设施栽培仙客来养分吸收规律及钙影响优质花形成机理的研究
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摘要
仙客来(cyclamen persicum L.)也称兔耳花、一品冠等，为报春花科仙客来属球茎花卉，它是利用C.persicum所特有的遗传变异性，经过一代又一代仔细的选择和种内杂交培育而形成的一类花卉，该花卉花形独特、花色丰富、花期长。在欧美、日本等发达国家已经形成与各品种配套的产业化生产技术。我国仙客来的栽培历史短，产业化栽培则刚起步，国外引进的品种在我国的栽培条件下缺乏相应的技术研究，导致种子、栽培介质、肥料品种等都未在引进的基础上实现本土化，商品率低、生产成本高、缺乏优质仙客来的管理技术等问题普遍存在。为了促进我国仙客来产业的可持续发展，本研究在总结国内外仙客来产业化生产的技术现状，着重研究了设施栽培条件下仙客来的养分吸收规律、不同施肥模式对仙客来生长发育和品质的影响和钙对花器官发育及优质花形成的影响，主要研究结果总结如下：
一、设施栽培仙客来生长发育特点和养分吸收规律研究
针对仙客来工厂化生产中存在的盲目施肥、肥料利用率低、生产成本高、生长发育受阻、产量和品质难以提高等问题，研究了我国仙客来主栽品种的生长发育特点和养分吸收规律，获得以下主要结果：
1．通过对我国3个仙客来主栽品种(S3、M1、NP3)的生长规律研究，结果表明：在供试条件下，3个品种仙客来的生长发育由于受高温胁迫的影响，在越夏期即(8月21日)以前生长缓慢；叶片数仅占总数的30～40％，干物重仅占到总数的9.8％～15.5％。越夏后，仙客来迅速恢复生长，出叶加快，叶片数迅速增加，干物质积累加快，但恢复能力品种间有差异，其中M1＞NP3＞S3，进入盛花期M1最早，S3最迟，NP3处于中间；达到商品期时(1月12日)最大叶片数S3为83，M1为84，NP3为81，都超过了精品仙客来叶片的要求；干物质积累最大值S3为49.9g plant~(-1)，M1为43.2g plant~(-1)，NP3为42.2g plant~(-1)；总花数S3为23，M1为31，NP3为29。
2．经仙客来3个主栽品种的全生育期养分需要量的测定，结果表明：在设施栽培条件下仙客来氮磷钾的需要量S3分别为1257.25mg plant~(-1)、218.79mgplant~(-1)、1171.69mg plant~(-1)，需要比例为5.7：1：5.4，氮磷钾总量为2647.73 mg plant~(-1)；M1分别为1214.14mg plant~(-1)、222.13mg plant~(-1)、1292.66mg plant~(-1)，需要比例为5.5：1：5.8，氮磷钾总量为2728.93 mg plant~(-1)；NP3分别为1003.78mg plant~(-1)、174.23mg plant~(-1)、992.82mg plant~(-1)，需要比例为5.8：1：5.7，氮磷钾总量为2170.83mg plant~(-1)；氮磷钾需要量，M1＞S3＞NP3。仙客来对钙、镁的需要量S3分别为184.50 mg plant~(-1)、85.40mg plant~(-1)；M1分别为203.51 mg plant~(-1)、97.05mg plant~(-1)；NP3分别为273.21 mg·plant~(-1)、127.14mg plant~(-1)；钙镁的需要量NP3＞M1＞S3。仙客来对微量元素的需要量各品种也有一定的差别，从总量看，M1的需要量大于S3和NP3。
3．考察仙客来各生育期养分吸收规律，可以看出：在试验条件下，仙客来在越夏前对氮磷钾的吸收量较少，占最大吸收量氮为5.3～7.9％，磷为7.0～14.4％，钾4.8～7.7％。但品种间有差异，其中M1和NP3的前期氮磷钾吸收积累量要大于S3；越夏期间品种S3对氮磷钾的吸收增加较少，而M1和NP3有缓慢增加；越夏后仙客来对氮磷钾的吸收迅速增加，吸收与积累的规律与仙客来植株的生长发育规律相一致。仙客来对钙、镁及微量元素的吸收动态与氮磷钾的吸收有相似的变化。
二、不同施肥模式对仙客来生长发育和养分吸收利用的研究
根据设施栽培仙客来的养分吸收特点以及优质仙客来生产要求，设置了三种不同的供肥模式，在三个主栽品种上试验，取得了如下结果：
1．根据仙客来的养分吸收规律及供试条件的生长发育状况设置的供肥模式2和模式3与常规供肥模式1相比，可以明显降低越夏期间仙客来的高温逆境胁迫，越夏后恢复生长快，增叶早而多，干物质积累迅速增加，花芽分化多，开花早而整齐，叶下花少，优质花增加。其中，模式3优于模式2；从3个品种看，M1、NP3品种的影响效果优于S3。
2．分析不同供肥模式对仙客来养分吸收的影响，可以发现：不同供肥模式对氮素吸收影响品种间比较一致，模式1＞模式3＞模式2，模式1吸收了过多的氮而不利于仙客来发育。对磷素的影响品种间不一致：品种S3是模式2＞模式3＞模式1，NP3是模式1＞模式3＞模式2，而M1在3种模式下的磷素吸收量差异不大，其中NP3品种在模式1下对磷的过多吸收影响了仙客来的生长发育及开花质量。对钾、钙、镁的影响都是模式2和模式3高于模式1。对微量元素的吸收的影响，模式间和品种间都有差别，但总体来说，供肥模式2更有利于微量元素的吸收。
3．计算不同供肥模式下仙客来养分利用率，可以明显看出：模式2和模式3与模式1相比显著提高了氮磷钾养分的利用率，其中氮利用率提高了1.9～2.5倍，磷利用率提高了3.5～4.6倍，钾利用率提高了，2.2～3.3倍。但不同供肥模式对养分利用率的影响品种间有一定的差异。
4．在中国东南部设施栽培条件下，高温逆境胁迫对仙客来生长发育带来深刻影响，从而导致产量降低，品质差，是优质仙客来生产的重要限制因素。如何弄清仙客来的抗逆境生理，提高仙客来抗高温逆境胁迫的能力，是仙客来生产者十分关心的问题。通过对高温胁迫下仙客来根系活力，叶片中SOD酶活性、MDA含量等有关细胞膜稳定性，可溶性糖含量等生理生化指标的研究，结果表明：模式2和模式3供肥条件下仙客来抗高温胁迫能力大于常规模式1。结合越夏期间仙客来叶片、花芽的生长状况，以及越夏后迅速恢复生长，提前进入旺长期，形成大株型优质花商品的实践也证明了这一点。
综上所述并结合生产实际，对于S3品种由于在供肥模式2和模式3的条件下，生长发育差异不大，而模式2施肥简单，利于操作，在生产中可以选择模式2施肥；NP3和M1品种在模式3下生长显著优于模式2，不仅越夏后叶片生长和花芽分化迅速，且提早上市，品质优，生产中可以选择模式3施肥。
