【作者】 刘悦秋；

【导师】 孙向阳；

【作者基本信息】 北京林业大学 ， 生态学， 2007， 博士

【摘要】 异株荨麻(Urticadioica L.subsp.dioica)原产于欧洲,是一种重要的食用和药用植物,具有很高的经济利用价值。通过对引种的异株荨麻形态结构、物候期、种子萌发条件以及对不同环境条件(光照、土壤水分和温度等)的生态适应性与抗逆生理生化特性的系统研究,明确了异株荨麻引种的关键影响因素与引种栽培条件,并根据异株荨麻原产地的地面气候资料,应用模糊相似优先比法对其在我国的适生区进行了区划。在此基础上,通过对施肥与其营养成分的关系、挥发油成分质谱分析以及黄酮类物质的提取与鉴定,探索了异株荨麻食用、药用开发利用价值。从而为异株荨麻的大面积引种栽培以及可利用价值研究奠定基础。获得的主要结果如下:通过石蜡切片和系统观察,明确了异株荨麻根、茎、叶解剖结构,并确定异株荨麻在北京地区的生育期为5～9月份,生长期为5月到11月中旬,播种最适宜时期为5月,实生苗生育期为105 d,生育期大于10℃的积温为1350℃,多年生苗生育期75～80d左右,生育期大于10℃的积温为1050℃;采用双因素随机区组设计方案,系统研究了温度和稀土、温度和赤霉素对异株荨麻种子萌发的影响。结果表明,异株荨麻种子在15～40℃恒温条件下均能发芽,但不同温度下种子发芽率和发芽势均差异显著,最适温度约为30℃。稀土溶液在25～500mg/L范围内,均可有效促进种子萌发,浓度为100 mg/L时,种子发芽势和发芽率均最高。赤霉素溶液浓度在25～600 mg/L范围内,均可促进种子萌发,最适宜浓度为400 mg/L。温度和稀土对异株荨麻种子发芽率无明显交互作用,而温度和赤霉素对促进种子发芽有显著的交互作用,各种组合中,以温度为30℃和35℃、赤霉素溶液浓度分别为400、500和600 mg/L的6个组合的交互效应对种子发芽率的提高最为显著。异株荨麻种子在中性条件下最易发芽,随着酸碱度的增加,种子发芽率逐渐降低;全光照和不同程度的遮荫(43%,58%,73%,87%,97%)对异株荨麻生长发育、光合特性以及荧光参数均有显著的影响,全光照到遮荫73%的光环境下异株荨麻均能正常生长发育,生育期快慢与遮荫程度呈负相关。遮荫程度小于73%条件下,随着遮荫程度的提高,异株荨麻株高、地茎、叶面积、叶片数、干物质积累量逐渐增多。而遮荫程度过高(超过73%)则不利于其生长和繁殖。异株荨麻的光补偿点和光饱和点均较低,净光合速率日变化曲线呈单峰型曲线。随着遮荫程度的提高,其光补偿点、光饱和点、暗呼吸速率、光合速率高峰值、日平均光合速率、蒸腾速率、气孔导度均依次降低。在不同光照条件下,蒸腾速率与气孔导度呈显著正相关。蒸腾速率和气孔导度与光合速率的相关性随遮荫条件的不同而异,全光照条件下蒸腾速率与光合速率呈显著正相关,而所有遮荫条件下相关性不显著。气孔导度与光合速率在所有光照下相关性均不显著。遮荫条件下叶片总叶绿素、叶绿素a、叶绿素b含量均随遮荫程度的提高呈上升趋势,而叶绿素a/b的值则随着遮荫程度的提高而下降。叶绿素荧光参数PSⅡ内禀光能转化效率(Fv/Fm)和潜在活性(Fv/Fo)日变化呈倒抛物线形。不同遮荫条件下Fv/Fm和Fv/Fo值均高于全光照的,且随着遮荫程度的提高其值均依次增加。说明,异株荨麻是一种耐荫性很强的植物。土壤水分含量对异株荨麻成活率、株高、地茎、叶面积以及体内水分分配等均有显著的影响作用。含量在85%条件下,异株荨麻株高、地茎生长量以及叶片数、单叶面积、单株叶面积以及干物质积累量均最高,但含水量过高(100%)或干旱胁迫处理(含水量低于60%)均不利于其生长。随着干旱胁迫程度的增加(土壤含水量下降),异株荨麻地上部含水量所占总水量的比值显著下降,而根部含水量所占总水量的比值显著上升,根比重和根冠比也显著上升。土壤含水量对异株荨麻光合特性也有显著的影响作用。龟水量为70%时,光补偿点、CO2补偿点、暗呼吸速率最低,而光饱和点、最大表观量子效率、最大净光合速率、RuBP羧化酶相对活性最高,随着土壤水分的增加或减少,光饱和点、最大表观量子效率、最大净光合速率和RuBP羧化酶相对活性显著下降,而光补偿点、暗呼吸速率和C02补偿点却呈上升趋势,且这种趋势随胁迫程度的加剧而愈加明显。不同水分条件下异株荨麻光合、蒸腾、气孔导度日变化均呈现单峰型曲线,其中土壤水分含量85%条件下异株荨麻光合速率、气孔导度、蒸腾速率最高,随着土壤水分含量的上升和下降,其值均显著降低。水分利用效率却随土壤水分降低呈增高趋势。土壤水分含量85%条件下,叶绿素a、叶绿素b、总叶绿素和类胡萝卜素含量均最高,随着土壤水分的上升和下降,叶绿体色素含量均呈显著下降趋势,但Chla/Chlb却随着土壤水分含量的下降一直呈现显著上升趋势。土壤含水量对异株荨麻活性氧清除酶(SOD、CAT和POD)、渗透调节物质(Pro、可溶蛋白和糖)以及膜透性和过氧化物(MDA)均有显著的影响作用。异株荨麻适宜在土壤水分含量60-100%(尤其85%)的生境中生长,在水分胁迫条件下异株荨麻通过改变体内代谢与合成的有关物质含量,增强对干旱的抵抗能力。比较异株荨麻在不同温度下的各项生长和光合指标,结果表明25℃下的地径和株高增长量最为显著,光合速率最大,生物产量最高,因此在20～25℃范围内,温度越高异株荨麻生长状况越好。对异株荨麻SOD、POD、MDA、Pro、可溶性糖、相对电导率的测定结果表明,异株荨麻20℃环境中生长时,受温度因子的抑制最小。不同在温度下异株荨麻各项生化指标存在显著相关性,表现为细胞膜透性、MDA、游离脯氨酸相互间为显著正相关,并与SOD活性、POD活性、可溶性糖含量间为显著负相关。采用正交试验设计,施用氮、磷、钾与微肥不同配比的肥料时,异株荨麻叶片中除粗脂肪外,其它营养物质如灰分、粗蛋白、维生素C、可溶性糖含量以及必需氨基酸和总氨基酸种类等均受到肥料显著的影响作用。综合平衡异株荨麻与施肥相关的5个品质指标,以各个指标达到最大值时各肥料不同水平出现的频率来表示,5个指标达到最优时的各种肥料配施组合为:尿素0.24g/kg土、过磷酸钙0.03g/kg土、硫酸钾0.05g/kg土、硫酸锰+硫酸硼0.0 1 7mg/kg土。采用GC-MS分析异株荨麻不同部位挥发油的活性物质,结果表明,不论是在叶部还是根部,异株荨麻生长期挥发油的化学成分都多于成熟期挥发油;异株荨麻叶片中的挥发油成分多于根部挥发油成分。而且不论是叶部还是根部,在从生长期到成熟期的生活过程中,挥发油中的大部分相同物质在总挥发油萃取物中所占的比例是增加的。挥发油中具有生物活性的物质很多,包括醇类物质、酮类物质、酯类物质、醛类物质、烯类物质、呋喃类物质、萘类物质、烷类物质和蒽类物质等。其中壬醛、2-甲基萘、石竹烯、α-法呢烯、苯甲酸己酯、甲氧基苯基肟、1-辛烯-3-醇即蘑菇醇、;3-辛酮、2-正戊基呋喃、壬醇、庚醛、十四醛、乙酸己酯、植醇、石竹烯、1-己醇、邻苯二甲酸双(2-甲基乙基)酯、丁香酚、1-庚醇、水杨酸甲酯等活性成分具有良好的工业、农业和医学价值。应用HPLC方法对不同生长期和不同器官的异株荨麻黄酮类化合物进行了定量分析。结果表明,异株荨麻地上部分黄酮含量要大于地下部分,成熟期的黄酮含量要大于生长期的黄酮含量,一年生,二年生,三年生根的黄酮含量随时间的变化逐年增高。无论是在不同的生长阶段还是不同的器官中,三种黄酮苷所占总黄酮的比例不同,槲皮素所占比例最大,其次是山萘酚,异鼠李素所占比例最低。应用模糊相似法分析异株荨麻在我国的适生区划,结果表明,最适宜引种区域为甘肃东南部的天水地区,其气候条件与原产地最为相似,是我国引种异株荨麻最为理想的地区。适宜引种区集中在我国北方北纬35～40°的中、东部地区,东北平原的吉林、辽宁两省以及青藏高原海拔3000～4000m的地区。次适宜引种区围绕适宜引种区形成外围圈,向北部延伸至黑龙江、内蒙古,向西达到甘肃、新疆,向东南到浙江、江西北部,向西南可达云贵高原。尚适宜引种地区主要在我国南部沿海,这些地区与原产地气候差异大,是我国广东、广西、海南、福建四省,江西、湖南、浙江南部部分地区。不适宜引种地区为我国新疆、内蒙部分地区。更多还原

