, survival and reproductive responses to temperature poses practical challenges because of their modality, variability among individuals and high mortality near the lower and upper threshold temperatures. We address this challenge with an integrated approach to the design of experiments and analysis of data based on maximum likelihood. This approach expands, simplifies and unifies the analysis of laboratory data parameterizing the thermal responses of insects in particular and poikilotherms in general. This approach allows the use of censored observations (records of surviving individuals that have not completed development after a certain time) and accommodates observations from temperature transfer treatments in which individuals pass only a portion of their development at an extreme (near-threshold) temperature and are then placed in optimal conditions to complete their development with a higher rate of survival. Results obtained from this approach are directly applicable to individual-based modeling of insect development, survival and reproduction with respect to temperature. This approach makes possible the development of process-based phenology models that are based on optimal use of available information, and will aid in the development of powerful tools for analyzing eruptive insect population behavior and response to changing climatic conditions.     

Host selection in insects: reproductive interference shapes behavior of ovipositing females

In nature, closely related species often utilize different host species, but it is still unclear what factors contribute to the evolution and maintenance of such diversified host selection. In this review, I describe how negative interspecific mating interactions (reproductive interference) can shape host selection by animals, focusing mainly on phytophagous and predatory insects. First, I explain an important premise of this hypothesis, which is that the adult reproductive site is the same as the feeding site for the offspring. Next, I describe several mathematical models and well-studied empirical systems to show that reproductive interference can sufficiently drive and maintain different host selection between phylogenetically related species. Then, I argue for the first time that reproductive interference can cause an oviposition preference in insects that is not optimal for the survival and development of the offspring, as a result of maternal adaptive behavior that maximizes the mother's own fitness. Furthermore, I argue that in insects, reproductive interference probably shapes oviposition behavior before the female alights on the host (e.g., habitat preference), without affecting post-alighting decision making. I would like to emphasize that these two arguments represent the novel approach to clarify the unrevealed pattern of complex insect oviposition behavior.     

Common-intersection hypothesis of development rate lines of ectotherms within a taxon revisited

The relationship between development rate of an ectotherm and temperature in experiments where insects or mites are reared under several constant temperatures, while non-linear over the entire range, can be approximately described by a line in the mid-temperature range. It was hypothesized that the development rate lines of ectotherms within a taxon such as a family would have a common intersection. Thus for a given temperature, the development time should be the same for all the species within any specified taxon. In the current study, the data on temperature-dependent development rates of species of the Aphididae and Tetranychidae families were used to test the validity of the common-intersection hypothesis. The hypothesis was supported with either dataset used. In addition, the current study tested the common-intersection hypothesis through mathematical analysis on the basis of two linear relationships (i.e., the linear relationship between temperature and development rate and that between the lower developmental threshold and the sum of effective temperatures). An attempt was made to combine the rate isomorphy hypothesis that all the lower developmental thresholds of different stages were assumed to be equal for a single species with the common-intersection hypothesis for related species within a taxon and hypothesize that if these two hypotheses hold, then for any developmental stages a common temperature of all stages should exist.     

Estimating temperature‐dependent developmental rates of potato tuberworm,Phthorimaea operculella (Lepidoptera: Gelechiidae)

Abstract The potato tuberworm, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), is the most destructive pest of potato, Solanum tuberosum L. (Solanaceae), in tropical and subtropical regions in both field and storeroom situations. The modeling of temperature‐dependent development can be useful in forecasting occurrence and population dynamics of the pests. Published developmental parameters for this pest vary greatly for many reasons. We determined temperature‐dependent development of P. operculella at seven constant temperatures (16, 20, 24, 28, 32, 34 and 36 °C). Developmental period of whole immature stage (egg to the end of the pupal stage) varied from 75.5 days at 16 °C to 17 days at 32 °C. The population failed to survive at 36 °C. The observed data was modeled to determine mathematical functions for simulating P. operculella development in each stage of development and overall. Two linear models, ordinary linear regression and the Ikemoto linear model were used to describe the relationship between temperature and development rate of the different stages of P. operculella and estimating the thermal constant and lower temperature threshold. The lower temperature threshold (t) and thermal constant (k) of whole immature stage were estimated to be 11.6 °C and 338.5 DD by Ikemoto linear model, and the estimated parameters were not substantially different with those estimated by ordinary linear models. Different models provided a better fit to the various developmental stages. Of the eleven nonlinear models fitted, the Beriere‐1, Logan‐6 and Lactin‐1 model was found to be the best for modeling development rate of egg, larva and pupa of P. operculella, respectively. Phenological models based on these findings can be part of a decision‐support tool to improve the efficiency of pest management programs.     