三、钙营养对仙客来花器官发育及优质花形成机理的研究
1．水培条件下，仙客来对钙的吸收与花器官发育关系密切。当营养液中钙浓度分别为0mmol L~(-1)、2.5 mmol L~(-1)和5 mmol L~(-1)时，仙客来对钙的吸收速率分别稳定在-0.6 mg plant~(-1) d~(-1)、5 mg plant~(-1) d~(-1)和15 mg plant~(-1) d~(-1)，花器官中钙含量分别对应为3.372g kg~(-1)d．wt、6.341 g kg~(-1)d．wt和8.697 g kg~(-1)d．wt，而它们相对应的平均开花数量分别为3、8和12，平均花茎高度分别20.2cm、24.7cm和28.3cm。因此，在一定浓度范围内增加营养液中钙的浓度可以提高开花品质。提高营养液中氮的浓度，仙客来对钙的吸收受到显著抑制，钙的平均吸收速率仅为2.5 mgplant~(-1) d~(-1)，花器官中钙含量为3.784 g kg~(-1)d．wt，其花茎高度甚至比缺钙处理还低19.8％。增加营养液中硼的浓度对仙客来钙吸收影响不显著，但促进了仙客来开花数量，同时花器官中钙和硼的比例与5mmol／L钙处理接近。
2．SEM扫描结果表明，花器官中钙含量显著较低的无钙处理和高氮处理其花茎维管组织发育不成熟，维管组织小，细胞排列凌乱，髓射线、木质部和韧皮部无明显区分；而其他花器官中钙含量较高的处理则花茎维管组织内细胞清晰可见，排列规则，髓射线、木质部和韧皮部各部分有明显区分界限，能见到具环形加厚现象的导管分子，其中以5mmol L~(-1)钙处理最明显。因此，钙对花茎组织细胞的发育的影响是其影响花器官发育的原因之一。
3．TEM超微结构表明，花器官中钙浓度显著较低的无钙处理和高氮处理下，花茎细胞叶绿体、线粒体、细胞核等细胞器明显受损。缺钙处理下细胞壁厚度非常不均匀，出现断裂；高氮处理下虽未见细胞壁、细胞膜断裂现象但细胞壁很薄。其它花器官中钙浓度较高的处理下细胞结构正常，细胞器较完整，未见破损现象的发生，随着钙浓度的增加5mmol L~(-1)钙处理下细胞结构较2.5mmol L~(-1)钙处理更完整清晰。
4．钙的细胞化学定位结果表明，花茎细胞中细胞壁和细胞间隙是Ca~(2+)分布的主要场所，细胞膜、核膜及细胞质中也有少量分布。5mmol L~(-1)钙处理下，Ca~(2+)沉淀颗粒最多；而缺钙处理和高氮处理下Ca~(2+)沉淀颗粒很少，几乎没有；增加硼的供应细胞间隙中Ca~(2+)颗粒分布较少，而细胞壁上Ca~(2+)颗粒增多且体积有所增大。对花茎细胞切片进行EDS能谱分析，测定不同钙条件下花茎细胞中钙的相对含量，测定结果与钙的细胞化学定位结果较一致，5mmol L~(-1)处理下花茎细胞中高密度电子体的Ca／O比值最大，花茎细胞钙含量最大。各处理Ca／O比值大小顺序为Ca_0＜+Ca+N＜+Ca≤+Ca+B＜++Ca。
5．Ca~(2+)吸收运输与Ca~(2+)-ATPase活性关系密切。在缺钙处理下，仙客来各器官钙含量显著降低，细胞内钙浓度也减少，各器官Ca~(2+)-ATPase活性也显著降低，可能导致Ca~(2+)信使系统的紊乱。增加外源钙浓度促进各器官中钙含量，也显著提高Ca~(2+)-ATPase活性；增加氮的供应抑制了钙的吸收，Ca~(2+)-ATPase活性也有所下降。
6．分析测定了不同供钙条件下仙客来花茎细胞壁各成分含量的变化，结果表明，随着花器官中钙含量的增加，纤维素含量显著增加，钙含量较低的缺钙处理和高氮处理下细胞壁中纤维素含量是钙含量较高处理的三分之一左右；而果胶含量和半纤维素含量变化随着钙浓度增加而降低；增加氮的供应使细胞壁中半纤维素所占的比重达到了65.6％。随着花器官中钙含量的增加，醛酸在果胶中含量也随着增加，纤维素中呈递减，半纤维素中变化不一；总糖含量的变化与醛酸含量变化规律较相似；蛋白质在半纤维素中含量增加，纤维素中降低，果胶中变化不大。因此，供钙对仙客来花器官发育的影响与细胞壁结构的变化直接相关。
7．不同供钙条件能显著影响仙客来花器官内源激素的含量。钙吸收正常的处理，随着花茎伸长至开花，IAA、ZTs和GA_3含量都增加。缺钙和钙吸收受阻处理中，赤霉素类物质在花茎伸长到盛花期间含量急剧下降，可能导致花茎伸长受阻；缺钙处理IAA含量水平变化与钙供应充足的处理呈相反趋势，也可能导致花茎弯曲、畸形现象；同时，花茎发育后期生长素类物质含量小可能是其维管组织发育不完善的主要因素。高浓度外源钙的处理使花器官中不仅IAA和ZTs含量都高于其他处理，而该处理导管增厚最显著，两者的对应关系密切。
8．设施栽培条件下研究了不同的供钙方式和供以不同钙源对仙客来开花的影响，结果表明，停止外源钙的供应对仙客来开花品质影响较大，花茎高度仅高出叶面2cm左右，达不到仙客来商品要求。CaEDTA在促进仙客来钙吸收上比Ca(NO_3)_2更有效，相对应在促进仙客来开花品质上也更有效。不同的供钙方式下仙客来营养器官钙含量差异较大；根部供钙下钙含量为根＞球茎＞叶片，叶面喷钙下钙含量叶片＞根＞球茎；但花器官中钙含量并没有因为供钙方式差异表现出显著差异。
Cyclamen (cyclamen persicum L.), also called Tu'er Flower, Yipin Guan, etc, belongs to corm flower of Cyclamen genus, Primula malacoides Franch family. By virtue of peculiar hereditary variability of C. persicum, it is a kind of flower fostered through selection generation after generation and intraspecific hybridization and cultivation. This flower has special flower forms, abundant flower color, and long florescence. In developed countries such as America, Europe, Japan, etc., the