基于改进的Lotka-Volterra种间竞争模型预测退化高寒草地人工恢复演替结果
|
摘要
结合青藏高原地区黑土滩型退化高寒草甸改建成人工草地后恢复过程中植物群落组分的数量特征变化,基于中心差分法、相对误差热图、ODE算法等研究方法进行计算机模拟,对Lotka-Volterra种间竞争模型进行非线性化改进,建立了适于高寒草地人工恢复演替过程中的竞争效应预测模型,与经典的Lotka-Volterra种间竞争模型进行对比验证,实证了改进模型的可解性和准确性。并分析了改进的Lotka-Volterra模型的系统动力学行为,预测人工恢复植物群落各组分的竞争结局,判断人工草地恢复演替状况。结果表明:(1)随着恢复年限的增加,栽培植物(垂穗披碱草或草地早熟禾)与原生植物在经历一个激烈的竞争阶段后逐渐趋于动态平衡,表明人工种植的垂穗披碱草、草地早熟禾等本土植物,可以促进青藏高原地区"黑土滩"型退化高寒草甸的有效恢复;(2)综合各分组的恢复演替阶段特征,可食牧草、栽培植物和顶极植物均从第7年开始竞争力呈现明显下降的趋势,因此建议在对人工草地的恢复管理中,7—10年间进行适度的人为干预,如施肥、灭鼠害以及适当去除有毒有害杂草等;(3)综合各类植物的竞争演替预测情况,均从第20年左右开始逐渐趋于动态平衡。因此,根据该模型分析,从生态恢复的角度来看,"黑土滩"型退化草地进行人工恢复至少需要20年以上,才能获得较为稳定的植物群落。
Combining the quantitative characteristic changes of plant community components in the restoration process of the "black soil patch" degraded alpine meadow in the Qinghai-Tibet Plateau after being converted into artificial grassland, computer simulations based on methods, such as the central difference method, relative error thermography, and ODE algorithm were conducted. A prediction model of competitive effects suitable for alpine meadow restoration succession was established through the nonlinear improvement of the Lotka-Volterra interspecific competition model. The model was compared with the classical Lotka-Volterra interspecific competition model, and the solvability and accuracy of the improved model were demonstrated. Furthermore, the dynamic behavior of the improved Lotka-Volterra model system was analyzed, the competitive outcome of the plant community components in the artificial restoration was predicted, and the restoration succession of the artificial grassland was judged. The results showed that:(1) With the increasing length of vegetation recovery, cultivated plants(Elymus nutans or Poa pratensis) and original plants will gradually tend to be dynamically balanced after undergoing a fierce competition phase. Accordingly, cultivated plants —— Elymus nutans or Poa pratensis can promote the effective restoration of the "black soil patch" degraded alpine meadow in the Qinghai-Tibet Plateau;(2) Synthesizing the characteristics of the restoration succession stage of each group, the competitiveness of edible forage, cultivated plants, and climax plants all showed an obvious downward trend since the 7 th year. In the restoration management of the artificial grassland, we recommend that moderate human interventions, such as fertilization, rodent control, poisonous and harmful weed removal be carried out in the next 7—10 years;(3) Analyzing the prediction results on competition succession of each group synthetically, all of tend to be dynamically balanced from the 20 th year. Consequently, according to the model, from the perspective of ecological restoration, it will take at least 20 years for the artificial restoration of the "black soil patch" degraded grassland to obtain a relatively stable plant community.
Combining the quantitative characteristic changes of plant community components in the restoration process of the "black soil patch" degraded alpine meadow in the Qinghai-Tibet Plateau after being converted into artificial grassland, computer simulations based on methods, such as the central difference method, relative error thermography, and ODE algorithm were conducted. A prediction model of competitive effects suitable for alpine meadow restoration succession was established through the nonlinear improvement of the Lotka-Volterra interspecific competition model. The model was compared with the classical Lotka-Volterra interspecific competition model, and the solvability and accuracy of the improved model were demonstrated. Furthermore, the dynamic behavior of the improved Lotka-Volterra model system was analyzed, the competitive outcome of the plant community components in the artificial restoration was predicted, and the restoration succession of the artificial grassland was judged. The results showed that:(1) With the increasing length of vegetation recovery, cultivated plants(Elymus nutans or Poa pratensis) and original plants will gradually tend to be dynamically balanced after undergoing a fierce competition phase. Accordingly, cultivated plants —— Elymus nutans or Poa pratensis can promote the effective restoration of the "black soil patch" degraded alpine meadow in the Qinghai-Tibet Plateau;(2) Synthesizing the characteristics of the restoration succession stage of each group, the competitiveness of edible forage, cultivated plants, and climax plants all showed an obvious downward trend since the 7 th year. In the restoration management of the artificial grassland, we recommend that moderate human interventions, such as fertilization, rodent control, poisonous and harmful weed removal be carried out in the next 7—10 years;(3) Analyzing the prediction results on competition succession of each group synthetically, all of tend to be dynamically balanced from the 20 th year. Consequently, according to the model, from the perspective of ecological restoration, it will take at least 20 years for the artificial restoration of the "black soil patch" degraded grassland to obtain a relatively stable plant community.
