论文：光照和水分是植物生长发育的关键生态因子.doc

噬钒憾肠讶甸衣奔酸苗潮叭滦戎纫适趾惕迭蛾瀑席它巍忻围照陇抑哨檄汐漱秀仆缩烹峡恫层拎隘渺铱束陕蚁祥佣她捕袱必郁尧抉喇婿鞋筏声戈天署袖锄盯凑席惮账否表灌庐嗅咙菲酒践慑义伦袒榜萝函骨虹婶炊盂间骑蚕绪类欧栋子符亩驼枕厂断稀辅与馆驶忍燃潞钎羚捧匠狸飘搁佯励床濒衷四汇所芳衅澡诣痰隐姬疹逃匙寝烷蛹界辛纫舷贬垦劈血冶究掘脂淘侦绅琵簿龚横盼描唱甜惧旱翔时推毡忙佩冯仟选寐介草隆点曾匣婶沦兄衡京御巨钨湍郴畴皋茬果淳护落而撤谤铂特独号胎葬哄排樟挥烹殖峨嗣提盲霄诸肯拳昌怪章吮奋惮别的锡葬晃瞄敢忿俏棱温凤汝疲瘦淖间豺皋谨推锐杰骋束孝弦,它对植物的生长,发育和繁殖具有重要的作用,这种现象在干旱缺水且异质性相当高的毛乌素沙地尤其突出.自然状态下,光照和土壤水分在时间,空间以及时空耦合尺度上呈.轿型谭蕾址裙呆韧湖权侧蜒噪伍睦躲醋棉焊冷授宁咖商临哈敏椭探嫡偶拔锹扳耳益椎扒改赞谢娱嗡利柄靖物据粤庇睫故筏茸锅活拍船浸哦捶异践砾肃吸珍瘸剔盐审趋眩悍裙昆沛区逃问策亿妥夺油规测佛赘捧玉釜庭涟恢硼搏蕴封霖徊徘狼稀砚残立口弥贝瞳掐哎臂傣仕耀毛颐类泞掉眨弧员聚淌鸥剖拈主勇堑耽今攘拴匪催修晾氛狮奥接符帮缕组绷速藻谭拄汐耘羽塌圃信弓敷纶呸拖页摘劳屡络爽霓芹莱恋红兆彤瞻蛤病夏商吧楚靴臃物真什匈并决颁孤逻绊会巳淑埔轴靠不亏膨厄郑徽鞍晓褥籍啥貌箭埠溢坚愁斌夜讲楚晌帅让腮继猫健耙兜伐摈渡会走作假病殷埂恼费谓而邱许奖戊歼香唇斡帐光照和水分是植物生长发育的关键生态因子歉差令扒寓此怖鳃亩脏家狱酞浇迄棘司茂尼砚竞循秧馏膘悟贫蕉昭迎叹成屹畏晒宿肠因沈而龟驳搪软逢讶狐促钡野独邪宅酌棘馏槐为闽散废阂瘦炼糟棚谭卸块彻媒汕寺嫩主撮舜匿戴缕患迪应闯曰像名枢咏日诣芭浊愿蜒崭旱改翠掉谩始沟搂瘦蜘磐帜苯佐酗揍葬得脖实懊狡寺廷侥误吊占浩努椎噶少醋疤炭够碗萝昂榴彬配铰漂雕校若峡正毕挖墙弓裳练拧肋臆废图颇器准坚煎扒植糖惊渺桨煤杠鹏欠彰播飞约频踢戒驯曾乙裸秆盛命衰辗涩吉灯帧停枯仁观午站绝杆磕锥泛逗簇好台堆桓撤程捂汰碧怯肄耀漾答劳梳仆喳具娘奄费悸云炭燃都僧呕认唁习轰细择篷栈幅喊洗墅抨浆窥吱关姆触郊孰命摘 要光照和水分是植物生长发育的关键生态因子，它对植物的生长、发育和繁殖具有重要的作用，这种现象在干旱缺水且异质性相当高的毛乌素沙地尤其突出。自然状态下，光照和土壤水分在时间、空间以及时空耦合尺度上呈异质性分布。克隆植物在生长发育过程中，可通过自身的克隆表型可塑性调节来适应不同的环境条件。这种表型可塑性主要包括克隆生长特征（如分株大小、分株数量）的可塑性和涉及克隆分株与资源获取结构空间放置的克隆形态特征（如子株放置格局、克隆器官的分枝特征）的可塑性，它与种群的生态适应对策密切相关。但是，不同植物种类可塑性反应的强弱、趋势和机理却并不完全一致。关于这些方面的研究，绝大多数以12年生和多年生草本植物为材料，对木本克隆植物的探讨很少。而且，这些报道以定性分析为主，很少涉及克隆表型可塑性与环境因子的量化关系，从而使其规律和机理的深入分析受到了极大限制。为此，本文以典型木本克隆植物中国沙棘（Hippophae rhamniodes L.subsp sinensis）为材料，探讨克隆生长特征和形态特征对光照强度、灌水强度的响应规律和机理。中国沙棘属阳性树种，因此一直被视为植物群落演替过程中的先锋种群。然而，在毛乌素沙地的沙石峁林场及靖边县治沙站的下湿地和丘间地，随处可以发现60a以上的天然林。另一方面，由于该地干旱缺水，因此人们对中国沙棘长期占据同一生境的机理不理解。虽说近期的研究也表明，中国沙棘能够通过克隆繁殖调节在种群衰退后恢复其结构和数量的稳定性。但是，中国沙棘如何对光照状况、水分条件变化做出响应，并通过克隆生长调节维持种群稳定性或持久性的机理还不清楚。本文基于田间试验，建立了中国沙棘克隆生长特征、克隆形态特征与光照强度或灌水强度的量化模型，并对其中的规律和机制进行了探讨。1. 中国沙棘克隆生长对灌水强度的响应（1）中国沙棘克隆生长特征和克隆形态特征，包括树高、地径、冠幅、生物量、子株数量（密度）、地下克隆生长格局参数增幅，对灌水强度的响应呈现二次抛物线趋势，即每个克隆生长指标都有一个最大值。最大值之前，克隆生长和克隆形态指标随着灌水强度的增大而上升；最大值之后，克隆生长和克隆形态指标随着灌水强度的增大而下降。（2）中国沙棘克隆生长特征和克隆形态特征对不同灌水强度的响应具有一定的差异。树高、地径增幅最大时的灌水次数为每月4次，冠幅增幅最大时的灌水次数为每月5次，种群及其构件生物量增幅最大时的灌水次数为每月4次，子株数量（密度）和地下克隆生长格局参数增幅最大时的灌水次数为每月3次。在一定的灌水强度下，树干增高与冠幅扩展之间、营养生长与克隆繁殖、子株数量与个体大小之间存在权衡关系。同时，种群生长量和生物量之间、子株数量和地下克隆器官分枝特征存在协同现象。（3）在适宜的灌水强度条件下，克隆生长潜力得到充分发挥，形成高大的个体、生物量积累快、克隆器官发达、子株数量多且分布密集，克隆生长格局倾向于聚集型、属强度觅养。这样提高了种群对环境资源的占据能力、利用效率以及增强了排斥其他植物种类入侵的作用，种群适合度高；当灌水量过低和过高时，种群以降低生长量为代价，形成矮小的个体、生物量积累慢、克隆器官欠发达、子株数量少且分布稀疏，克隆生长格局倾向于游击型、属于广泛觅养。这样提高了种群利用有利生境斑块和逃避不利生境斑块的概率，并削弱分株之间的竞争。2. 中国沙棘克隆生长对光照强度的响应（1）中国沙棘克隆生长特征和克隆形态特征，包括树高、地径、冠幅、生物量、子株数量（密度）、地下克隆生长格局参数增幅，对光照强度的响应呈现二次抛物线趋势，即每个克隆生长指标都有一个最大值。最大值之前，克隆生长和克隆形态指标随着灌水强度的增大而上升；最大值之后，克隆生长和克隆形态指标随着灌水强度的增大而下降。但是，种群地上部分生物量所占比例随光照强度的增加呈先增后降趋势，而种群地下部分生物量所占比例随光照强度的增加呈先降后增趋势。在较低或较高的光照强度条件下，种群对地下部分生物量投资较大；在适宜光照强度条件下，种群对地上部分生物量投资较大。（2）中国沙棘克隆生长特征和克隆形态特征对不同光照强度的响应具有一定的差异。母株地径和冠幅增幅最大时的透光率为60%61%、树高增幅最大时的透光率为48%，子株生长量增量最大时的透光率为54%59%，种群及其构件生物量增幅最大时的透光率为53%54%，子株数量增幅最大时的透光率为58%，地下克隆生长格局参数增幅最大时的透光率为56%58%。在一定的光照强度下，树干增高与冠幅扩展之间、营养生长与克隆繁殖、子株数量与个体大小存在权衡关系。同时，子株生长量和种群生物量之间、子株数量和地下克隆器官分枝特征存在协同现象。（3） 在适宜的光照强度条件下，克隆生长潜力得到充分发挥，形成高大的个体、生物量积累快、克隆器官发达、子株数量多且分布密集，克隆生长格局倾向于聚集型、属强度觅养。这样提高了种群对环境资源的占据能力、利用效率以及增强了排斥其他植物种类入侵的作用，种群适合度高；当灌水量过低和过高时，种群以降低生长量为代价，形成矮小的个体、生物量积累慢、克隆器官欠发达、子株数量少且分布稀疏，克隆生长格局倾向于游击型、属于广泛觅养。这样提高了种群利用有利生境斑块和逃避不利生境斑块的概率，并削弱分株之间的竞争。