related industrialization production technology of different varieties has been formed. In our country, the planting history of the cyclamen is short and the industrialized culture just starts. Because of lack of corresponding technical research, the breeds introduced abroad cannot realize the localization in the aspects of seed, culturemedium, and fertilizer. The problems such as low commodity rate, high production cost and lack of management skills of the high-quality cyclamen generally exist. In order to accelerate the sustainable development of cyclamen industry of our country, this research, based on the technological status quo of domestic and international industrialization production of cyclamen, investigates the nutrient absorption rule of cyclamen under the cultivation conditions, different fertilizer mode impact on the growth and development and quality, and calcium influence on the development of flowers and the formation of high-quality flowers. The main results are summarized as follows:
Ⅰ. Growth and development characteristics and nutrient absorption rules of Cyclamen cultivated under facility conditions
Aiming at the problems of blind fertilizer application, low fertilizer utilization ratio, high production cost, obstruction of growth and development, difficulty of the improvement of output and quality, etc., the growth and development characteristics and nutrient absorption rules of the main cultivated cyclamen breeds in our country have been studied and the following results are obtained:
1. The study on the growth rules of three cultivars (S3, M1, NP3) of cyclamen of our country shows that, under the testing conditions, the growth and development of three cultivar breeds are slow before the over-summer period (August 21) due to the influence of high temperature stress; The number of leaves accounts for 30～40% of the total number, and the dry matter weight only accounts for 9.8% - 15.5% of the total number. However, after the over-summer period, cyclamen resumes to grow rapidly, leaf emergence accelerates, the number of leaves rapidly adds, and the accumulation of dry matter accelerates, but there are differences on the repair capacity among the breeds, wherein M1＞NP3＞S3, and M1 firstly starts into the full florescence, S3 is the last and NP3 is in the middle; when reaching the period of commodity (January 12), the maximum number of leaf for S3 is 83, for M1 is 84, and for NP3 is 81, all of which exceed the requirements of leaves of choice cyclamen; the maximum of dry mater accumulation for S3 is 49.9g·plant~(-1), for M1 is 43.2 g·plant~(-1), and for NP3 is 42.2 g·plant~(-1); and the total number of flowers for S3 is 23, for M1 is 31, for NP3 is 29.
