内蒙古典型草原几种优势植物生态适应对策研究
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摘要
为了揭示草原的退化机理，采用野外放牧强度控制实验和盆栽刈割实验相结合，从克隆形态、生殖、资源(生物量及营养成分)分配格局等方面研究了退化草原优势植物冷蒿(Artemisia frigida)、星毛委陵菜(Potentilla acaulis)对不同放牧强度的生态适应对策；采用水分模拟实验，从资源分配角度探讨了典型草原优势植物羊草(Leymus chinensis)、大针茅(Stipa grandis)、冷蒿、星毛委陵菜对未来降水变化的响应。
(1)形态可塑性是植物种群克服环境异质性的重要途径之一，克隆植物的形态可塑性是植物利用异质性资源的生态适应对策。
冷蒿、星毛委陵菜的克隆形态特征对放牧强度表现出显著的可塑性反应。冷蒿间隔子长度零牧＞轻牧＞重牧＞中牧；分枝强度重牧＞中牧＞轻牧＞零牧。星毛委陵菜间隔子长度轻牧＞零牧＞中牧＞重牧；分枝强度重牧＞中牧＞轻牧＞零牧。
(2)繁殖对策是指生物对环境的生殖适应趋势，是资源或能量向生存、生长和生殖等活动中最适分配的结果。植物在不同生境中的繁殖对策反映出植物对环境的适应能力和在该生境中的生殖潜能。
冷蒿种群头状花序中管状花的数量及种群结实量轻牧＞零牧＞中牧＞重牧，零牧与重牧之间差异显著；花序中种子数为零牧＞轻牧＞中牧＞重牧，零牧、轻牧与重牧之间差异显著。
冷蒿有性生殖分配从零牧到重牧依次为10.61％，8.54％，6.41％，0.92％，其中零牧与中牧和重牧、轻牧与重牧间差异显著；刈割实验得到相似的结果。
星毛委陵菜生殖枝密度零牧与重牧间差异显著；种群生殖枝分化率在总体上随放牧强度的增加呈下降趋势，零牧显著高于中牧和重牧，但与轻牧间差异不显著；生殖枝上花序数目呈增加趋势，但各处理间差异不显著。
按比例刈割冷蒿再生量1/4刈割＞不刈割＞2/4刈割＞3/4刈割，且按比例刈割冷蒿的再生生长优于留茬高度刈割；刈牧显著影响冷蒿生物量分配格局，尤其是叶和花的分配；冷蒿有性生殖分配随刈割强度的增加而降低，繁殖方式
发生改变，主要通过克隆生长维持和扩大种群。
（3）植物在不同环境条件下的资源分配格局反映了植物发育对环境的响应
规律和适应对策。
随放牧强度的增加，冷蒿种群总生物量呈递减趋势；星毛委陵菜种群总生
物量则先减少后增加。随放牧强度增加，冷蒿地上部分生物量显著下降。
冷蒿种群生物量分配的总体格局是：根＞叶＞茎。星毛委陵菜种群生物量
分配格局为：根＞叶＞横生匍匐茎＞叶柄。
（4）碳、氮、磷等元素是构成植物体的重要元素，其含量及分配状况影响
植物个体的生长与发育，进而影响种群的竟争能力，这是群落演替过程中物种
替代的重要原因之一。因此，植物营养元素含量及其分配的变化是植物适应放
牧生境的重要生态对策之一，这与植物种类和放牧强度有关。
冷蒿叶C／N比随放牧强度的增大而增大，茎CM比在一定范围内随牧压
增加而增大，之后则减小。地下部分CM比在一定范围内随放牧率增大而减
小，之后放牧率再增大，CM比又增加。放牧使星毛委陵菜地上部分CM比
减小，地下部分CM比增加。
（5）具有不同生态适应对策的物种共存时，有利于充分利用环境资源，是
天然草地群落稳产、高产的基础。随着放牧强度的增加，群落中的增加种和侵
入种逐渐替代原来的减少种，使群落组成与结构发生变化。
轻度放牧时，冷蒿小禾草草原群落物种多样性及初级生产能力最高，并随
着放牧强度的增加而下降；其中，克氏针茅和羊草等种群的重要值随放牧强度
增大而显著降低，冷蒿让位于更加耐牧的星毛委陵菜和寸草苔。中牧条件下糙
隐子草重要值最高；过度放牧时，群落组成中以星毛委陵菜占绝对优势，变蒿、
刺穗萎、伏毛山毒草等比例有所增加，一年生植物狗尾草开始呈斑块状分布。
内蒙古典型草原草地畜牧业可持续发展适宜放牧强度为：暖季不超过2。67
羊ban‘，即每个羊单位应占有夏季草场不少于 0．4Allll‘，于 5月底或 6月初
开始放牧，这样，既可使羊草草原或大针茅草原不会退化到冷蒿草原，又可使
冷蒿退化草原有可能得以恢复，达到利用式改良的目的。否则，内蒙古典型草
原将最终退化趋同于星毛委陵菜群落，直至沙化，草地生产力下降，可利用率
降低。
（6）羊草、大针茅、冷蒿。星毛委陵菜种群生物量及其分配格局对降水变
化的响应差异显著。水分充足时，植物生物量相对多地分配到地上部分，有利
二
于获取更多的光能，提高竞争能力；水分溃缺时，植物根系生物量增多，有利
于吸收更多的水分和营养物质，提高竟争能力。因此，植物可以通过改变资源
分配格局而提高其竞争和适应能力。不同植物的生长对水分胁迫的敏感性不
同，这种因物种而异的特点，会引起旱生植物种群优势度的变化，最终引起生
态系统结构与功能的改变。
门）水分变化对草原植物养分含量及其分配格局的影响随物种的不同而不
同。水分条件
The research was conducted at Inner Mongolia Grassland Ecosystem Research Station of Chinese Academy of Sciences (43?26' -44?08' N, 116?04' -117?05' E) . It was studied that clonal morphological plasticity, reproductive strategy and resource allocation patterns of Artemisia frigida and Potentilla acaulis under different grazing intensities (zero grazing, light grazing: 1.33 sheep-hm'2, moderate grazing: 4.00 sheep-hm"2, heavy grazing: 6.67 sheep-hnr2) . Resource allocation patterns of four dominant species (Leymus chinensis, Stipa grandis, Artemisia frigida and Potentilla acaulis) in response to precipitation changes were investigated by simulation experiment.
