辽东半岛结缕草种群动态及其生活史格局研究
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摘要
结缕草(Zoysia japonica Steud.)是一种典型的无性系草本植株，既具有地下根状茎又具有地上匍匐茎。我国大面积的野生结缕草分布在山东半岛和辽东半岛，是当地宝贵的乡土资源植物。本文以分布在辽东半岛的结缕草为研究对象，通过对结缕草实验无性系建立、观测和天然种群两个生长季的取样研究，分析了结缕草实验无性系的扩展动态规律及内在扩展机理，分析了不同生境条件下结缕草天然种群的动态及机理，按着结缕草生活史模型，阐述了结缕草种群的生活史格局。所得主要结论如下：
1.在栽培条件下，经过一个生长季的生长，结缕草实验无性系大小的可塑性较大，但均遵循着规律性的时间生长和空间扩展。随着生长时间的进程，无性系分株构件种群以指数函数规律增加，匍匐茎主茎长度以直线函数规律伸长，主茎间隔子数量则以对数函数规律增加。在空间扩展上，无性系半径随着时间以直线函数规律延伸，面积随着时间以指数函数规律扩展，分株种群的数量则随着半径的延伸以指数函数规律增加，随着面积的扩展以直线函数规律增加，主茎间隔子数量随着半径的延伸以直线函数规律增加，随着面积的扩展以对数函数规律增加。
2.对于天然结缕草种群，在两个生长季内未火烧样地的分株构件种群波动较小，火烧样地的在6、7月份出现密度峰值，波动较大，林下样地的分株密度在2005年的5月和10月出现峰值，在2006年波动很小。3个样地结缕草芽构件种群的变化趋势相似，在10月份达到密度峰值。火烧样地和未火烧样地的苗构件种群在生长季内的变化趋势相似，林下样地的则与其分株种群的变化趋势相似。结缕草根状茎的长度在3个样地中均是在生长季的两端高，在生长季的中间期较低。
3.未火烧样地的枯落物生物量随着时间逐渐减少，而火烧样地枯落物生物量在整个生长季内虽有上下波动但幅度不大。在生长季内各月份火烧样地的其他植物种株数、生物量均小于未火烧样地的。火烧清除了样地的枯落物，改善了结缕草生长的生态条件，同时火烧能够导致其他植物种的植物体及种子死亡，而对结缕草自身的影响较小，因为结缕草主要靠营养繁殖，其主要繁殖体根状茎储藏在地下，分株直立茎的分生生长点基本贴于地表或潜在地下，而且受到叶鞘的包覆保护。这样，火烧样地结缕草种群各构件的数量整体上均高于未火烧样地的。林下样地由于受到光照和土壤条件的限制，各构件的数量整体上低于未火烧样地的。
4.在未火烧样地、火烧样地和林下样地各月份的结缕草种群阶段结构中，整体上营养株所占的百分比均最高，平均64.5%以上，芽所占的百分比次之，平均为18.5%，除了生殖株以外苗所占的百分比最小，平均为14.2%。在结缕草种群生物量结构中，整体上茎生物量所占的百分比均最高，平均为51.7%，叶鞘生物量所占的百分比次之，平均为24.9%，叶片生物量的百分比最小，平均为23.3%。
5.火烧样地在5-6月间结缕草种群的增长率最大（λ=1.1247），而在9-10月间种群增长率（λ=0.557）明显小于1。未火烧样地在8-9月间种群增长率最大（λ=1.1374），而在9-10月间种群增长率（λ=0.674）明显小于1。林下样地在8-9和9-10月间种群增长率分别为λ=1.1141和λ=1.1904，在整个生长季未出现负增长（λ <1）。3个样地结缕草种群生活史的主路径均是“营养株路径”，生长季的各月份间营养株的保持对种群增长率的贡献最大，维系着种群的相对稳定。种群增长率的增加主要来自于转移等级的繁殖和生长对种群增长率的贡献。
6.营养繁殖在无性系植物种群生活史格局中处于核心地位。结缕草种群营养繁殖格局在天然结缕草种群和实验结缕草种群中明显不同。天然结缕草种群的营养繁殖体由根状茎、匍匐茎和分株直立茎的分蘖节组成，而且以根状茎为主，根状茎对芽和苗的贡献率分别大于95%和90%；实验结缕草种群的营养繁殖体由分蘖节和匍匐茎组成，二者扮演着缺一不可、同等重要的作用。实验结缕草种群营养繁殖力随着时间以指数函数增加；天然结缕草种群营养繁殖力随着时间在不同时期内大小差异较大，且不同样地间繁殖力的时间格局存在不同，火烧样地和未火烧样地在生殖期以前和果后营养期均表现出较高的营养繁殖力，而林下样地在生长季的前期其营养繁殖力较低，至7-8和9-10月份间各出现一个峰值。
7.不同生境结缕草种群有性生殖株的数量特征主要受控于环境资源的限制，但不同生境结缕草种群有性生殖株的株高、花序长的不同变化及其比率关系蕴涵着其个体生殖生长具有自身调节策略。火烧并未改变每花序种子数、花序长、生殖生长比率、每花序种子重及生殖分配与生殖株高度间的定量关系，但火烧极显著地增加了生殖株密度，极显著地降低了结缕草每花序种子数，对千粒重没有显著影响，种子单位产量是未火烧的1.85倍。火烧样地土壤种子库中结缕草种子的数量也相应地高于未火烧样地的。
8.牛、羊通过自己的粪便散布结缕草种子，牛粪、羊粪中结缕草种子的数量分别为101.70、6.88粒/g，牛粪和羊粪中结缕草种子的发芽率均高于对照的，牛粪中建植的种苗数是134.40株/牛粪样，羊粪中建植的种苗数是0.17株/羊粪球。羊粪的质地对种苗的建植有不利的影响，牛粪的没产生不利的影响，而且在自然界中牛粪样块还可以作为更新斑块，更有利于结缕草种群的有性更新。