2. The determination of the nutrient requirements in the whole breeding time of the three cultivars of cyclamen shows that, under the cultivation conditions, the requirements of nitrogen, phosphorus and potassium for S3 are respectively 1257.25mg·plant~(-1), 218.79mg·plant~(-1), 1171.69mg·plant~(-1), the ratio between them is 5.7:1:5.4, and the total amount of the three elements is 2647.73 mg·plant~(-1); the requirements of nitrogen, phosphorus and potassium for M1 are respectively 1214.14mg·plant~(-1), 222.13mg·plant~(-1), 1292.66mg·plant~(-1), the ratio between them is 5.8:1:5.7, and the total amount of the three elements is 2170.831 mg·plant~(-1); while the requirements of nitrogen, phosphorus and potassium for NP3 are respectively 1003.78mg·plant~(-1), 174.231mg·plant~(-1), 992.82mg·plant~(-1), the ratio between them is 5.5:1:5.8, and the total amount of the three elements is 2728.93 mg·plant~(-1); comparing the required amount of the three elements for the three breeds, the result is M1＞S3＞NP3. The requirements of calcium, magnesium for S3 are respectively 184.50 mg·plant~(-1), 85:40mg·plant~(-1); for M1 are respectively. 203.51 mg·plant~(-1), 97.05mg·plant~(-1); while for NP3 are respectively 273.21 mg·plant~(-1), 127.14mg·plant~(-1); comparing the total amount of calcium and magnesium for the three breeds, the result is NP3＞M1＞S3. The requirements of microelement for cyclamen are also different between different breeds. For the total amount, M1 is greater than S3, NP3.
3. Investigation on the nutrient, absorption rules of cyclamen in different cultivation period shows that, under the testing conditions, the absorption amount of nitrogen, phosphorus and potassium before over-summer period is relatively less, accounting for 5.3～7.9%, 7.0～14.4%, 4.8～7.7% of the total absorption amount of nitrogen phosphorus and potassium respectively. But there are differences among the varieties. The absorption accumulation amount of M1 and NP3 is greater than S3 during the early period; the absorption accumulation rate is relatively less for S3 and slowly increases for M1 and NP3; the absorption of nitrogen, phosphorus and potassium of cyclamen greatly increases after over-summer period, and the rules of absorption and accumulation keeps pace with the growth and development of cyclamen. The absorption trends of calcium, magnesium and microelements are similar to the absorption of nitrogen, phosphorus and potassium.
Ⅱ. Research on the growth and nutrient absorption and utilization under different fertilization modes
Three different fertilization modes are established based on the nutrient absorption characteristics and production requirements of cyclamen cultivated in the different facilities. The following results have been obtained on the three different breeds:
1. Based on the nutrient absorption rules and growth of testing conditions, three fertilizer supply modes are established. Compared with fertilizer supply model, Mode 2 and mode 3 can obviously decrease the high temperature stress during the over-summer period, resume the growth more quickly, and increase leaves earlier and more; accumulates and increases dry matters rapidly, splits up more bud, blooms early and neatly with less flowers below leaves, and quality flower increases. Among them, mode 3 is superior to mode 2; for the three varieties, the influence effects of M1, NP3 are superior to S3.
2. Analysis of different fertilization supply modes effects on the nutrient absorption shows that, different fertilizer supply modes have similar effects on the nitrogen absorptions; that the impacts of mode 1 is greater than mode 3, and mode 3 is greater than mode 2. For mode 1, too much nitrogen absorption is not good for the development of cyclamen. However, the effects on element of phosphorus are inconsistent among different varieties: for S3, mode 2＞mode 3＞mode 1, for NP3, mode 1＞mode 3＞mode 2, but for M1, there is no obvious difference on the phosphorus absorption under the three modes. Among them, for NP3, the surplus absorption in phosphorus under mode 1 influences the development and growth and bloom quality of cyclamen. As to the impacts on potassium, calcium, magnesium, mode 2 and mode 3 are higher than mode 1. For the impact on absorption of the microelements, there are differences among the modes and among the breeds. But as a whole, the fertilizer supply mode 2 is more favorable to the absorption of microelements.
3. Calculation on the nutrient utilization ratio under different fertilizer supply modes obviously shows that, compared with mode 1, mode 2 and mode 3 have obviously improved the utilization ratio of the nitrogen phosphorus and potassium nutrients, among them the utilization ratio of nitrogen has been increased by 1.9～2.5 times, utilization ratio of phosphorus by 3.5～4.6 times, while utilization ratio of potassium has been increased by 2.2～3.3 times. But for the impact on nutrient utilization ratio of different fertile modes, there are some differences among the breeds.