1) Morphological plasticity was one of the important ecological strategies to utilizing heterogeneous resources. Spacer length and branching intensity of Artemisia frigida and Potentilla acaulis responded to grazing intensities significantly. Spacer length of Artemisia frigida was the longest under zero grazing and shortest under moderate grazing while that of Potentilla acaulis was the longest under light grazing and shortest under heavy grazing. Branching intensity of both species increased significantly along the grazing gradient. Difference in morphological plasticity of these two species indicated that they had different ecological strategies to grazing intensities.
2) Reproductive strategy was the trade-offs of resource allocation to maintenance, growth and production. It indicated adaptive ability and reproductive potentiality of plant population.
Index of reproductive shoots density, tubular flower number and seeds per area of Artemisia frigida were significantly different between grazing treatments. Sexual reproduction allocation was 10.61%, 8.54%, 6.41%, 0.92% respectively from zero grazing to heavy grazing. There were significant differences between zero grazing and moderate grazing, zero grazing and heavy grazing, light and heavy grazing. The results of clipping experiment were identical to that of grazing experiment. Sexual reproduction ability of Artemisia frigida was affected significantly
by grazing intensities. There was significant difference in reproductive shoot density between zero grazing and heavy grazing. The number of inflorenscences per reproductive shoot of Potentilla acaulis increased with the increase of grazing intensities. But there was no significant difference between treatments.
Results of clipping experiment showed that regrowth ability of Artemisia frigida under proportional clipping was superior to that under stubble clipping. Biomass allocation patterns of Artemisia frigida were significantly different between treatments, particularly allocation to leaves and flowers. Sexual reproduction allocation decreased with the increase of grazing or clipping intensities, which suggested that reproductive mode of Artemisia frigida changed under heavy grazing. Biomass allocation in priority to regenerative organism and reproductive mode alteration to clonal growth to sustain and propagate population were important ecological strategies to heavy grazing.
3) Biomass allocation pattern indicated plant s adaptive strategy. ANOVA results showed significant effects of grazing treatments on population total biomass, above-ground biomass and R/S ratio. Population total biomass in Artemisia frigida decreased while that in Potentilla acaulis decreased firstly and then increased with the increase of grazing intensities The above-ground biomass in Artemisia frigida decreased significantly with the increase of grazing intensities. The reverse trend was true for Potentilla acaulis. Trends in R/S were similar to that in above-ground biomass.
Biomass allocation pattern of Artemisia frigida was: roots > leaves >stems. ANOVA results did not show significant effects of treatment on biomass of leaves, roots and their allocation in Artemisia frigida except stem and its allocation. The difference in stem biomass and its allocation was significant between no grazing and light grazing, light and heavy grazing.
