【作者】 王小红；

【导师】 周祖基；

【作者基本信息】 四川农业大学 ， 森林培育学， 2007， 博士

【摘要】 竹子是重要的森林资源，同时某些竹种也是“国宝”大熊猫的主要食物来源。四川省雅安地区大面积水竹开花不仅给当地农民的实际生产造成很大损失，对中国雅安碧峰峡大熊猫繁育基地大熊猫的食物来源也造成一定的威胁。本论文以碧峰峡周边地区的开花水竹和慈竹为研究对象，对其开花期间生物学特性及物质代谢规律进行了研究，并在此基础上对水竹花后复壮和更新进行了试验。主要结果如下：1开花水竹林的出笋时间比未开花竹稍晚，但整个发笋过程与未开花竹基本一致，出笋时间持续约60天。水竹发笋和成竹率较高，平均成竹率为73.4％，年平均发笋数和成竹数分别达到了2.44株／m²和1.76株／m²，每亩产笋量可达一千株以上；开花竹林的出笋量和成竹率明显低于未开花竹林，总的出笋量只有未开花竹林的24％，年平均发笋数只有0.62株／m²，且平均成竹率也只有37.8％，说明随着竹子开花，竹子的生活力降低，竹林的生产力明显衰退。未开花和开花水竹的高生长均符合Logistic生长曲线，但开花水竹高生长能力明显弱于未开花水竹，平均最终高度只有未开花竹的36.9％。对环境因子的定期观察和分析结果表明，影响水竹出笋的决定因素是气温和地温。在影响水竹高生长的生态因子中，最低气温与高生长之间的Logistic模型模拟拟合度最好，从一定程度上说明最低气温是影响水竹高生长的主要生态因子。同时，根据水竹发笋和生长规律可以选择合理的移栽和施肥时间。2水竹在雅安地区的开花表现为成片开花，花期较长，从11月开始到次年的4月中旬左右结束，持续约5个月。自然授粉条件下，水竹结实率极低，对水竹进行人工授粉试验仍未能得到种子。水竹属于复穗状花序，镜下检测发现，花药比较丰富且花粉粒活力较高，但柱头和子房常常发育不良或者畸形，且柱头可授性也极低，几乎为零；对水竹花进行解剖检查，发现在水竹花里有少量幼虫，有虫的水竹雌雄蕊完全被破坏。推测水竹丰富的花药和花粉与雅安地区多雨潮湿的气候条件可能为该类昆虫的繁殖提供了有利条件，导致其幼虫破坏水竹雌雄花器官的发育。与水竹相比，慈竹花期较长，通常从12月开始至次年6月左右结束，历时约8个月。慈竹花粉丰富，但花粉粒活力较低。目前没有发现慈竹结实，初步认为与花粉粒活力较低和雅安独特的多雨气候有关。3雅安地区刺竹和水竹开花年（2001年-2006年）1月均温分别高出未开花年（1985年-2000年）和多年（1985年-2006年）平均值0.29℃和0.225℃；7月均温分别高出未开花年的多年平均值0.958℃和0.697℃；开花年年降水量分别比未开花年和多年平均值少124.9mm和90.8mm。而对开花和未开花水竹林小生境生态因子的调查结果显示，处于阳坡、光照条件好的样地开花明显多于处于阴坡、光照条件较差的样地；林内温度和湿度与竹子开花显著相关，开花样地内早上9点和中午2点平均温度和湿度均比未开花样地高。总的来看，充足的光照、较高的林内温度、湿度以及高温和少雨在一定程度上起到了诱导和促进竹子开花的作用。4对水竹和慈竹开花期间叶片相关代谢物质含量的测定结果显示，激素GA₃对水竹和慈竹的开花有一定促进作用，ABA和ZT表现为抑制作用，IAA对竹类植物开花的影响不大。未开花水竹和慈竹的叶绿素a、叶绿素b及叶绿素总量变化基本一致，且两者间差异不显著。开花水竹和慈竹尤其是开花逆转水竹叶绿素a、叶绿素b以及叶绿素总量变化有明显的波动，且伴随开花进程含量持续增加。开花水竹和慈竹可溶性糖、总糖含量以及蔗糖酶活性在开花期间均呈上升趋势，而未开花水竹和慈竹蔗糖酶活性呈下降趋势；开花水竹和慈竹中淀粉和总糖含量、淀粉酶及蔗糖酶活性在开花初期均显著增加，随后减小，后期又增加。水竹和慈竹叶片中可溶性蛋白质含量是随生育期推进逐渐增加，但开花竹含量和增加幅度明显小于未开花和开花逆转水竹；开花水竹和慈竹GS酶活性显著低于未开花和开花逆转水竹。开花水竹和慈竹可溶性糖／可溶性蛋白质均高于开花竹，而总糖／全氮比值无显著差异。测定和分析结果说明进入开花阶段后，水竹和慈竹碳代谢活动增强而氮代谢降低，这从一定程度上反映出碳氮代谢的强弱能预示着植物从营养生长向生殖生长的转化。5通过对开花初期水竹和慈竹与代谢有关的指标进行主成分分析，筛选出对水竹和慈竹开花起关键作用的主要代谢因子。分析结果显示，GA₃、GA₃／ABA和淀粉是这两种竹子开花的关键物质，其次是可溶性糖、淀粉酶和GS酶活性；而ZT、ZT／ABA、淀粉酶是对开花逆转过程起重要作用的关键物质。即一定浓度的GA₃、GA₃／ABA以及淀粉和可溶性糖起到了促进竹子开花的作用，而合适浓度的ZT和ZT／ABA又可以促使水竹开花的逆转。为使竹子开花的原理更趋明朗化，在前面所有研究的基础上，再结合其他植物开花研究的成果，建立了水竹开花进程的初级模拟图。模型显示竹子开花是一个漫长而复杂的过程，如果在此过程中任一环节被影响或打断均有可能加速或者停止生长和开花边程，这也正是竹类植物开花具有更多变数的原因。6通过对开花水竹林采用不同复壮措施进行复壮更新和抑制开花进程试验，并结合大熊猫的实际喂养需求，认为最佳复壮措施为在水竹始花时进行皆伐，砍去竹林内所有竹子，然后在第二年新发竹林内只砍除开花竹。该措施下竹鞭中可溶性糖、总糖、可溶性蛋白质、总糖／蛋白质总量最高，蔗糖酶和谷氨酰胺合成酶活性最大。这一结果很好解释了竹子开花期间的物质代谢机制、证实了地下茎的生长状态对竹类植物的发笋成竹能力起关键作用，说明地下茎中主要营养物质含量和相关酶活性的大小与其生长状态以及开花进程有着非常重要的联系。更多还原

【Abstract】 Bamboo is the important plant both as the forest resource and as the main food of Giant Panda, which is the on the verge of depopulation. Therefore, the great occurrence of Phyllostachys heteroclada olive flowering, resulting in the death of the bamboo, not only damaged the coming of local peasants, but also endangered the survival of bamboo. This study has investigated biological characteristics and carbon, nitrogen metabolism and rejuvenation of Phyllostachys heteroclada olive and