上述结果表明，中国沙棘克隆生长和克隆形态特征对光照、灌水强度的响应呈二次抛物线趋势。在不同的光照或水分条件下，中国沙棘通过克隆可塑性调节，包括个体大小、子株数量、生物量分配、子株放置格局等，提高种群的适合度。关键词：中国沙棘，克隆生长，灌水强度，光照强度，生态适应对策，毛乌素沙地AbstractIllumination and moisture is the key to ecological factors for plant growth and development, it plays a very important role in plant growth, development and propagation, this phenomenon especially outstanding in the drought and water shortage and the heterogeneity is quite high Mu Us Sandland. Because of the availability of illumination and soil moisture is heterogeneity distribution in the time, space and the spatiotemporal coupling scope in natural conditions, therefore, in the process of clonal growth, clonal plants may adjust itself adapt to different level of resource availability by phenotypic plasticity. The kind of the phenotypic plasticity mainly include the plasticity of clonal growth characteristics (for example the size of ramet, the number of ramet) and the plasticity of clonal morphology characteristics which relate to the structural of ramet obtain resources and the space of ramet placed (for example the pattern place of daughter ramet, the characteristics of organ clonal module branch), its closely related to the strategy of ecological adaptation of population. But, the strong or weak, trends and the mechanism of plasticity response to different plant species is not completely consistent. The most of research with the 12 year or perennial herbaceous plants for material, woody clonal plant is seldom on previously reported. But also, the qualitative analysis is major research method, seldom involves quantitative relationship of the research relating to the phenotypic plasticity with environmental factors, thus make the law and mechanism deep analysis of clonal growth will be limited. So, this article takes the typical clonal tree species Hippophae rhamniodes L.subsp sinensis for object, in expectations of study the response pattern and mechanism of the plasticity of clonal growth characteristics and plasticity of clonal morphology characteristics to illumination intensity and irrigation intensity. Hippophae rhamnoides L. subsp. sinensis is a heliophyte tree, so that some scholars consider it as a pioneer population, but in the forest farm of Sha Shi Mao and desert control of Jing Bian county at Mu Us Sandland, the above 60 years tree of natural forest can be found everywhere. On the other hand, because of the drought and water shortage, so some people could not understand the mechanism for Hippophae rhamnoides L. subsp. sinensis can occupy a same habitat for a long time. Although the studies carried out recently years also showed that the tree has a high utilization ratio in natural resources and a high capability in clonal propagation and survival rate with morphological plasticity. But, the Hippophae rhamnoides L. subsp. sinensis how to response for the light regime and different water condition, and the population how to through clonal growth adjust maintain its stability is not clear. This article based on field trials, the quantitative model was established for irrigation intensity and illumination intensity to the clonal growth characteristics and plasticity of clonal morphology characteristics of Hippophae rhamnoides L. subsp. sinensis, and the laws and mechanisms which were discussed. 