引文
[1] 丁明军,张镱锂,刘林山,王兆锋.1982—2009年青藏高原草地覆盖度时空变化特征.自然资源学报,2010,25(12):2114- 2122.
[2] 林慧龙,董世魁.高寒地区多年生禾草混播草地种间竞争效应分析.草业学报,2003,12(3):79- 82.
[3] 曹广民,杜岩功,梁东营,王启兰,王长庭.高寒嵩草草甸的被动与主动退化分异特征及其发生机理.山地学报,2007,25(6):641- 648.
[4] 郭军乐,郑建宗,李春宁.不同退化状态高寒草甸人工恢复与自然恢复比较.陕西林业科技,2008,(4):1- 5,68- 68.
[5] 刘敏,龚吉蕊,王忆慧,张梓榆,徐沙,罗亲普.豆禾混播建植人工草地对牧草产量和草质的影响.干旱区研究,2016,33(1):179- 185.
[6] Mandimba G R.Contribution of nodulated legumes on the growth of Zea mays L.under various cropping systems.Symbiosis,1995,19(2/3):213- 222.
[7] Wilson F E A,Harvey B M R,McAdam J H,Walton D W H.The response of Whitegrass [Cortaderia pilosa (D′Urv.) Hack.] to nitrogen nutrition.Grass and Forage Science,2001,56(1):84- 91.
[8] 李里,刘伟.退化草地植物功能群和物种丰富度与群落生产力关系的研究.草地学报,2011,19(6):917- 921,999- 999.
[9] 樊江文,钟华平,杜占池,韩彬,梁飚.草地植物竞争的研究.草业学报,2004,13(3):1- 8.
[10] Hallam T G,Clark C E,Jordan G S.Effects of toxicants on populations:a qualitative approach II.First order kinetics.Journal of Mathematical Biology,1983,18(1):25- 37.
[11] Smith F E.Population dynamics in daphnia magna and a new model for population growth.Ecology,1963,44(4):651- 663.
[12] 王慧忠,张新全.Lotka—Volterra数学模型在草地管理中的应用.草业学报,2006,15(1):54- 57.
[13] 孙飞达,龙瑞军,蒋文兰,郭正刚,聂学敏.三江源区不同鼠洞密度下高寒草甸植物群落生物量和土壤容重特性研究.草业学报,2008,17(5):111- 116.
[14] 尚占环,董全民,施建军,周华坤,董世魁,邵新庆,李世雄,王彦龙,马玉寿,丁路明,曹广民,龙瑞军.青藏高原“黑土滩”退化草地及其生态恢复近10年研究进展——兼论三江源生态恢复问题.草地学报,2018,26(1):1- 21.
[15] 马玉寿.三江源区“黑土型”退化草地形成机理与恢复模式研究[D].兰州:甘肃农业大学,2006.
[16] 张蕊,王媛,马丽娜,桑潮,王力,郭瑞英,汪海波,尚占环.三江源区退化人工草地、“黑土滩”和天然草地植物群落物种多样性.草地学报,2014,22(6):1171- 1178.
[17] 江小雷,张卫国,杨振宇,杜国祯.不同演替阶段鼢鼠土丘群落植物多样性变化研究[J].应用生态学报,2004,15(5):814- 818.
[18] Sun G W,Cui Q W,Bo S.A new mathematical model of interspecific competition —— an expansion of the classical Lotka-Volterra competition equations.Ecological Modelling,1990,58(1/4):273- 284.
[19] Law R,Murrell D J,Dieckmann U.Population growth in space and time:spatial logistic equations.Ecology,2003,84(1):252- 262.
[20] 张大勇,赵松岭.森林自疏过程中密度变化规律的研究.林业科学,1985,21(4):369- 374.