【Abstract】 The stinging nettle, Urtica dioica L.subsp.dioica is an important nutritious and medical plant with great economic values. In this paper, systemic studies on configuration, development, seeds germination, and ecological adaptability in different environmental conditions of light, water and temperature stress of Utica dioica were researched, and the key factors and most appropriate cultivate conditions of Utica dioica introduction were illuminated subsequently. Extensive studies on the appropriate introduced area of Utica dioica in China by using fuzzy distance similitude priority ration on the basis of climatic data between the original and introduce area were also reviewed. Last, the nutritious and medical values of Utica dioica were discussed by way of studies on the relationship between fertilizer and nutrition, GC-MS analysis of volatile and extraction and identification of flavones substance. Results obtained from above are as follows:Structure of Utica doica roots stems and leaves dissection were observed with the paraffine slice. The period of fecundity of Utica dioica in Bejing is from may to September, and the growth period is from may to november, the optimum planting period is may, and seeds duration is 105 days, accumulate temperature above 10℃ of the fecundity duration is 1350 ℃, and the fecundity duration of multiply years seeds is about 75-80 days, accumulate temperature above 10℃ is 1050 ℃.The effects of temperature, lanthanon and temperature, gibberellin on seed germination to Urtica dioica L.subsp.dioica were studied respectively in laboratory using double factors randomized block design. Results indicated there were significant different germination rates at different temperatures range of 15 to 40 ℃. The optimum germination temperature of the seeds was 30 to 35 ℃. Both the lanthanon and gibberellin solution could improve the seeds germination at the concentration of 25～500 mg/L and 25～600 mg/L respectively, the optimum germination lanthanon and gibberellin solution concentration were 100 mg/L and 400mg/L respectively. The combined temperature and lanthanon have shown no interacting effect on seed germination while effect was significant between the combination of temperature and gibberellin, among all the combination of temperature and gibberellin, the 6 combinations of 30 and 35 ℃ combined with 400, 500 and 600 mg/L GA₃ have generated the best effect to seed germination. The seeds easily germinated at the litmusless while antiblastic at the acidic and basic water.The effects of full daylight and different shades （43% , 58%, 73%, 87% and 97%） on growth, photosynthetic characteristics and chlorophyll fluorescence parameters of Urtica dioica were examined. The results indicated that Urtica dioica L.subsp.dioica could grow and develop normally from the full daylight to 73% shade, the plant height, stem diamer, leaves areas and dry matter accumulation increase with increasing of shades degrees, while not benefit the plant growth exceeding the 73% shade. There were lower light compensation point （LCP） and light saturation point （LSP） and dark respiration rate （DR.）, which decrease with the increasing of shade degree. Diurnal variation of net photosynthetic rate （P_n） presents only one peak under full daylight and all shade Conditions, 0 the daily maximum and average net photosynthetic rate decrease significantly with increasing of shade degree, while the solar energy utilization efficiency increase. Diurnal variations of transpiration rate and stomatal conductance in leaves are similar to that of