放牧对克氏针茅功能性状的影响
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摘要
植物功能性状是植物对周围环境的适应及响应,进一步了解植物的功能性状,对于深入了解由放牧导致的草地退化具有重要意义。于内蒙古大学生态学研究基地开展4年(2012-2015年)放牧控制试验,通过研究克氏针茅(Stipa krylovii)在放牧干扰下的个体功能性状,探讨内蒙古典型草地生态系统中克氏针茅对放牧的响应和适应对策。2016年,在克氏针茅种群生物量高峰期(8月)测定了连续4年放牧(2012-2015年)和无牧条件下(对照)的植株高度(PH)、总叶面积(TLA)、茎叶比(SLW)、茎质量(SW)、总质量(AB)、总叶质量(TLQ)、比叶面积(SLA)、叶数(LN)、营养枝数(NNB)、丛幅(WCC)共10个性状指标。结果表明:(1)4年放牧显著降低了植株高度和比叶面积(P<0.05),与对照相比,植株高度和比叶面积分别下降了31%和58%;显著增加了克氏针茅的营养枝数和叶数(P<0.05);与对照相比,营养枝数与叶数分别增加了50%和63%;(2)其中比叶面积、营养枝数、叶数这3个功能性状为敏感性指标,其余7个为惰性指标;(3)生物量与株高、茎质量、营养枝数、总叶质量、茎叶比、丛幅、叶面积、和叶数存在极显著的正相关关系(P<0.01),与比叶面积呈现显著负相关关系;(4)株高、茎质量、总叶质量、总质量、丛幅、叶面积之间存在显著正相关关系。该研究认为,克氏针茅各功能性状对放牧干扰的不同响应,是其对干扰产生的生态对策,进而维持种群的稳定性。
Plant functional traits represent how plants adapt and response to the surrounding environment. Studying plant functional traits is of great significance for understanding the mechanism of grassland degradation caused by grazing. To explore the response and adaptation strategies of Stipa krylovii to grazing in the typical steppe ecosystems in Inner Mongolia, a total of 10 trait indicators including plant height(PH), total leaf area(TLA), stem/leaf ratio(SLW), stem weight(SW), Aboveground biomass(AB), Total leaf quality(TLQ), specific leaf area(SLA), number of leaves(NL), number of nutrient branches(NNB), width of clusters(WCC) were respectively measured in biomass peak of Stipa krylovii population(August) under consecutive grazing(year 2012-2015) and no grazing(control) conditions. Our results showed that:(1) Compared to the control, 4-year consecutives grazing significantly decreased the PH and SLA by 31% and 58%(P<0.05), but evidently increased the NNB and NL by 50% and 63%(P<0.05)respectively;(2) NNB and NL were sensitive indicators, and the other seven traits were inertindicators;(3) There was a significantly positive correlation between biomass and PH, SW, NNB, TLQ, SLW, WCC, TLA and LN(P<0.01), while the biomass was negatively correlated with SLA; And(4) there were also significantly positive correlation among PH, SW, TLQ, AB, WCC and TLA.Our study suggests that the different responses of various functional traits of Stipa krylovii to grazing disturbance are ecological countermeasures for disturbance to maintain the stability of the population.
Plant functional traits represent how plants adapt and response to the surrounding environment. Studying plant functional traits is of great significance for understanding the mechanism of grassland degradation caused by grazing. To explore the response and adaptation strategies of Stipa krylovii to grazing in the typical steppe ecosystems in Inner Mongolia, a total of 10 trait indicators including plant height(PH), total leaf area(TLA), stem/leaf ratio(SLW), stem weight(SW), Aboveground biomass(AB), Total leaf quality(TLQ), specific leaf area(SLA), number of leaves(NL), number of nutrient branches(NNB), width of clusters(WCC) were respectively measured in biomass peak of Stipa krylovii population(August) under consecutive grazing(year 2012-2015) and no grazing(control) conditions. Our results showed that:(1) Compared to the control, 4-year consecutives grazing significantly decreased the PH and SLA by 31% and 58%(P<0.05), but evidently increased the NNB and NL by 50% and 63%(P<0.05)respectively;(2) NNB and NL were sensitive indicators, and the other seven traits were inertindicators;(3) There was a significantly positive correlation between biomass and PH, SW, NNB, TLQ, SLW, WCC, TLA and LN(P<0.01), while the biomass was negatively correlated with SLA; And(4) there were also significantly positive correlation among PH, SW, TLQ, AB, WCC and TLA.Our study suggests that the different responses of various functional traits of Stipa krylovii to grazing disturbance are ecological countermeasures for disturbance to maintain the stability of the population.