ANOVA results showe
(1)形态可塑性是植物种群克服环境异质性的重要途径之一，克隆植物的形态可塑性是植物利用异质性资源的生态适应对策。
冷蒿、星毛委陵菜的克隆形态特征对放牧强度表现出显著的可塑性反应。冷蒿间隔子长度零牧＞轻牧＞重牧＞中牧；分枝强度重牧＞中牧＞轻牧＞零牧。星毛委陵菜间隔子长度轻牧＞零牧＞中牧＞重牧；分枝强度重牧＞中牧＞轻牧＞零牧。
(2)繁殖对策是指生物对环境的生殖适应趋势，是资源或能量向生存、生长和生殖等活动中最适分配的结果。植物在不同生境中的繁殖对策反映出植物对环境的适应能力和在该生境中的生殖潜能。
冷蒿种群头状花序中管状花的数量及种群结实量轻牧＞零牧＞中牧＞重牧，零牧与重牧之间差异显著；花序中种子数为零牧＞轻牧＞中牧＞重牧，零牧、轻牧与重牧之间差异显著。
冷蒿有性生殖分配从零牧到重牧依次为10.61％，8.54％，6.41％，0.92％，其中零牧与中牧和重牧、轻牧与重牧间差异显著；刈割实验得到相似的结果。
星毛委陵菜生殖枝密度零牧与重牧间差异显著；种群生殖枝分化率在总体上随放牧强度的增加呈下降趋势，零牧显著高于中牧和重牧，但与轻牧间差异不显著；生殖枝上花序数目呈增加趋势，但各处理间差异不显著。
按比例刈割冷蒿再生量1/4刈割＞不刈割＞2/4刈割＞3/4刈割，且按比例刈割冷蒿的再生生长优于留茬高度刈割；刈牧显著影响冷蒿生物量分配格局，尤其是叶和花的分配；冷蒿有性生殖分配随刈割强度的增加而降低，繁殖方式
发生改变，主要通过克隆生长维持和扩大种群。
（3）植物在不同环境条件下的资源分配格局反映了植物发育对环境的响应
规律和适应对策。
随放牧强度的增加，冷蒿种群总生物量呈递减趋势；星毛委陵菜种群总生
物量则先减少后增加。随放牧强度增加，冷蒿地上部分生物量显著下降。
冷蒿种群生物量分配的总体格局是：根＞叶＞茎。星毛委陵菜种群生物量
分配格局为：根＞叶＞横生匍匐茎＞叶柄。
（4）碳、氮、磷等元素是构成植物体的重要元素，其含量及分配状况影响
植物个体的生长与发育，进而影响种群的竟争能力，这是群落演替过程中物种
替代的重要原因之一。因此，植物营养元素含量及其分配的变化是植物适应放
牧生境的重要生态对策之一，这与植物种类和放牧强度有关。
冷蒿叶C／N比随放牧强度的增大而增大，茎CM比在一定范围内随牧压
增加而增大，之后则减小。地下部分CM比在一定范围内随放牧率增大而减
小，之后放牧率再增大，CM比又增加。放牧使星毛委陵菜地上部分CM比
减小，地下部分CM比增加。
（5）具有不同生态适应对策的物种共存时，有利于充分利用环境资源，是
天然草地群落稳产、高产的基础。随着放牧强度的增加，群落中的增加种和侵
入种逐渐替代原来的减少种，使群落组成与结构发生变化。
轻度放牧时，冷蒿小禾草草原群落物种多样性及初级生产能力最高，并随
着放牧强度的增加而下降；其中，克氏针茅和羊草等种群的重要值随放牧强度
增大而显著降低，冷蒿让位于更加耐牧的星毛委陵菜和寸草苔。中牧条件下糙
隐子草重要值最高；过度放牧时，群落组成中以星毛委陵菜占绝对优势，变蒿、
刺穗萎、伏毛山毒草等比例有所增加，一年生植物狗尾草开始呈斑块状分布。
内蒙古典型草原草地畜牧业可持续发展适宜放牧强度为：暖季不超过2。67
羊ban‘，即每个羊单位应占有夏季草场不少于 0．4Allll‘，于 5月底或 6月初
开始放牧，这样，既可使羊草草原或大针茅草原不会退化到冷蒿草原，又可使
冷蒿退化草原有可能得以恢复，达到利用式改良的目的。否则，内蒙古典型草
原将最终退化趋同于星毛委陵菜群落，直至沙化，草地生产力下降，可利用率
降低。
（6）羊草、大针茅、冷蒿。星毛委陵菜种群生物量及其分配格局对降水变
化的响应差异显著。水分充足时，植物生物量相对多地分配到地上部分，有利
二
于获取更多的光能，提高竞争能力；水分溃缺时，植物根系生物量增多，有利
于吸收更多的水分和营养物质，提高竟争能力。因此，植物可以通过改变资源
分配格局而提高其竞争和适应能力。不同植物的生长对水分胁迫的敏感性不
同，这种因物种而异的特点，会引起旱生植物种群优势度的变化，最终引起生
态系统结构与功能的改变。
门）水分变化对草原植物养分含量及其分配格局的影响随物种的不同而不
同。水分条件
The research was conducted at Inner Mongolia Grassland Ecosystem Research Station of Chinese Academy of Sciences (43?26' -44?08' N, 116?04' -117?05' E) . It was studied that clonal morphological plasticity, reproductive strategy and resource allocation patterns of Artemisia frigida and Potentilla acaulis under different grazing intensities (zero grazing, light grazing: 1.33 sheep-hm'2, moderate grazing: 4.00 sheep-hm"2, heavy grazing: 6.67 sheep-hnr2) . Resource allocation patterns of four dominant species (Leymus chinensis, Stipa grandis, Artemisia frigida and Potentilla acaulis) in response to precipitation changes were investigated by simulation experiment.