net photosynthetic rate. The values of transpiration rate and stomatal conductance also decrease as shade degrees increase. There are a positive correlation between transpiration rate and stomatal conductance under full daylight and all shade conditions while no significant positive correlation between the transpiration rate and net photosynthetic rate except that under the full daylight. Under full daylight and all shades there are no significant positive correlation between stomatal conductance and net photosynthetic rate. The content of total chlorophyll, chlorophyll a and chlorophyll b in leaves grown under all shades conditions are also significant higher than those under full daylight and increases with increasing of shade degrees while the chlorophyll a/b decreases. Diurnal variation of chlorophyll fluorescence parameters （Fv/Fm and Fv/Fo） presents the inverse parabola style, the value of Fv/Fm and Fv/Fo under all shade conditions are also significant higher than those under full daylight and increases with increasing of shade degrees. Results obtained from above showed that the stinging nettle was a shade tolerant plant by decreasing the light saturation point, light compensation point, dark respiration rate and chlorophyll a/b while increasing the content of total chlorophyll, chlorophyll a and chlorophyll b, and the value Fv/Fm and Fv/Fo under the shade conditions.The survival, height, stem and leaves growth and distribution of water content of Urtica dioica L.subsp.dioica were significantly different in different soil water stress. The water content of 85% is the most suitable for the accumulation of biomass. The photosynthetic characteristics were also affected significantly by different soil water stress. The lowest light compensation point and the highest light saturation point and apparent quantum efficiency occurred in 85% water content, while the light compensation point increase and the light saturation point and apparent quantum efficiency decreased with increasing and decreasing of the soil water content. The net photosynthetic rate and R.uBP activity were also the highest in the 85% soil water content, while the dark respiration rate and CO₂ compensation point were the lowest. With increasing and decreasing of the soil water content, the net photosynthetic rate and RuBP activity decreased and dark respiration rate and CO₂ compensation point increased. Diurnal variation of transpiration rate and stomatal conductance in leaves also presents only one peak in different soil water contents. The content of total chlorophyll, chlorophyll a and chlorophyll b in leaves grown in 85% soil water content were significant higher thaa others and increases with increasing and decreasing of soil water content, though the Chla/Chlb increased with decreasing of soil water content. The SOD activity, CAT activity, POD activity, Proline content, Dissoluble protein content, Dissoluble sugar content, MDA content, Conductance rate of Utica dioica in different water stress level were affected significantly. The Utica dioica is suitable to cultivate in moist soil environment.The temperature ranges from 20℃ to 25 ℃ is suitable for Urtica dioica L.subsp. dioica growth. the physiological and chemical index of Utica dioica were affected significantly under the temperature stress by determination of SOD and POD activity and MDA, Pro, Dissoluble sugar content and relative Conductance rate. The MDA content and relative Conductance rate increase significantly as