[21] 李新运,赵善伦,尤作亮,余锦.一种自适应的种群增长模型及参数估计.生态学报,1997,17(3):311- 316.
[22] 马玉寿,尚占环,施建军,董全民,王彦龙,杨时海.黄河源区"黑土滩"退化草地群落类型多样性及其群落结构研究.草业科学,2006,23(12):6- 11.
[23] 王长庭,龙瑞军,王启兰,刘伟,景增春,张莉.三江源区不同建植年代人工草地群落演替与土壤养分变化.应用与环境生物学报,2009,15(6):737- 744.
[24] Afraimovich V,Tristan I,Huerta R,Rabinovich M I.Winnerless competition principle and prediction of the transient dynamics in a Lotka-Volterra model.Chaos:An Interdisciplinary Journal of Nonlinear Science,2008,18(4):043103.
[25] 冯瑞章.人为扰动对高寒草地土壤有机碳及微生物学特征的影响[D].西宁:中国科学院西北高原生物研究所,2009.
[26] 杨慧茹.青海草地早熟禾人工草地演替规律及群落稳定性研究[D].西宁:青海大学,2012.
[27] 曹广民,龙瑞军.三江源区"黑土滩"型退化草地自然恢复的瓶颈及解决途径.草地学报,2009,17(1):4- 9.
[28] 周华坤,赵新全,赵亮,韩发,古松.高山草甸垂穗披碱草人工草地群落特征及稳定性研究.中国草地学报,2007,29(2):13- 25.
[29] 王启兰,王长庭,刘伟,曹广民,龙瑞军.三江源区不同建植年限人工草地植物群落与土壤微生物生理类群的变化.应用生态学报,2009,20(11):2646- 2651.
[30] 马玉寿,施建军,董全民,王彦龙,盛丽.人工调控措施对“黑土型”退化草地垂穗披碱草人工植被的影响.青海畜牧兽医杂志,2006,36(2):1- 3.
[31] 王元素,蒋文兰,洪绂曾,王堃.人工混播草地群落稳定性研究进展.中国草地,2005,27(4):58- 63,73- 73.
[32] 张成霞,南志标.放牧对草地土壤理化特性影响的研究进展.草业学报,2010,19(4):204- 211.
[33] 李春刚.放牧干扰对高寒草甸群落特征的影响.贵州农业科学,2010,38(5):128- 132.
[34] 董世魁,丁路明,徐敏云,龙瑞军,胡自治.放牧强度对高寒地区多年生混播禾草叶片特征及草地初级生产力的影响.中国农业科学,2004,37(1):136- 142.
[2] 林慧龙,董世魁.高寒地区多年生禾草混播草地种间竞争效应分析.草业学报,2003,12(3):79- 82.
[3] 曹广民,杜岩功,梁东营,王启兰,王长庭.高寒嵩草草甸的被动与主动退化分异特征及其发生机理.山地学报,2007,25(6):641- 648.
[4] 郭军乐,郑建宗,李春宁.不同退化状态高寒草甸人工恢复与自然恢复比较.陕西林业科技,2008,(4):1- 5,68- 68.
[5] 刘敏,龚吉蕊,王忆慧,张梓榆,徐沙,罗亲普.豆禾混播建植人工草地对牧草产量和草质的影响.干旱区研究,2016,33(1):179- 185.
[6] Mandimba G R.Contribution of nodulated legumes on the growth of Zea mays L.under various cropping systems.Symbiosis,1995,19(2/3):213- 222.
[7] Wilson F E A,Harvey B M R,McAdam J H,Walton D W H.The response of Whitegrass [Cortaderia pilosa (D′Urv.) Hack.] to nitrogen nutrition.Grass and Forage Science,2001,56(1):84- 91.
[8] 李里,刘伟.退化草地植物功能群和物种丰富度与群落生产力关系的研究.草地学报,2011,19(6):917- 921,999- 999.
[9] 樊江文,钟华平,杜占池,韩彬,梁飚.草地植物竞争的研究.草业学报,2004,13(3):1- 8.
[10] Hallam T G,Clark C E,Jordan G S.Effects of toxicants on populations:a qualitative approach II.First order kinetics.Journal of Mathematical Biology,1983,18(1):25- 37.
[11] Smith F E.Population dynamics in daphnia magna and a new model for population growth.Ecology,1963,44(4):651- 663.