9.结缕草种群的总生物量及各构件生物量均随着种群密度的增大而以线性函数增加，各构件生物量与总生物量以及各构件生物量间亦均表现为线性函数关系,总生物量增量的56.43％分配给分株，43.57％分配给根状茎，分株构件生物量增量的49.77％分配给叶片，50.23％分配给叶鞘，叶片、叶鞘和根状茎构件生物量的积累按1：1.1：1.6比例形成。
Zoysia japonica Steud. is a typical clonal grass plant with underground rhizomesand aboveground stolons, which is mainly distributed in the Liaodong and JiaozhouPeninsulas in China. By establishing the experimental Z. japonica clone and samplingthe natural Z. japonica populations, this paper analysed the dynamics of Z. japonicaclone and populations, and studied the life history patterns of Z. japonica populations.The results indicated that:
1. The ramets of Z. japonica clones increased with time by exponent function,the main stolon elongated with time by linear function, and the number of spacers onthe main stolon increased with time by logarithm function. The radius and area ofclones expanded with time by linear and exponent function, respectively.
2. The number of Z. japonica ramets changed little in un-burned site andunder-forest site in2006, and large in burned site and in under-forest site in2005. Thechanging trends of bud numbers were similar in three sites. The length of rhizomeswas longer at the beginning and end of the growth season than in the middle of thegrowth season in these three sites.
3. The mass of litter decreased in un-burned site and almost did not change inburned site with time. The number and biomass of other species were both lower inburned site than un-burned site. The burning improved the ecological condition andcould lead other species and their seeds to die. Thus, the numbers of all Z. japonicamodules were higher in burned site than un-burned site. The modular numbers werelower in under-forest site than un-burned site due to the limitation of light and soil.
4. In the structure of stage classes, the percentage of vegetative adult ramets waslargest (more than64.5%), and the those of buds and juvenile ramets were18.5%and14.2%, respectively. In the structure of biomass, the percentage of rhizome mass waslargest (51.7%), and the those of buds and foliage mass were24.9%and23.3%,respectively.