Neosinocalamus affinis Keng f during the period of flowering. And the main result as followed:1. The development rule of Phy. Her. olive was investigated. The shooting time of flowering is later than that of the non-flowering bamboos. The development of shooting of flowering individuals is similar with that of the non-flowerings, and it has spent 60 days on shooting. The shooting production and proportion of grown up bodies of the flowering bamboos is clearly few than that of non-flowering bamboos, which indicated that both the bamboos’ vitality and forest production decreased evidently during the period of flowering. The height growth of flowering and non-flowering bamboos is consistent with the Logistic of curve of growth, but the capacity of height growth of non-flowering is evidently stronger.2. The whole population flowering at the same time is the main type of Phy. Het. olive in Ya’an, with lasting five months. Neither pollinating with artificial ways nor natural ones made bamboos seeding successfully. With observing the flower apparatus and microspore fertility, it was suggested that normal chapiter and germen and the damage of worms mainly contributed to the non-seeding of bamboos. Comparing with Phy. Het. olive, Neo. Aff Keng f has longer flowering period with eight months. The great number of pollens were found to have lower fertility, and by now seeding was not observed for Neo. Aff Keng f. The rain-rich weather is the main reason of Neo. Aff Keng f sterility.3. Both the average air temperature in January and July of lowering years （2001 to 2006） are higher than that of non-flowering （1985 to 2000）years, the average air temperature in January is higher than average of non-flowering years 0.29℃and 0.225℃of average from 1985 to 2006; the average air temperature in July of flowering years is also higher, it is higher than average of non-flowering years 0.958℃and 0.697℃of average from 1985 to 2006. The mean year-precipitation of flowering years is fewer than non-flowering years, it in flowering years is lower than non-flowering years 124.9 mm and 90.Smm from 1985 to 2006. The statistics also showed that bamboo flowering had the significant correlation with the forest temperature and humidity, both the temperature and humidity of flowering bamboo forest were higher than that of non-flowering forest. And the bamboos growing at the sunshiny area are more prone to flower than those growing at downsun area. generally, enough sunshine, higher forest temperature and humidity and dry can lead bamboo to flower.4. The content of some matter is related to metabolism in Pky. Het. olive and Neo. Aff Keng f leaves were measurated during flowering, the result showed that hormone GA₃ promote flowering, ABA and ZT restrain flowering, the influence of IAA is not obvious. The change of chlorophyll a, chlorophyll b and total amount