玫瑰(Rosa rugosa)根系生物学特性及对水肥空间异质性响应的研究
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摘要
本文以玫瑰(Rosa rugosa)野生类型和栽培品种为试材,研究了盆栽、根窖栽培和大田栽培玫瑰根系周年生长动态、根系生长与地上部的相关性以及野生玫瑰和栽培玫瑰根系生理生化代谢动态;探讨了营养元素、土壤紧实度、土壤质地、施肥量等多种因素对玫瑰生长、根构型与生理特性的影响以及玫瑰根系对水肥空间异质性的响应。结果如下:
     一、以玫瑰野生类型和栽培品种为试材,采用盆栽法、根钻法和改良根窖法等对玫瑰根系生长动态进行调查,对根系生长与地上部的相关性、根系的生理生化代谢等进行了研究,结果表明:
     1.玫瑰新根在一年中有3次生长高峰,高峰期出现的时间因栽培方式、植株年龄不同而有所变化。盆栽幼株的生长高峰分别出现在6月、7月和9月,大田栽植幼株的高峰出现时间比盆栽略有延迟。大田多年生玫瑰根系生长高峰分别出现在3月末、5月和9月。盆栽玫瑰根系发生数量明显多于大田玫瑰。大田多年生玫瑰野生类型和栽培品种根系生长高峰出现的时间基本一致。
     2.玫瑰根系生长与地上部生长既相互促进又相互矛盾,呈交替生长现象。新栽植株的根系大量生长之后出现新梢生长高峰,而新梢旺长又抑制新根的发生。大田多年生玫瑰第1次生长高峰是为了积累营养物质供生枝、展叶、开花所需,而地上部迅速生长又抑制根系生长;花后进入第2次根系生长高峰,随着果实的生长,根系生长量减少,果实成熟以后又迅速增加,出现根系生长高峰。
     3.在新根生长季节,玫瑰根系中可溶性糖和淀粉含量在春季形成1次高峰,然后总体呈下降趋势,生长后期随着根系生长而缓慢积累;游离氨基酸含量随地上部的快速生长而增加,春、夏季形成3次高峰,秋季随气温的下降而急剧上升;蛋白质含量变化相反;SOD和POD活性变化基本一致,春、夏季形成3次高峰,秋季随着地上部生长停止而呈上升趋势,但变化幅度不大。春夏季节野生玫瑰根系中淀粉、游离氨基酸、蛋白质含量和SOD、POD活性均明显高于栽培玫瑰,但秋季随着生长的停止而栽培玫瑰高于野生玫瑰;可溶性糖含量则野生玫瑰一直高于栽培玫瑰。不同时期碳氮物质含量和SOD、POD活性的不同也影响着野生玫瑰与栽培玫瑰的抗性以及进入休眠的时间。
     二、以玫瑰实生苗为试材,在沙培条件下研究氮磷钾营养亏缺对玫瑰幼苗根系总长度、表面积、体积、根尖数、分枝角度、分形维数等根构型参数与根系活力的影响。结果表明:氮磷钾营养亏缺条件下,玫瑰幼苗细根数量占总根量的比例减小;根系密度增大;根系平均直径变大;根系分枝角度整体增大,根系趋向水平分布;根系活力显著下降,氮磷钾营养亏缺时分别比对照下降19.08%、29.96%和55.59%;根系分形维数减小;其中缺氮或缺磷处理后,玫瑰幼苗侧根数量、长度、根系总长度、表面积、根系体积明显增大,缺钾与之相反,上述指标分别比对照下降了8.70%、3.95%、47.75%、56.03%和70.00%;缺钾比缺氮、缺磷时根系活力和分形维数下降得更多。
     三、采用盆栽方式,以盆栽玫瑰实生苗为试材,研究土壤紧实度对玫瑰幼苗根构型和氮代谢的影响。结果表明,在不同土壤紧实度条件下玫瑰幼苗生长指标及生物量顺序为:1.30g·cm~(–3)处理>1.50g·cm~(–3)处理>1.10g·cm~(–3)处理;根冠比随土壤容重的增加而下降,1.50g·cm~(–3)处理根冠比下降显著。土壤紧实度的增加抑制了地下部的生长,表现为根系长度、侧根数量、主根直径等均随土壤容重的增加而降低,根系平均直径和总面积有所增加;根系活力以1.30g·cm~(–3)处理的最高,1.50g·cm~(–3)处理的最低;根系中硝态氮、氨态氮含量和硝酸还原酶活性(NR)变化趋势一致,均是1.30g·cm~(–3)处理的最高,1.50g·cm~(–3)处理次之,1.10 g·cm~(–3)处理最低;谷氨酰胺合成酶活性(GS)随着容重的增加而升高。试验结果表明土壤紧实度显著影响玫瑰植株氮代谢活动,进而影响植株对养分的吸收和利用。容重1.30g·cm~(–3)的土壤紧实度较适宜于玫瑰生长。
     四、采用盆栽方式,对栽培于沙土、壤土、黏土中的玫瑰幼苗根构型与生理特性进行了研究,结果表明,壤土中生长的玫瑰幼苗侧根数最多,主根直径和根平均直径、根系总长度和表面积、地上部生长量均最大;沙土中生长的玫瑰侧根数最少,主根直径和根平均直径、根系总长度和表面积、地上部生长量最小;黏土中玫瑰地上部生长量、根构型参数介于二者之间,但根冠比明显小于沙土和壤土;沙土和壤土处理的玫瑰根活力明显高于黏土,且以壤土中最高;根系中氨态氮、硝态氮含量和硝酸还原酶的活性(NR)变化一致,均以壤土处理的最高,沙土处理的最低,谷氨酰胺合成酶活性(GS)表现为黏土>壤土>沙土处理。
     五、以玫瑰实生苗为试材,探讨了在盆栽条件下不同施肥量对玫瑰根构型及根系活力的影响。结果表明,玫瑰幼苗植株高度、茎粗、地上部鲜重随施肥量的增加而增加,但施肥量过大(9g/盆)则呈下降趋势;根冠比随施肥量增加而减小;主根直径,根系平均直径、根尖数、根系总长和表面积等根构型参数均随施肥量增加而逐渐增加,C处理(9g/盆)比B处理(6g/盆)减少。根系活力在施肥量增加时呈上升趋势,以B处理(6g/盆)的根系活力最大,适当施肥有利于玫瑰的生长发育,但施肥过量则对植株生长不利。
     六、利用基质培的试验方法,研究了温室条件下玫瑰幼苗在养分和水分空间异质条件下根构型、根系生物量的分布、地上部生长量、根冠比、细根直径及比根长等特征。结果表明,玫瑰根系对养分和水分空间异质性反应敏感,空间异质性对玫瑰根系生长和分布影响明显;水肥空间异质性处理的玫瑰幼苗地上部生长量及根系总量明显小于空间同质性,根冠比下降;在施肥区和浇水区根系生长快,生物量高,而在非施肥区和非浇水区根系生长受到抑制,生物量低;施肥区与非施肥区相比,细根直径减小,比根长增大,有利于养分和水分的吸收运输,而非浇水区,虽然细根直径下降,但是比根长小,获取水分的能力下降,生物量小,导致细根直径下降,不利于植株生长。水分空间异质性对玫瑰生长的影响大于养分空间异质性。
With wild varieties and cultivars of Rosa rugosa as experimental material, this thesis studied the root annual growth pattern in pots and open field, correlation between root growth and ground parts, root physiological and biochemical metabolic pattern of rose roots;and it worked over the growth, root architecture and physiological characters affected by nutrient elements, soil compaction, soil texture and fertilizer application amount, and responses to heterogeneous nutrient and water of Rosa rugosa roots. The results were as followed:
     1. We survey the growth pattern of rose roots in wild varieties and cultivars of Rosa rugosa by the way of potting, root drill and modified root cellar and study the correlation between the roots growth and the top growth, root physiological and biochemical metabolic pattern of rose roots. The results showed that:
     (1)There are three growth peaks of rose new roots each year, and the peak period depends on the cultivating ways and ages of plants. The growth peaks of young plants in pots are on June, July and September, but that of the open filed are slightly later. The root growth peaks of perennial plants in open field are at the end of March, May and September. The roots quantity of potted rose is obviously higher than that in open field. There were almost no difference in the period of root growing peaks between the perennial plants of wild varieties and cultivars.
     (2)The growth of roots and the new shoot of Rosa rugosa grow in both helping each other and mutually contradictory, showing the growth of alternating phenomenon. To the new cultured plants, after the roots growing quickly there is the new shoots growth peak, but the shoots growing quickly restrain the new roots grow in return. The first growth peak of the perennial rose plants in open field is for accumulating nutrient substance to meet the need of branching, leaves growing and flowering. In return the shoot growing quickly will restrain the roots growing. There is the second root growing peak after flowering, the quantities of roots reduce with the fruits growing, and grow speedily after fruits mature, and appear the new root growth peak.
     (3)The content of sugar and starch in rose roots in the growing season, would be in downward tendency after forming an absorption peak in spring. And then it would accumulate more with the slowing growth in later period. The content of amino acid was increased with the rapid growth of the aerial parts of rose roots. Meanwhile, three time peaks would be formed in spring and summer and the content of amino acid was increasing rapidly with the temperature dropping in autumn. However, the content of protein is opposite. The activities of SOD and POD were almost the same, which became three times of active peaks in spring and summer, and would be in a rising tendency with the growth of the aerial parts, but the changes were not so much. The content of starch, amino acid and protein, as well as the SOD, POD activities in roots of wild roses in spring and summer were significantly higher than that of rose cultivars, but that in autumn were reversed. The content of sugar in wild roses has been higher than that of rose cultivars. The content of carbon and nitrogen substances in different periods and the activities of SOD and POD also affect the resistance and the dormancy time of wild roses and rose cultivars.