5. The population growth rate (λ) of Z. japonica in burned site was highest（λ=1.1247） between May and June, and less than one （λ=0.557）betweenSeptember and October. The population growth rate in un-burned site was highest（λ=1.1374）betweenAugust and September, and less than one （λ=0.674）betweenSeptember and October. The population growth rate in under-forest site was highest（λ=1.1904） between September and October, and more than one in all growthseason. The main life history pathway was adult ramet pathway‘in three sites, whichmeant that the stasis of adult ramets made the biggest contribution to λ to keep thepopulation relatively steady. The increased λ resulted from the contributions ofvegetative propagation and growth of transition classes.
6. The propagules were composed of the rhizomes, stolons and ramets. Thecontribution rates of rhizome to buds and juvenie ramets were more than95%and90%in the natural Z. japonica populations, respectively. However, in experimental Z. japonica clones, the propagules were only composed of the stolons and ramets whichplayed the same important roles. The vegetative fertility of experimental Z. japonicaclone was increased with time by exponent function. The fertility of the natural Z.japonica populations varied between different periods and its time patterns weredifferent among sites.
7. Burning extremely significantly decreased the number of seed per spike, andinsignificantly affected the weight of1000seeds, but extremely significantlyincreased the density of reproductive ramets. Thus, the seed yield in burned site was1.85times as high as in non-burning site. The quantitative relationships of thosequantitative characters to the height of reproductive ramets were same in burned andun-burned sites, respectively.
8. The number of Z. japonica seeds in droppings of cattle and sheep per gramwas101.7and6.88, respectively. The ingestions of cattle and sheep increased thegermination percentage, germination energy, and vigor index of Z. japonica seeds,and shortened their germination time, but did not affect the viability; A number ofseedlings (134.40seedlings/pat) were established from cattle dung pat. A fewseedlings (0.17seedlings/pellet) were established from sheep pellets. These resultssuggest that cattle and sheep have the relevant potential of endozoochory as Z.japonica seed dispersers.
9. The total biomass and the biomass of modules all increased with thepopulation size(population density), the biomasses of modules also increased withtotal biomass, and the biomasses of leaf, leaf sheath modules also increased with thebiomass of ramets all by the linear equation. The56.43％and43.57％of theincreased total biomass was allocated to ramets and rhizoms, and the50.23％and49.77％of the increased biomass of ramet was allocated to leaf sheath and leaf,respectively.