[12] 王慧忠,张新全.Lotka—Volterra数学模型在草地管理中的应用.草业学报,2006,15(1):54- 57.
[13] 孙飞达,龙瑞军,蒋文兰,郭正刚,聂学敏.三江源区不同鼠洞密度下高寒草甸植物群落生物量和土壤容重特性研究.草业学报,2008,17(5):111- 116.
[14] 尚占环,董全民,施建军,周华坤,董世魁,邵新庆,李世雄,王彦龙,马玉寿,丁路明,曹广民,龙瑞军.青藏高原“黑土滩”退化草地及其生态恢复近10年研究进展——兼论三江源生态恢复问题.草地学报,2018,26(1):1- 21.
[15] 马玉寿.三江源区“黑土型”退化草地形成机理与恢复模式研究[D].兰州:甘肃农业大学,2006.
[16] 张蕊,王媛,马丽娜,桑潮,王力,郭瑞英,汪海波,尚占环.三江源区退化人工草地、“黑土滩”和天然草地植物群落物种多样性.草地学报,2014,22(6):1171- 1178.
[17] 江小雷,张卫国,杨振宇,杜国祯.不同演替阶段鼢鼠土丘群落植物多样性变化研究[J].应用生态学报,2004,15(5):814- 818.
[18] Sun G W,Cui Q W,Bo S.A new mathematical model of interspecific competition —— an expansion of the classical Lotka-Volterra competition equations.Ecological Modelling,1990,58(1/4):273- 284.
[19] Law R,Murrell D J,Dieckmann U.Population growth in space and time:spatial logistic equations.Ecology,2003,84(1):252- 262.
[20] 张大勇,赵松岭.森林自疏过程中密度变化规律的研究.林业科学,1985,21(4):369- 374.
[21] 李新运,赵善伦,尤作亮,余锦.一种自适应的种群增长模型及参数估计.生态学报,1997,17(3):311- 316.
[22] 马玉寿,尚占环,施建军,董全民,王彦龙,杨时海.黄河源区"黑土滩"退化草地群落类型多样性及其群落结构研究.草业科学,2006,23(12):6- 11.
[23] 王长庭,龙瑞军,王启兰,刘伟,景增春,张莉.三江源区不同建植年代人工草地群落演替与土壤养分变化.应用与环境生物学报,2009,15(6):737- 744.
[24] Afraimovich V,Tristan I,Huerta R,Rabinovich M I.Winnerless competition principle and prediction of the transient dynamics in a Lotka-Volterra model.Chaos:An Interdisciplinary Journal of Nonlinear Science,2008,18(4):043103.
[25] 冯瑞章.人为扰动对高寒草地土壤有机碳及微生物学特征的影响[D].西宁:中国科学院西北高原生物研究所,2009.
[26] 杨慧茹.青海草地早熟禾人工草地演替规律及群落稳定性研究[D].西宁:青海大学,2012.
[27] 曹广民,龙瑞军.三江源区"黑土滩"型退化草地自然恢复的瓶颈及解决途径.草地学报,2009,17(1):4- 9.
[28] 周华坤,赵新全,赵亮,韩发,古松.高山草甸垂穗披碱草人工草地群落特征及稳定性研究.中国草地学报,2007,29(2):13- 25.
[29] 王启兰,王长庭,刘伟,曹广民,龙瑞军.三江源区不同建植年限人工草地植物群落与土壤微生物生理类群的变化.应用生态学报,2009,20(11):2646- 2651.
[30] 马玉寿,施建军,董全民,王彦龙,盛丽.人工调控措施对“黑土型”退化草地垂穗披碱草人工植被的影响.青海畜牧兽医杂志,2006,36(2):1- 3.
[31] 王元素,蒋文兰,洪绂曾,王堃.人工混播草地群落稳定性研究进展.中国草地,2005,27(4):58- 63,73- 73.
[32] 张成霞,南志标.放牧对草地土壤理化特性影响的研究进展.草业学报,2010,19(4):204- 211.
[33] 李春刚.放牧干扰对高寒草甸群落特征的影响.贵州农业科学,2010,38(5):128- 132.
[34] 董世魁,丁路明,徐敏云,龙瑞军,胡自治.放牧强度对高寒地区多年生混播禾草叶片特征及草地初级生产力的影响.中国农业科学,2004,37(1):136- 142.
© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号 地址：北京市海淀区学院路29号 邮编：100083 电话：办公室：(+86 10)66554848；文献借阅、咨询服务、科技查新：66554700 |