引文
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李西良,刘志英,侯向阳,等,2015.放牧对草地植物功能性状及其权衡关系的调控[J].植物学报,50(2):159-170.LI X L,LIU Z Y,HOU X Y,et al.,2015.Plant Functional Traits and Their Trade-offs in Response to Grazing:A Review[J].Chinese Bulletin of Botany,50(2):159-170.
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秦洁,韩国栋,乔江,等,2016.内蒙古不同草地类型中羊草地上生物量对放牧强度的响应[J].中国草地学报,38(4):76-82.QIN J,HAN G D,QIAO J,et al.,2016.Response of Leymus chinensis above-ground Biomass to Grazing Intensity in Different Grasslands of Inner Mongolia[J].Chinese Journal of Grassland,38(4):76-82.
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卫智军,刘文亭,吕世杰,等,2016.荒漠草地短花针茅种群年龄对放牧调控的响应机制[J].生态环境学报,25(12):1922-1928.WEI Z J,LIU W T,LV S J,et al.,2016.Response Mechanism of Stipa Breviflora Age to Graze in Desert Grassland[J].Ecology and Environmental Sciences,25(12):1922-1928.
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徐惠敏,白天晓,安娜,等,2016.不同刈割制度对典型草地羊草功能性状的影响[J].中国草地学报,37(6):60-65.XU H M,BAI T X,AN N,et al.,2016.Effect of Different Cutting System on Leymus chinensis Functional Traits[J].Chinese Journal of Grassland,37(6):60-65.
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BAI Y F,WU J G,CLARK C M,et al.,2012.Grazing alters ecosystem functioning and C?N?P stoichiometry of grasslands along a regional precipitation gradient[J].Journal of Applied Ecology,49(6):1204-1215.
BAI Y F,WU J G,PAN Q M,et al.,2007.Positive linear relationship between productivity and diversity:evidence from the Eurasian Steppe[J].Journal of Applied Ecology,44(5):1023-1034.
BELSKY A J,CARSON W P,JENSON C L,et al.,1993.Overcompensation by plants:Herbivory optimization or red herring[J].Evolutionary Ecology,7(1):109-121.
BERNARD-VERDIER M,NAVAS M L,VELLEND M,et al.,2012.Community assembly along a soil depth gradient:contrasting patterns of plant trait convergence and divergence in a Mediterranean rangeland[J].Journal of Ecology,100(6):1422-1433.
BULLOCK J M,FRANKLIN J,STEVENSON M J,et al.,2001.A plant trait analysis of responses to grazing in a long-term experiment[J].Journal of Applied Ecology,38(2):253-267.
CHEN S P,BAI Y F,LIN G H,et al.,2005.Effects of grazing on photosynthetic characteristics of major steppe species in the Xilin River Basin,Inner Mongolia,China[J].Photosynthetica,43(4):559-565.
CORNER E J H,1949.The durian theory or the origin of the modern tree[J].Annals of Botany,13(4):367-414.
DAVID J,ELDRIDGE,ALISTAIR G B,et al.,2016.Ecosystem structure,function,and composition in rangelands are nega-tively affected by livestock grazing[J].Ecological Applica-ions,26(4):1273-1283.
DIAZ S,LAVOREL S,DE BELLO F,et al.,2007.Incorporating plant functional diversity effects in ecosystem service assess-ments[J].Proceedings of the National Academy of Sciences of the United State of America,104(52):20684-20689.
ELDRIDGE D J,POORE A G,LETNIC M,et al.,2016.Ecosystem structure,function,and composition in rangelands are nega-tively affected by livestock grazing[J].Ecological Applications,26(4):1273-1283.
KLIMKOWSKA A,BEKKER R M,VAN DIGGELEN R,et al.,2010.Species trait shifts in vegetation and soil seed bank during fen degra-dation[J].Plant Ecology,206(1):59-82.
KNAPP A K,SMITH M D,2001.Variation among biomes in temporal dynamics of aboveground primary production[J].Science,291(5503):481-484.
LAVOREL S,GRIGULIS K,LAMARQUE P,et al.,2011.Using plant functional traits understand the landscape distribution of multiple ecosystem services[J].Journal of Ecology,299(1):135-147.
LEMAIRE G,HODGSON J,NABRINGER C,et al.,2000.Grassland Ecophysiology and Grazing Ecology[J].International symposium on grassland ecophysiology and grazing ecology,42(3):981-982.
LI B,YONG S P,LI Z H,1988.The vegetation of the Xilin river basin and its utilization[J].Research on Grassland Ecosystem(3):84-183.
MOONEY K A,HALITSCHKE R,KESSLER A,et al.,2010.Evolutionary trade-offs in plants mediate the strength of trophic cascades[J].Science,327(5973):1642-1644.
PENG S S,PIAO S L,CIAIS P,et al.,2013.Asymmetric effects of daytime and night-time warming on Northern Hemisphere vegetation[J].Nature,501(7465):88-92.