Modelling development of reptile embryos under fluctuating temperature regimes

An increase in temperature, within bounds, will accelerate development of reptile embryos, and morphogenesis can be normal over a range of temperatures despite those varying rates of development. Less well understood is the form of the relationship that best describes variation in developmental rate with temperature. In this article, we apply a linear degree.hour model, an empirical curvilinear model, a biophysical model, and a polynomial model to data on rates of embryonic development and temperature in the pig-nosed turtle Carettochelys insculpta from northern Australia. The curvilinear models, which have been applied with success to development of insects, describe the embryonic development of turtles well. When fluctuating temperatures extend beyond the constant temperatures that support successful incubation, the curvilinear models continue to perform well, whereas the linear model predictions fail. Sensitivity analysis indicates that under some circumstances, incubation duration may be increased by diel temperature fluctuations, independent of an influence of mean temperature. In other circumstances, incubation duration may be decreased, and in still other circumstances, diel temperature fluctuations will have no impact on incubation duration. This adds an additional dimension to our understanding of how thermal regimes can be selected or manipulated by reptiles to optimise incubation duration and the timing of offspring emergence.     

土壤-植物-大气连续体水热、CO2通量估算模型研究进展

土壤-植物-大气连续体(SPAC)水热、CO2通量的准确估算对理解陆地和大气的物质和能量交换过程有着重要意义.重点阐述了基于过程的土壤-植物-大气连续体水热、CO2通量模型,综述了统计模型、综合模型及基于遥感的模型的发展过程.其中水热通量统计模型包括基于温度和湿度以及基于温度和辐射的方法;CO2通量统计模型包括基于气候因子或蒸散因子以及基于光能利用率的方法.水热通量过程模型包括大叶、双源、多源和多层的水热传输物理模型;CO2通量过程模型包括叶片尺度及由大叶、双叶和多层方法扩展到冠层尺度的生理生态模型以及光合-蒸腾耦合模型.综合模型包括生物物理模型、生物化学模型和生物地理模型.统计模型形式简单,资料易得,对大范围的水热通量模拟具有指导意义;过程模型准确的揭示了水热和CO2通量传输的物理和生理过程,是大尺度综合模型的基础.未来生态系统水热、CO2通量估算模型将集成各种技术手段进行多尺度网络观测和大尺度机理模拟.     

Mapping members of the Anopheles gambiae complex using climate data

Abstract. Climate is the most important factor governing the distribution of insects over large areas. Warmth and moisture are essential for most insects' reproduction, development and survival. Here, it is shown that the principal vectors of malaria in Africa, members of the Anopheles gambiae complex, flourish within specific climate envelopes. By identifying these climatic conditions empirically, using climate or environmental databases, it is possible to map the distribution and relative abundance of mosquito species, and their chromosomal forms, at continental scales. Alternatively, mathematical models based on a fundamental understanding of how mosquitoes are affected by different climate factors, such as temperature and humidity, can also be employed to map distributions. Empirical or process‐driven models based on climate, or other environmental variables, provide simple tools for mapping the distribution and relative abundance of vectors at a coarse scale over large areas.     

Modelling development time of Lipaphis erysimi (Hemiptera: Aphididae) at constant and variable temperatures

The development period from birth to adult of virginoparae of the turnip aphid, Lipaphis erysimi (Kaltenbach), at 14 constant, 15 alternating and 15 natural temperature regimes were modelled to determine mathematical functions for simulating aphid development under a wide range of natural conditions. The day-degree model, the logistic equation, and the Wang model were used to describe the relationships between temperature and development rate at constant and alternating temperatures. The three models were then used with a Weibull function describing the distribution of development times, to simulate the development of individuals of cohorts at natural temperature regimes. Comparison of the observed with simulated distributions of adult emergence indicates that all three models can simulate the development of L. erysimi equally well when temperature does not go below 6 degrees C (the notional low temperature threshold of the day-degree model) or above 30 degrees C. When accumulation of temperatures below 6 degrees C becomes substantial, only the logistic curve offers accurate simulations; the other two models give falsely longer durations of development. When accumulation of temperatures above 30 degrees C becomes substantial, the logistic curve and the Wang model offer more accurate simulations than the day-degree model, which tends to produce shorter durations of development. Further analysis of the data reveals that development rate of this aphid at a given unfavourable high temperature may vary with time. Methods for accurately simulating the development time of L. erysimi in the field are suggested. The significance of modelling insect development at low and high temperatures by non-linear models is discussed.     

Modelling the effects of pre-exposure and post-exposure vaccines in tuberculosis control

Epidemic control strategies alter the spread of the disease in the host population. In this paper, we describe and discuss mathematical models that can be used to explore the potential of pre-exposure and post-exposure vaccines currently under development in the control of tuberculosis. A model with bacille Calmette-Guerin (BCG) vaccination for the susceptibles and treatment for the infectives is first presented. The epidemic thresholds known as the basic reproduction numbers and equilibria for the models are determined and stabilities are investigated. The reproduction numbers for the models are compared to assess the impact of the vaccines currently under development. The centre manifold theory is used to show the existence of backward bifurcation when the associated reproduction number is less than unity and that the unique endemic equilibrium is locally asymptotically stable when the associated reproduction number is greater than unity. From the study we conclude that the pre-exposure vaccine currently under development coupled with chemoprophylaxis for the latently infected and treatment of infectives is more effective when compared to the post-exposure vaccine currently under development for the latently infected coupled with treatment of the infectives.