the temperature increasing or decreasing from 20℃, while the SOD and POD activity decrease at high or low temperature. Combined all the growth and photosynthetic index at different temperatures, 25℃ is the most appropriate at which the plant, stem growth increment and photosynthetic rate and biomass reach maximum. There have significant relativity among all the physiological and chemical index of Utica dioica at different temperatures, with positive correlation among the amphithecium permeation and MDA and Pro content which negative correlate to SOD and POD actiyity and dissoluble sugar content.Nutrirnent of ash and crude protein and V_C and soluble sugar and amino acids of Urtica dioica L.subsp. dioica were significantly affected by the fertilizer of N and P and K and microelement by using the orthogonal design experiment. Different ration of fertilizers were also obtained when different single nutrition index reach the highest. Synthesis analysis to all the nutrition index indicated five nutriment index reach the highest when fertilized with 0.24g/kg carbamide and 0.03g/kg calcium phosphate and 0.05g/kg kalium sulphate and 0.017mg/kg manganese and boron sulphate.The chemical components of volatile oil extracted from different growth period and apparatus of Urtica dioica L.subsp.dioica were analyzed and identified by GC-MS. The results indicated that the chemical components identified during the growth were much more than those during the autumn either extracted from the leaves or roots, and the ratio of chemical components identified of the total contents increased from growth to autumn. The main constituents of volatile oils of Urtica dioica L.subsp.dioica includes mellow and ester and aldehyde and alkene and furan and naphthalene and alkyl and anthracene. Most of the chemical components have high value of industry and agriculture and medicine.Content of flavanone and flavanone glucoside in various growth period and apparatus of Urtica dioica L.subsp. dioica were analyzed using HPLC. The results indicated that Contents of flavanone in overground apparatus and in autumn were higher than those of underground and in growth period respectively. In the roots, Contents of flavanone increased as increasing of growth duration from 1 to 3 years. The ration of three flavanone glucoside including quercetin and karmpferol and isorgamnetin were different from different growth periond and apparatus of Urtica dioica L.subsp.dioica, among them, the content of quercetin was the largest, followed by karmpferol and isorgamnetin. These ressults offered useful values for the synthesis utilization of Urtica dioica L.subsp.dioica.The most suitable introduced area of Utica doica in China is the Tianshui area in southeastern Gansu province for its climate conditions are very close to the origin area using the dark similarity analysis. The most suitable introduced area of Urtica dioica L.subsp.dioica distributed at middle-east area of 35-40°N including Hebei, Beijing, Shandong, north of Henan, Shanxi, shanxi, north of Jiangsu, east of Gansu, Jiling, Liaoning province and Qinzang tableland. The suitable introduced area distributed at Henongjiang, Inner Mogolia, xingjiang, Lingxia, Jiangsu, Anhui, Hubei, Sichuan, Guizhou, Yunnan province and north of Zhejiang, Jiangxi and Hunan. The unsuited introduced area is the seaside of south China, including Guangdong,Guangxi, Hainan and Fujian province and parts of south of Jiangxi/Hunan and Zhejiang and Sinkiang and Inner Mongolia.更多还原