4. Under the facilities culture conditions in the southeast of China, high-temperature stress makes a great influence on the growth of development of cyclamen, which leads to the reduction of output and bad quality. It is an important restricting factor for the production of high-quality cyclamen. How to clarify the stress physiology and improve the capacity of resisting high-temperatures stress is the question that cyclamen producers care about very much. Through the study on biochemical and physiological index on relevant cell membrane steady and soluble sugar contents of root activity, SOD enzyme activity of the leaf, MDA content, it shows that, the capability of resisting high-temperature stress of mode 2 and mode 3 is greater than that of the regular mode 1, which can be also demonstrated by the fact of the characteristics of leaf and bud growth conditions of cyclamen during the over-summer period, the rapid growth and quick stepping into the prosperous growth period in advance after over-summer period, and forming of large high-quality flowers.
Considering the above descriptions and the production practice, for S3, there is no obvious difference in growth and development under the fertilizer supply between mode 2 and mode 3, and it is simple for fertilizing and application in mode 2, so mode 2 can be chosen for fertilizing in the production; the growth conditions of NP3, and M1 are obviously better than mode 2. Not only the leaf growth and bud splitting up is quickly after over-summer period, but also the products can come into the markets in advance with excellent quality, so mode 3 can be chosen for fertilizing in the production.
Ⅲ. Research on the mechanism of floral organs development and high-quality flower formation influenced by calcium nutrition
1. Under the water culture condition, the absorption of calcium is closely related to the floral organs development. When the concentration of calcium is Ommol L~(-1), 2.5 mmol L~(-1) and 5 mmol L~(-1) respectively in nutrient solution, the absorption rates of calcium are steadily around -0.6mg plant~(-1) d~(-1), 5mg plant~(-1) d~(-1) and 15 mg plant~(-1) d~(-1) respectively; the corresponding contents of calcium in the floral organs are respectively 3.372g kg~(-1)d.wt, 6.341 g kg~(-1) d.wt and 8.697 g kg~(-1) d.wt; the corresponding bloom quantity are 3, 8 and 12 respectively; the average height of pedicel is 20.2cm, 24.7cm and 28.3cm respectively. Therefore, the increase of calcium concentration within a certain scope can improve the quality of blooming. The improvement of nitrogen concentration will obviously inhibit the absorption of calcium. The average absorption rate of calcium is 2.5 mg plant~(-1) d~(-1), the content of calcium in floral organs is 3.784 g kg~(-1)d.wt, and the height of pedicel is even 19.8% lower than that under the condition of calcium deficiency. The increase of boron concentration in the nutrient solution has no obvious influence on the absorption of calcium, but it accelerates the quantity of blooming, and the ratio between calcium and boron in the floral organs is close to the results of the group treated with 5mmol L~(-1) calcium.
2. The result of SEM scan shows that under the acalcerosis treatment and high nitrogen treatment for the floral organs containing relatively lower calcium, the vascular tissue of scape is small with irregular cell arrangement and indistinct difference of medullary ray, xylem and phloem. However, under the treatment for other flower organs containing higher calcium, the cell in the vascular tissue of scape which is regularly arranged, is clear, and there is distinct difference in medullary ray, xylem and phloem. Moreover, annular and thickening vessel element can be observed, of which the treatment for 5mmol L~(-1) calcium is the most obvious. Therefore, one of the reasons that calcium has influence on the growth of flower organs is that it has influence on the growth of the scape cell.
3. TEM ultrastructure shows that under the acalcerosis treatment and high nitrogen treatment, the chloroplast, chondriosome and cell nucleus of the scape cell will be damaged obviously. Under acalcerosis treatment, the thickness of cell wall is exceptionally inhomogeneous and broken, but under high nitrogen treatment, although there is no breakage of cell wall and cell membrane, the cell wall is very thin. Under the treatment of other flower organs with relatively high calcium concentration, the cell structure is normal, and the organelle is complete without any breakage. As the increase by 5mmol L~(-1) of calcium concentration, the cell structure becomes clearer and more complete compared with the treatment group of 2.5mmol L~(-1) calcium.
4. The results for the cytochemistry allocation of calcium shows that, the cell wall and intercellular space in the scape cell are the main distribution places of Ca~(2+), which are also distributing in the cell membrane, nuclear membrane and cytoplasm with a little amount. Under 5mmol L~(-1) calcium treatment, there will be maximum amount of Ca~(2+) precipitation grain, and there will be little Ca~(2+) precipitation grain under the acalcerosis treatment and high nitrogen treatment. The distribution amount of Ca~(2+) precipitation grain in the intercellular space of the new added boron is relatively small, while the amount and volume of Ca~(2+) grain on the cell wall increases. Take a section of the scape cell for EDS energy spectrum analysis, and measure the comparative calcium content of scape cell under different calcium conditions. The measurement result is comparatively consistent with the result of cytochemistry allocation of calcium. Under 5mmol L~(-1) calcium treatment, the Ca/O ratio of high density electronic body in the scape cell is relatively high. The order of the Ca/O ratio under all kinds of treatment should be Ca_0＜+Ca+N＜+Ca＜+Ca+B＜++Ca.