     2. The sand culture system was used to study the effect of N, P, K-deficiency on the parameters of root architecture and activity of Rosa rugosa young seedlings. The results showed that proportion of tiny root decreased. The density and diameter of root were increased significantly, and the branches angles enlarged, and the roots tended to horizontal distribution, root activity obviously declined 19.08%、29.96% and 55.59% than that of control respectively, and the root fractal dimension diminished significantly under N, P, K-deficiency condition. Under N, P-deficiency condition, the length and number of lateral root, the total length, surface area and volume of root increased significantly. Under K-deficiency condition, on the contrary, the above specifications decreased 8.70%、3.95%、47.75%、56.03% and 70.00% by comparison.The root activity and fractal dimension under K -deficiency condition was less than under N, P -deficiency.
     3. The experiment was conducted with potted seedlings of Rosa rugosa in order to study the effect of soil compaction with different SBD(1.10 g·, 1.30 g·cm~(-3),1.50g·cm~(-3))on the root architecture and nitrogen metabolism of the seedlings. The result showed that the values of growth index and biological yield under different SBD were 1.30g·cm~(-3)>1.50g·cm-3>1.10g·cm~(-3).The ratio of R/S of plant was decreased with soil compaction increasing, and the R/S of the 1.50g·cm~(-3) SBD was significantly lower than that of the 1.10g·cm~(-3) and 1.30g·cm~(-3) SBD.The root growth of the potted seedlings were inhibited as the soil compaction was increased, including root length, lateral root number and tap root diameter, but the average diameter and surface area of root was increased. The root activity, NO_3~—-N and NH_4+-N contents, NR activity in root were the largest in the 1.30g·cm~(-3)SBD, then the 1.50g·cm~(-3) SBD and the 1.10g·cm~(-3) was the lowest.GS activity in root was increased as the soil compaction increasing. The experiment indicated that the soil compaction was a crucial factor to affect nitrogen metabolism and influence the absorption and utilization of nutrients in plants. And 1.30g·cm~(-3) SBD was favorable for plants to grow.
     4. A pot experiment was carried out to explore the characteristics of growth, root architecture and physiological characters of seedling of Rosa rugosa planted in sandy, loam and clay soil. The results showed that, the top growth quantity of in loam was the highest, the diameter of main root and average was the biggest, quantity of lateral root was the most, total length of root and surface area was the biggest; all these of them in sand were the lowest, and these of them in clay were the moderate, but the R/S of Rosa rugosa in clay was lower distinctly than in sand and loam. the root activity of Rosa rugosa in sand and loam were higher than that in clay, the root activity of Rosa rugosa in loam was the highest; the content of NH_4~+-N and NO3--N of roots, NR activity in root was almost the same, all of them were the highest in loam and lowest in sand; GS activity in root was the largest in clay, then in loam and the lowest in sand.
     5. The seedlings of Rosa rugosa were used to study the effect of different fertilizer application amount on the growth, root architecture and activity under pot culture condition. The results showed that the plant height, term diameter, top fresh weight increased with the increasing fertilizer application amount, but above three indices decreased when the fertilizer application amount was higher(9g/pot);the R/S decreased with the increasing fertilizer application amount. The diameter of tap root, average diameter, number, total root length, root surface area and root activity increased with the increasing fertilizer application amount, reached the highest level supplied with 6g/pot.Rational fertilizer application amount was conducive to growth and development of Rosa rugosa.
     6. The medium culture was used to study the effect of heterogeneous nutrient and water on root architecture, root mass distribution, top mass, R/S, fine root diameter and SRL in greenhouse. The result showed that the root of Rosa rugosa was responded sensitively to water and nutrient heterogeneity, growth and distribution of root were affected obviously by space heterogeneity; growth quantity of up ground parts, total root and R/S treated in water and fertilizer heterogeneity were less than in homogeneity, and the R/S was reduced; roots grew quickly and total growth quantity was added clearly in fertilizer and irrigated area, and the growth were restrained in no nutrient and water area. Compared with the no fertilizer area, the diameter of fine root decreased and increased SRL were beneficial for absorbing and transporting of nutrient and water, in the no irrigation area, though diameter of fine root decreased, but the low SRL reduced the ability of absorbing water, reduced the total quantity of growth and then were bad to rose growing. Rose growing affected by water heterogeneity was more obviously than that by fertilizer heterogeneity.
引文
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