of chlorophyll content in Phy. Het. olive and Neo. Aft. Keng f leaves showing no obvious difference. The change of chlorophyll a, chlorophyll b and total amount of chlorophyll content in flowering Pky. Her. olive and Neo. Aft. Keng f leaves, especially flower reversed Pky. Her. olive leaves obvious undulate, and the content increased along with flowering. The content of soluble sugar, total amount of sugar and activity of invertase in flowering Phy. Het. olive and Neo. Aff Keng f leaves are all increased at begginning of flower, activity of invertase in non-flowering Phy. Het. olive and Neo. Aff Keng f leaves reduced. The content of soluble protein in Phy. Het. olive and Neo. Aff Keng f leaves is increased along with growth, but the growing rate of flowering is smaller than non-flowering; the activity of GS of flowering bamboo is higher than non-flowering; the content of soluble sugar,/ soluble protein in flowering bamboo larger than non-flowering, total amount of sugar/total amount of protein is not obvious different between flowering and non-flowering bamboo. The result showed that the intensity of carbon and nitrogen metabolism can forebode the transform from vegetable growth to florescence.5 The key metabolism factors of blooming of Phyllostachys heteroclada olive and Neosinocalamus affinis Keng f were analyzed by using the principal component analysis method. The results show that GA₃, GA₃/ABA and starch enzymeis are the most important foctors, followed by soluble sugar, activity of starch enzyme and GS. Certain content of GA, starch and soluble sugar and GA/ABA could promote bamboo flower, while right content of ZT and ZT/ABA could restrain bamboo flowering. In order to show bamboo flowering more clearly, form the primary simulating model of bamboo flowering was constructed. And the model showed that the flowering of bamboo is lengthy and complicated. Any broken for the tache would possibly accelerate or terminate growing and florescence course, which could be the reason why bamboo florescence is so variable.6. The rejuvenation experiments that can restrain Phy. Het. olive florescence were carried out, according of feeding demand of local panda. The best measure of rejuvenation of Phy. Her. olive was suggested to fell all the plants at the beginning of florescence and only fell flowering plants next year. This measure led to high content of soluble sugar、total sugar、soluble protein and the ratio of total sugar and protein in rhizome with the most active GS. This result explained well that the mechanism of metabolize during florescence periods, and approved that the condition of rhizome growth was a key to reflect clone-growth and ability of bamboo shooting. All of the main nutrient matters content, GS activity of rhizome and morphological characteristics has important relationship with florescence procedure.更多还原