5. Absorption and transportation of Ca~(2+) are closely related to the activity of Ca~(2+)-ATPase. Under Calcium-deficiency treatment, the calcium contents in all organs of cyclamen decrease obviously, calcium concentration in cells also reduces and activities of Ca~(2+) -ATPase in all organs also markedly decease. This may cause disorder of Ca~(2+) messenger system. Increase of calcium concentration from external source can improve calcium contents in all organs and obviously promote the activity of Ca~(2+) -ATPase. Increase of nitrogen supply inhibits the absorption of calcium and also decreases the activity of Ca~(2+) -ATPase.
6. The changes of contents of all components in cell wall of cyclamen under conditions with different calcium supply are analyze and measured. The result indicates that with the increase of calcium content in organs, the cellulose content obviously increases. The cellulose content in cell wall under calcium-deficiency and nitrogen-efficiency is about 1/3 of that under treatment with comparatively high calcium content. However, the contents of pectin and hemicellulose decrease with the increase of calcium concentration. The increase of nitrogen supply makes the proportion of hemicellulose reach 65.6%, With the increase of calcium content in flower organs, the content of aldehydic acid increases in pectin, decreases, in cellulose and changes without regular rules in hemicellulose. The rule of change of total sugar content is similar to that of change of aldehydic acid content. The content of protein increases in hemicellulose, decreases in cellulose and changes a little in pectin. Therefore, calcium supply has direct effect on growth of organs of cyclamen and changes of structure of cell wall.
7. The different calcium condition can obviously affect the cyclamen flower organ endogene hormone content. At the calcium absorption normal treatment, IAA, ZTs and GA3 content are increasing along with the floral elongates to blossoms. At the treatments with lacking the calcium and the calcium absorption blocked, the gibberelin class matter content drops suddenly during the period of floral elongates to blossoms. It may cause the pedicel elongation to be blocked. The change of the IAA content level was opposite within the lacking calcium treatment and the calcium supply sufficient treatment. It also possibly causes the abnormal phenomenon of pedicel curving etc. At the same time, the lower content of growth hormone class matter is possibly the imperfect primary factor of vascular tissue growth during later period of pedicel growth. At the treatment with high calcium content supplying, both the IAA and the ZTs content are higher and the incrassation phenomenon of vessel is obviously. Maybe, there are some corresponding relations.