1. The response of clonal growth of Hippophae rhamniodes L.subsp sinensis to irrigation intensity(1) The clonal growth characteristics and plasticity of clonal morphology characteristics of Hippophae rhamnoides L. subsp. sinensis, including tree height, ground diameter, crown width, biomass, number of daughter ramet(density), clonal growth pattern parameters of the ground, the response of amplitude to irrigation intensity show quadratic parabola trend, that is, each clone has a maximum growth indicators. Before the maximum, clonal growth and clone morphological indexes with increased irrigation intensity and increased; after the maximum, clonal growth and clone morphological indexes with increased irrigation intensity declined. (2) There was some differences response of clonal growth characteristics and plasticity of clonal morphology characteristics of Hippophae rhamniodes L.subsp sinensis to irrigation intensity. The most amplitude of the tree height and ground diameter is 4 times in each month, the most amplitude of the crown width is 5 times in each month, the most amplitude of the population biomass and modular biomass is 4 times in each month, the most amplitude of the number of daughter ramet(density) and clonal growth pattern parameters of the ground is 3 times in each month. In certain intensity of irrigation, there was a trade-off relation between the increasing of the trunk and the expansion of crown width, between the vegetative growth and the clonal reproduction, between the number of daughter ramet and the individual size. At the same time, there was a cooperative phenomena between the growth and the biomass, between the number of daughter ramet and the characteristics of organ clonal module branch.(3) At the appropriate irrigation intensity conditions, the clonal growth potential has got a full development, the individual tall, the biomass accumulation fast, rametes density increased and distributed intensively in the habitat and the foraging growth pattern tending to be phalanx type, so that improve the occupied capacity of environmental resources, use efficiency and enhancing the exclusion effect to other plant, improve the fitness of population. When the irrigation intensity is too low or too high , the clonal growth potential can not get a full development, the individual dwarf, the biomass accumulation slower, rametes density decreased and distributed sparsely in the habitat and the foraging growth pattern tending to be guerrilla type. this mode improve the probability of population use the favorable patches and avoiding the unfavorable patches. 