1.在栽培条件下，经过一个生长季的生长，结缕草实验无性系大小的可塑性较大，但均遵循着规律性的时间生长和空间扩展。随着生长时间的进程，无性系分株构件种群以指数函数规律增加，匍匐茎主茎长度以直线函数规律伸长，主茎间隔子数量则以对数函数规律增加。在空间扩展上，无性系半径随着时间以直线函数规律延伸，面积随着时间以指数函数规律扩展，分株种群的数量则随着半径的延伸以指数函数规律增加，随着面积的扩展以直线函数规律增加，主茎间隔子数量随着半径的延伸以直线函数规律增加，随着面积的扩展以对数函数规律增加。
2.对于天然结缕草种群，在两个生长季内未火烧样地的分株构件种群波动较小，火烧样地的在6、7月份出现密度峰值，波动较大，林下样地的分株密度在2005年的5月和10月出现峰值，在2006年波动很小。3个样地结缕草芽构件种群的变化趋势相似，在10月份达到密度峰值。火烧样地和未火烧样地的苗构件种群在生长季内的变化趋势相似，林下样地的则与其分株种群的变化趋势相似。结缕草根状茎的长度在3个样地中均是在生长季的两端高，在生长季的中间期较低。
3.未火烧样地的枯落物生物量随着时间逐渐减少，而火烧样地枯落物生物量在整个生长季内虽有上下波动但幅度不大。在生长季内各月份火烧样地的其他植物种株数、生物量均小于未火烧样地的。火烧清除了样地的枯落物，改善了结缕草生长的生态条件，同时火烧能够导致其他植物种的植物体及种子死亡，而对结缕草自身的影响较小，因为结缕草主要靠营养繁殖，其主要繁殖体根状茎储藏在地下，分株直立茎的分生生长点基本贴于地表或潜在地下，而且受到叶鞘的包覆保护。这样，火烧样地结缕草种群各构件的数量整体上均高于未火烧样地的。林下样地由于受到光照和土壤条件的限制，各构件的数量整体上低于未火烧样地的。
4.在未火烧样地、火烧样地和林下样地各月份的结缕草种群阶段结构中，整体上营养株所占的百分比均最高，平均64.5%以上，芽所占的百分比次之，平均为18.5%，除了生殖株以外苗所占的百分比最小，平均为14.2%。在结缕草种群生物量结构中，整体上茎生物量所占的百分比均最高，平均为51.7%，叶鞘生物量所占的百分比次之，平均为24.9%，叶片生物量的百分比最小，平均为23.3%。
5.火烧样地在5-6月间结缕草种群的增长率最大（λ=1.1247），而在9-10月间种群增长率（λ=0.557）明显小于1。未火烧样地在8-9月间种群增长率最大（λ=1.1374），而在9-10月间种群增长率（λ=0.674）明显小于1。林下样地在8-9和9-10月间种群增长率分别为λ=1.1141和λ=1.1904，在整个生长季未出现负增长（λ <1）。3个样地结缕草种群生活史的主路径均是“营养株路径”，生长季的各月份间营养株的保持对种群增长率的贡献最大，维系着种群的相对稳定。种群增长率的增加主要来自于转移等级的繁殖和生长对种群增长率的贡献。
6.营养繁殖在无性系植物种群生活史格局中处于核心地位。结缕草种群营养繁殖格局在天然结缕草种群和实验结缕草种群中明显不同。天然结缕草种群的营养繁殖体由根状茎、匍匐茎和分株直立茎的分蘖节组成，而且以根状茎为主，根状茎对芽和苗的贡献率分别大于95%和90%；实验结缕草种群的营养繁殖体由分蘖节和匍匐茎组成，二者扮演着缺一不可、同等重要的作用。实验结缕草种群营养繁殖力随着时间以指数函数增加；天然结缕草种群营养繁殖力随着时间在不同时期内大小差异较大，且不同样地间繁殖力的时间格局存在不同，火烧样地和未火烧样地在生殖期以前和果后营养期均表现出较高的营养繁殖力，而林下样地在生长季的前期其营养繁殖力较低，至7-8和9-10月份间各出现一个峰值。
7.不同生境结缕草种群有性生殖株的数量特征主要受控于环境资源的限制，但不同生境结缕草种群有性生殖株的株高、花序长的不同变化及其比率关系蕴涵着其个体生殖生长具有自身调节策略。火烧并未改变每花序种子数、花序长、生殖生长比率、每花序种子重及生殖分配与生殖株高度间的定量关系，但火烧极显著地增加了生殖株密度，极显著地降低了结缕草每花序种子数，对千粒重没有显著影响，种子单位产量是未火烧的1.85倍。火烧样地土壤种子库中结缕草种子的数量也相应地高于未火烧样地的。
8.牛、羊通过自己的粪便散布结缕草种子，牛粪、羊粪中结缕草种子的数量分别为101.70、6.88粒/g，牛粪和羊粪中结缕草种子的发芽率均高于对照的，牛粪中建植的种苗数是134.40株/牛粪样，羊粪中建植的种苗数是0.17株/羊粪球。羊粪的质地对种苗的建植有不利的影响，牛粪的没产生不利的影响，而且在自然界中牛粪样块还可以作为更新斑块，更有利于结缕草种群的有性更新。
9.结缕草种群的总生物量及各构件生物量均随着种群密度的增大而以线性函数增加，各构件生物量与总生物量以及各构件生物量间亦均表现为线性函数关系,总生物量增量的56.43％分配给分株，43.57％分配给根状茎，分株构件生物量增量的49.77％分配给叶片，50.23％分配给叶鞘，叶片、叶鞘和根状茎构件生物量的积累按1：1.1：1.6比例形成。
Zoysia japonica Steud. is a typical clonal grass plant with underground rhizomesand aboveground stolons, which is mainly distributed in the Liaodong and JiaozhouPeninsulas in China. By establishing the experimental Z. japonica clone and samplingthe natural Z. japonica populations, this paper analysed the dynamics of Z. japonicaclone and populations, and studied the life history patterns of Z. japonica populations.The results indicated that:
1. The ramets of Z. japonica clones increased with time by exponent function,the main stolon elongated with time by linear function, and the number of spacers onthe main stolon increased with time by logarithm function. The radius and area ofclones expanded with time by linear and exponent function, respectively.