8. Under different facility cultivation condition, studies on the influence of difficult calcium-supplying methods and different calcium resources on cyclamen-blooming show that, stopping the external supply of calcium has greater impact on the quality of cyclamen-blooming, with the height only about 2cm higher than the leave which is beyond the need of the cyclamen goods. CaEDTA is more effective than Ca(NO_3)_2 to promote cyclamen's absorption of calcium, as well as to improve cyclamen-blooming quality. Due to different calcium-supplying methods, tremendous differences exist in calcium content of nutrition organ of the cyclamen, for example, as to calcium supplied by root method, calcium content of root is higher than corm, which is higher than leaves. However it turns out to be different for calcium sprayed on leaves method, with which the calcium content of leaves is higher than root, and root higher than corm. Nevertheless, there is no obvious difference in calcium content in its floral organs resulted from different calcium-supplying methods.
一、设施栽培仙客来生长发育特点和养分吸收规律研究
针对仙客来工厂化生产中存在的盲目施肥、肥料利用率低、生产成本高、生长发育受阻、产量和品质难以提高等问题，研究了我国仙客来主栽品种的生长发育特点和养分吸收规律，获得以下主要结果：
1．通过对我国3个仙客来主栽品种(S3、M1、NP3)的生长规律研究，结果表明：在供试条件下，3个品种仙客来的生长发育由于受高温胁迫的影响，在越夏期即(8月21日)以前生长缓慢；叶片数仅占总数的30～40％，干物重仅占到总数的9.8％～15.5％。越夏后，仙客来迅速恢复生长，出叶加快，叶片数迅速增加，干物质积累加快，但恢复能力品种间有差异，其中M1＞NP3＞S3，进入盛花期M1最早，S3最迟，NP3处于中间；达到商品期时(1月12日)最大叶片数S3为83，M1为84，NP3为81，都超过了精品仙客来叶片的要求；干物质积累最大值S3为49.9g plant~(-1)，M1为43.2g plant~(-1)，NP3为42.2g plant~(-1)；总花数S3为23，M1为31，NP3为29。
2．经仙客来3个主栽品种的全生育期养分需要量的测定，结果表明：在设施栽培条件下仙客来氮磷钾的需要量S3分别为1257.25mg plant~(-1)、218.79mgplant~(-1)、1171.69mg plant~(-1)，需要比例为5.7：1：5.4，氮磷钾总量为2647.73 mg plant~(-1)；M1分别为1214.14mg plant~(-1)、222.13mg plant~(-1)、1292.66mg plant~(-1)，需要比例为5.5：1：5.8，氮磷钾总量为2728.93 mg plant~(-1)；NP3分别为1003.78mg plant~(-1)、174.23mg plant~(-1)、992.82mg plant~(-1)，需要比例为5.8：1：5.7，氮磷钾总量为2170.83mg plant~(-1)；氮磷钾需要量，M1＞S3＞NP3。仙客来对钙、镁的需要量S3分别为184.50 mg plant~(-1)、85.40mg plant~(-1)；M1分别为203.51 mg plant~(-1)、97.05mg plant~(-1)；NP3分别为273.21 mg·plant~(-1)、127.14mg plant~(-1)；钙镁的需要量NP3＞M1＞S3。仙客来对微量元素的需要量各品种也有一定的差别，从总量看，M1的需要量大于S3和NP3。
3．考察仙客来各生育期养分吸收规律，可以看出：在试验条件下，仙客来在越夏前对氮磷钾的吸收量较少，占最大吸收量氮为5.3～7.9％，磷为7.0～14.4％，钾4.8～7.7％。但品种间有差异，其中M1和NP3的前期氮磷钾吸收积累量要大于S3；越夏期间品种S3对氮磷钾的吸收增加较少，而M1和NP3有缓慢增加；越夏后仙客来对氮磷钾的吸收迅速增加，吸收与积累的规律与仙客来植株的生长发育规律相一致。仙客来对钙、镁及微量元素的吸收动态与氮磷钾的吸收有相似的变化。
二、不同施肥模式对仙客来生长发育和养分吸收利用的研究
根据设施栽培仙客来的养分吸收特点以及优质仙客来生产要求，设置了三种不同的供肥模式，在三个主栽品种上试验，取得了如下结果：
1．根据仙客来的养分吸收规律及供试条件的生长发育状况设置的供肥模式2和模式3与常规供肥模式1相比，可以明显降低越夏期间仙客来的高温逆境胁迫，越夏后恢复生长快，增叶早而多，干物质积累迅速增加，花芽分化多，开花早而整齐，叶下花少，优质花增加。其中，模式3优于模式2；从3个品种看，M1、NP3品种的影响效果优于S3。
2．分析不同供肥模式对仙客来养分吸收的影响，可以发现：不同供肥模式对氮素吸收影响品种间比较一致，模式1＞模式3＞模式2，模式1吸收了过多的氮而不利于仙客来发育。对磷素的影响品种间不一致：品种S3是模式2＞模式3＞模式1，NP3是模式1＞模式3＞模式2，而M1在3种模式下的磷素吸收量差异不大，其中NP3品种在模式1下对磷的过多吸收影响了仙客来的生长发育及开花质量。对钾、钙、镁的影响都是模式2和模式3高于模式1。对微量元素的吸收的影响，模式间和品种间都有差别，但总体来说，供肥模式2更有利于微量元素的吸收。
3．计算不同供肥模式下仙客来养分利用率，可以明显看出：模式2和模式3与模式1相比显著提高了氮磷钾养分的利用率，其中氮利用率提高了1.9～2.5倍，磷利用率提高了3.5～4.6倍，钾利用率提高了，2.2～3.3倍。但不同供肥模式对养分利用率的影响品种间有一定的差异。