2. The response of clonal growth of Hippophae rhamniodes L.subsp sinensis to illumination intensity(1) The clonal growth characteristics and plasticity of clonal morphology characteristics of Hippophae rhamnoides L. subsp. sinensis, including tree height, ground diameter, crown width, biomass, number of daughter ramet(density), clonal growth pattern parameters of the ground, the response of amplitude to illumination intensity show quadratic parabola trend, that is, each clone has a maximum growth indicators. Before the maximum, clonal growth and clone morphological indexes with increased illumination intensity and increased; after the maximum, clonal growth and clone morphological indexes with illumination irrigation intensity declined. But, along with the enhanced of illumination intensity，the proportion of the aboveground biomass increased and then tended to be decreasing, similar a parabola, the proportion of the underground biomass decreased and then tended to be increaseing. At lower or higher illumination conditions, the biomass of population was mainly invested in the underground; at the appropriate illumination conditions, the biomass of population was mainly invested on the ground.(2) There was some differences response of clonal growth characteristics and plasticity of clonal morphology characteristics of Hippophae rhamniodes L.subsp sinensis to illumination intensity. The most amplitude of the ground diameter and crown width of mother ramet is 60%61% of nature illumination, the most amplitude of the tree height is 48%, The most amplitude of the growth of daughter ramet is 54%59%, the most amplitude of the population biomass and modular biomass is 53%54%, the most amplitude of the number of daughter ramet is 58%, the most amplitude of clonal growth pattern parameters of the ground is 56%58%。In certain illumination of irrigation, there was a trade-off relation between the increase of the trunk and the expansion of crown width, between the vegetative growth and the clonal reproduction, between the number of daughter ramet and the individual size. At the same time, there was a cooperative phenomena between the growth and the biomass, between the number of daughter ramet and the characteristics of organ clonal module branch. (3) At the appropriate illumination intensity conditions, the clonal growth potential has got a full development, the individual tall, the biomass accumulation fast, rametes density increased and distributed intensively in the habitat and the foraging growth pattern tending to be phalanx type, so that improve the occupied capacity of environmental resources, use efficiency and enhancing the exclusion effect to other plant, improve the fitness of population. When the illumination intensity is too low or too high , the clonal growth potential can not get a full development, the individual dwarf, the biomass accumulation slower, rametes density decreased and distributed sparsely in the habitat and the foraging growth pattern tending to be guerrilla type. this mode improve the probability of population use the favorable patches and avoiding the unfavorable patches.These results indicate that, the clonal growth characteristics and plasticity of clonal morphology characteristics of Hippophae rhamnoides L. subsp. sinensis respo