SHAN Y M,CHEN D M,GUAN X X,et al.,2011.Seasonally dependent impacts of grazing on soil nitrogen mineralization and linkages to ecosystem functioning in Inner Mongolia grassland[J].Soil Biology and Biochemistry,43(9):1943-1954.
冯祥超,张璐,李雅茹,等,2017.季节性放牧对典型草地土壤微生物碳氮周转的影响[J].中国草地学报,39(2):73-84.FEN X C,ZHANG L,LI Y R,et al.,2017.Effect of Ftmr Forage on Production Performance of Lambs[J].Chinese Journal of Grassland,39(2):73-84.
李江文,王忠武,任海燕,等,2017.荒漠草地建群种短花克氏针茅功能性状对长期放牧的可塑性响应[J].西北植物学报,37(9):1854-1863.LI J W,WANG Z W,REN H Y,et al.,2017.Plastic Response of Individual Functional Traits in Stipa breviflora to Long term Grazing in a Desert Steppe[J].Acta Botanica Boreali-Occidentalia Sinica,37(9):1854-1863.
李西良,侯向阳,吴新宏,等,2014.草甸草地羊草茎叶功能性状对长期过度放牧的可塑性响应[J].植物生态学报,38(5):440-451.LI X L,HOU X Y,WU X H,et al.,2014.Plastic responses of stem and leaf functional traits in Leymus chinensis to long-term grazing in a meadow steppe[J].Chinese Journal of Plant Ecology,38(5):440-451.
李西良,刘志英,侯向阳,等,2015.放牧对草地植物功能性状及其权衡关系的调控[J].植物学报,50(2):159-170.LI X L,LIU Z Y,HOU X Y,et al.,2015.Plant Functional Traits and Their Trade-offs in Response to Grazing:A Review[J].Chinese Bulletin of Botany,50(2):159-170.
刘文亭,卫智军,吕世杰,等,2016.内蒙古荒漠草原短花针茅叶片功能性状对不同草地经营方式的响应[J].生态环境学报,25(3):385-392.LIU W T,WEI Z J,LV S J,et al.,2016.Response of Grassland Using Modes to Leaf Trait of Stipa breviflora in Desert Steppe of Inner Mongolia[J].Ecology and Environmental Sciences,25(3):385-392.
刘晓娟,马克平,2015.植物功能性状研究进展[J].中国科学:生命科学,45(4):325-339.LIU X J,MA K P,2015.Plant Functional Traits:Concepts,Applications and Future Directions[J].Science China Press,45(4):325-339.
刘钟龄,王炜,郝敦元,等,2002.内蒙古草地退化与恢复演替机理的探讨[J].干旱区资源与环境,16(1):84-91.LIU Z L,WANG W,HAO D Y,et al.,2002.Probes on the Degeneration and Recovery Succession Mechanisms of Inner Mongolia Steppe[J].Journal of Arid Land Resources and Environment,16(1):84-91.
戚德辉,温仲明,杨士梭,等,2015.基于功能性状的铁杆蒿对环境变化的响应与适应[J].应用生态学报,26(7):12-18.QI D H,WEN Z M,YANG S S,et al.,2015.Trait-based responses and adaptation of Artemisia sacrorum to environmental changes[J].Chinese Journal of Applied Ecology,26(7):12-18.
秦洁,韩国栋,乔江,等,2016.内蒙古不同草地类型中羊草地上生物量对放牧强度的响应[J].中国草地学报,38(4):76-82.QIN J,HAN G D,QIAO J,et al.,2016.Response of Leymus chinensis above-ground Biomass to Grazing Intensity in Different Grasslands of Inner Mongolia[J].Chinese Journal of Grassland,38(4):76-82.
任继周,2012.放牧,草地生态系统存在的基本方式:兼论放牧的转型[J].自然资源学报,27(8):1259-1275.REN J Z,2012.Grazing,the Basic Form of Grassland Ecosystem and Its Transformation[J].Journal of Natural Resources,27(8):1259-1275.
王炜,梁存柱,刘钟龄,等,2000.草原群落退化与恢复演替中的植物个体行为分析[J].植物生态学报,24(3):268-274.WANG W,LIANG C Z,LIU Z L,et al.,2000.Analysis of the Plant Individual Behaviour During the Degradatton and Restoring Succession in Steppe Community[J].Chinese Journal of Plant Ecology,24(3):268-274.
卫智军,刘文亭,吕世杰,等,2016.荒漠草地短花针茅种群年龄对放牧调控的响应机制[J].生态环境学报,25(12):1922-1928.WEI Z J,LIU W T,LV S J,et al.,2016.Response Mechanism of Stipa Breviflora Age to Graze in Desert Grassland[J].Ecology and Environmental Sciences,25(12):1922-1928.
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