2. The number of Z. japonica ramets changed little in un-burned site andunder-forest site in2006, and large in burned site and in under-forest site in2005. Thechanging trends of bud numbers were similar in three sites. The length of rhizomeswas longer at the beginning and end of the growth season than in the middle of thegrowth season in these three sites.
3. The mass of litter decreased in un-burned site and almost did not change inburned site with time. The number and biomass of other species were both lower inburned site than un-burned site. The burning improved the ecological condition andcould lead other species and their seeds to die. Thus, the numbers of all Z. japonicamodules were higher in burned site than un-burned site. The modular numbers werelower in under-forest site than un-burned site due to the limitation of light and soil.
4. In the structure of stage classes, the percentage of vegetative adult ramets waslargest (more than64.5%), and the those of buds and juvenile ramets were18.5%and14.2%, respectively. In the structure of biomass, the percentage of rhizome mass waslargest (51.7%), and the those of buds and foliage mass were24.9%and23.3%,respectively.
5. The population growth rate (λ) of Z. japonica in burned site was highest（λ=1.1247） between May and June, and less than one （λ=0.557）betweenSeptember and October. The population growth rate in un-burned site was highest（λ=1.1374）betweenAugust and September, and less than one （λ=0.674）betweenSeptember and October. The population growth rate in under-forest site was highest（λ=1.1904） between September and October, and more than one in all growthseason. The main life history pathway was adult ramet pathway‘in three sites, whichmeant that the stasis of adult ramets made the biggest contribution to λ to keep thepopulation relatively steady. The increased λ resulted from the contributions ofvegetative propagation and growth of transition classes.
6. The propagules were composed of the rhizomes, stolons and ramets. Thecontribution rates of rhizome to buds and juvenie ramets were more than95%and90%in the natural Z. japonica populations, respectively. However, in experimental Z. japonica clones, the propagules were only composed of the stolons and ramets whichplayed the same important roles. The vegetative fertility of experimental Z. japonicaclone was increased with time by exponent function. The fertility of the natural Z.japonica populations varied between different periods and its time patterns weredifferent among sites.
7. Burning extremely significantly decreased the number of seed per spike, andinsignificantly affected the weight of1000seeds, but extremely significantlyincreased the density of reproductive ramets. Thus, the seed yield in burned site was1.85times as high as in non-burning site. The quantitative relationships of thosequantitative characters to the height of reproductive ramets were same in burned andun-burned sites, respectively.
8. The number of Z. japonica seeds in droppings of cattle and sheep per gramwas101.7and6.88, respectively. The ingestions of cattle and sheep increased thegermination percentage, germination energy, and vigor index of Z. japonica seeds,and shortened their germination time, but did not affect the viability; A number ofseedlings (134.40seedlings/pat) were established from cattle dung pat. A fewseedlings (0.17seedlings/pellet) were established from sheep pellets. These resultssuggest that cattle and sheep have the relevant potential of endozoochory as Z.japonica seed dispersers.
9. The total biomass and the biomass of modules all increased with thepopulation size(population density), the biomasses of modules also increased withtotal biomass, and the biomasses of leaf, leaf sheath modules also increased with thebiomass of ramets all by the linear equation. The56.43％and43.57％of theincreased total biomass was allocated to ramets and rhizoms, and the50.23％and49.77％of the increased biomass of ramet was allocated to leaf sheath and leaf,respectively.
引文
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