1) Morphological plasticity was one of the important ecological strategies to utilizing heterogeneous resources. Spacer length and branching intensity of Artemisia frigida and Potentilla acaulis responded to grazing intensities significantly. Spacer length of Artemisia frigida was the longest under zero grazing and shortest under moderate grazing while that of Potentilla acaulis was the longest under light grazing and shortest under heavy grazing. Branching intensity of both species increased significantly along the grazing gradient. Difference in morphological plasticity of these two species indicated that they had different ecological strategies to grazing intensities.
2) Reproductive strategy was the trade-offs of resource allocation to maintenance, growth and production. It indicated adaptive ability and reproductive potentiality of plant population.
Index of reproductive shoots density, tubular flower number and seeds per area of Artemisia frigida were significantly different between grazing treatments. Sexual reproduction allocation was 10.61%, 8.54%, 6.41%, 0.92% respectively from zero grazing to heavy grazing. There were significant differences between zero grazing and moderate grazing, zero grazing and heavy grazing, light and heavy grazing. The results of clipping experiment were identical to that of grazing experiment. Sexual reproduction ability of Artemisia frigida was affected significantly
by grazing intensities. There was significant difference in reproductive shoot density between zero grazing and heavy grazing. The number of inflorenscences per reproductive shoot of Potentilla acaulis increased with the increase of grazing intensities. But there was no significant difference between treatments.
Results of clipping experiment showed that regrowth ability of Artemisia frigida under proportional clipping was superior to that under stubble clipping. Biomass allocation patterns of Artemisia frigida were significantly different between treatments, particularly allocation to leaves and flowers. Sexual reproduction allocation decreased with the increase of grazing or clipping intensities, which suggested that reproductive mode of Artemisia frigida changed under heavy grazing. Biomass allocation in priority to regenerative organism and reproductive mode alteration to clonal growth to sustain and propagate population were important ecological strategies to heavy grazing.
3) Biomass allocation pattern indicated plant s adaptive strategy. ANOVA results showed significant effects of grazing treatments on population total biomass, above-ground biomass and R/S ratio. Population total biomass in Artemisia frigida decreased while that in Potentilla acaulis decreased firstly and then increased with the increase of grazing intensities The above-ground biomass in Artemisia frigida decreased significantly with the increase of grazing intensities. The reverse trend was true for Potentilla acaulis. Trends in R/S were similar to that in above-ground biomass.
Biomass allocation pattern of Artemisia frigida was: roots > leaves >stems. ANOVA results did not show significant effects of treatment on biomass of leaves, roots and their allocation in Artemisia frigida except stem and its allocation. The difference in stem biomass and its allocation was significant between no grazing and light grazing, light and heavy grazing.
ANOVA results showe
引文
1 安宝林．1986．草原退化及其治理的战略措施．中国草地．(3)：56-59．
2 白永飞，陈佐忠．2000．锡林河流域羊草草原植物种群和功能群的长期变异性及其对群落稳定性的影响．植物生态学报．24(6)：641-647．
3 白永飞，李德新，许志信，魏志军．1999．牧压梯度对克氏针茅生长和繁殖的影响．生态学报．19(4)：479-484．
4 白永飞，李凌浩，王其兵，张丽霞，张焱．陈佐忠．2000．锡林河流域草原群落植物多样性和初级生产力沿水热梯度变化的样带研究．植物生态学报．24(6)：667-673．
5 宝音陶格涛，李艳梅，贾建芬，杨持．2000．牧压梯度下冷蒿有性繁殖器官变化特征的观察分析．内蒙古大学学报(自然科学版)．31(3)：311-313．
6 陈玲，阿里木斯，杨持．1997．草原植物种群营养元素生殖分配规律研究(Ⅰ)．内蒙古大学学报(自然科学版)．28(4)：568～571．
7 陈敏，王艳华，1985．栽培条件下羊草生物学特性的观察．草原生态系研究(第1集)．北京：科学出版社，212-223．
8 陈尚，李自珍，王刚．1997．克隆植物生长型的研究进展．生态学杂志，16(4)：59-63．
9 陈尚，马艳，李白珍，王刚．1999．克隆植物种子繁殖和营养繁殖的适合度分析和度量．生态学报，19(2)：287-290．
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