4．在中国东南部设施栽培条件下，高温逆境胁迫对仙客来生长发育带来深刻影响，从而导致产量降低，品质差，是优质仙客来生产的重要限制因素。如何弄清仙客来的抗逆境生理，提高仙客来抗高温逆境胁迫的能力，是仙客来生产者十分关心的问题。通过对高温胁迫下仙客来根系活力，叶片中SOD酶活性、MDA含量等有关细胞膜稳定性，可溶性糖含量等生理生化指标的研究，结果表明：模式2和模式3供肥条件下仙客来抗高温胁迫能力大于常规模式1。结合越夏期间仙客来叶片、花芽的生长状况，以及越夏后迅速恢复生长，提前进入旺长期，形成大株型优质花商品的实践也证明了这一点。
综上所述并结合生产实际，对于S3品种由于在供肥模式2和模式3的条件下，生长发育差异不大，而模式2施肥简单，利于操作，在生产中可以选择模式2施肥；NP3和M1品种在模式3下生长显著优于模式2，不仅越夏后叶片生长和花芽分化迅速，且提早上市，品质优，生产中可以选择模式3施肥。
三、钙营养对仙客来花器官发育及优质花形成机理的研究
1．水培条件下，仙客来对钙的吸收与花器官发育关系密切。当营养液中钙浓度分别为0mmol L~(-1)、2.5 mmol L~(-1)和5 mmol L~(-1)时，仙客来对钙的吸收速率分别稳定在-0.6 mg plant~(-1) d~(-1)、5 mg plant~(-1) d~(-1)和15 mg plant~(-1) d~(-1)，花器官中钙含量分别对应为3.372g kg~(-1)d．wt、6.341 g kg~(-1)d．wt和8.697 g kg~(-1)d．wt，而它们相对应的平均开花数量分别为3、8和12，平均花茎高度分别20.2cm、24.7cm和28.3cm。因此，在一定浓度范围内增加营养液中钙的浓度可以提高开花品质。提高营养液中氮的浓度，仙客来对钙的吸收受到显著抑制，钙的平均吸收速率仅为2.5 mgplant~(-1) d~(-1)，花器官中钙含量为3.784 g kg~(-1)d．wt，其花茎高度甚至比缺钙处理还低19.8％。增加营养液中硼的浓度对仙客来钙吸收影响不显著，但促进了仙客来开花数量，同时花器官中钙和硼的比例与5mmol／L钙处理接近。
2．SEM扫描结果表明，花器官中钙含量显著较低的无钙处理和高氮处理其花茎维管组织发育不成熟，维管组织小，细胞排列凌乱，髓射线、木质部和韧皮部无明显区分；而其他花器官中钙含量较高的处理则花茎维管组织内细胞清晰可见，排列规则，髓射线、木质部和韧皮部各部分有明显区分界限，能见到具环形加厚现象的导管分子，其中以5mmol L~(-1)钙处理最明显。因此，钙对花茎组织细胞的发育的影响是其影响花器官发育的原因之一。
3．TEM超微结构表明，花器官中钙浓度显著较低的无钙处理和高氮处理下，花茎细胞叶绿体、线粒体、细胞核等细胞器明显受损。缺钙处理下细胞壁厚度非常不均匀，出现断裂；高氮处理下虽未见细胞壁、细胞膜断裂现象但细胞壁很薄。其它花器官中钙浓度较高的处理下细胞结构正常，细胞器较完整，未见破损现象的发生，随着钙浓度的增加5mmol L~(-1)钙处理下细胞结构较2.5mmol L~(-1)钙处理更完整清晰。
4．钙的细胞化学定位结果表明，花茎细胞中细胞壁和细胞间隙是Ca~(2+)分布的主要场所，细胞膜、核膜及细胞质中也有少量分布。5mmol L~(-1)钙处理下，Ca~(2+)沉淀颗粒最多；而缺钙处理和高氮处理下Ca~(2+)沉淀颗粒很少，几乎没有；增加硼的供应细胞间隙中Ca~(2+)颗粒分布较少，而细胞壁上Ca~(2+)颗粒增多且体积有所增大。对花茎细胞切片进行EDS能谱分析，测定不同钙条件下花茎细胞中钙的相对含量，测定结果与钙的细胞化学定位结果较一致，5mmol L~(-1)处理下花茎细胞中高密度电子体的Ca／O比值最大，花茎细胞钙含量最大。各处理Ca／O比值大小顺序为Ca_0＜+Ca+N＜+Ca≤+Ca+B＜++Ca。
5．Ca~(2+)吸收运输与Ca~(2+)-ATPase活性关系密切。在缺钙处理下，仙客来各器官钙含量显著降低，细胞内钙浓度也减少，各器官Ca~(2+)-ATPase活性也显著降低，可能导致Ca~(2+)信使系统的紊乱。增加外源钙浓度促进各器官中钙含量，也显著提高Ca~(2+)-ATPase活性；增加氮的供应抑制了钙的吸收，Ca~(2+)-ATPase活性也有所下降。
6．分析测定了不同供钙条件下仙客来花茎细胞壁各成分含量的变化，结果表明，随着花器官中钙含量的增加，纤维素含量显著增加，钙含量较低的缺钙处理和高氮处理下细胞壁中纤维素含量是钙含量较高处理的三分之一左右；而果胶含量和半纤维素含量变化随着钙浓度增加而降低；增加氮的供应使细胞壁中半纤维素所占的比重达到了65.6％。随着花器官中钙含量的增加，醛酸在果胶中含量也随着增加，纤维素中呈递减，半纤维素中变化不一；总糖含量的变化与醛酸含量变化规律较相似；蛋白质在半纤维素中含量增加，纤维素中降低，果胶中变化不大。因此，供钙对仙客来花器官发育的影响与细胞壁结构的变化直接相关。
7．不同供钙条件能显著影响仙客来花器官内源激素的含量。钙吸收正常的处理，随着花茎伸长至开花，IAA、ZTs和GA_3含量都增加。缺钙和钙吸收受阻处理中，赤霉素类物质在花茎伸长到盛花期间含量急剧下降，可能导致花茎伸长受阻；缺钙处理IAA含量水平变化与钙供应充足的处理呈相反趋势，也可能导致花茎弯曲、畸形现象；同时，花茎发育后期生长素类物质含量小可能是其维管组织发育不完善的主要因素。高浓度外源钙的处理使花器官中不仅IAA和ZTs含量都高于其他处理，而该处理导管增厚最显著，两者的对应关系密切。
8．设施栽培条件下研究了不同的供钙方式和供以不同钙源对仙客来开花的影响，结果表明，停止外源钙的供应对仙客来开花品质影响较大，花茎高度仅高出叶面2cm左右，达不到仙客来商品要求。CaEDTA在促进仙客来钙吸收上比Ca(NO_3)_2更有效，相对应在促进仙客来开花品质上也更有效。不同的供钙方式下仙客来营养器官钙含量差异较大；根部供钙下钙含量为根＞球茎＞叶片，叶面喷钙下钙含量叶片＞根＞球茎；但花器官中钙含量并没有因为供钙方式差异表现出显著差异。
Cyclamen (cyclamen persicum L.), also called Tu'er Flower, Yipin Guan, etc, belongs to corm flower of Cyclamen genus, Primula malacoides Franch family. By virtue of peculiar hereditary variability of C. persicum, it is a kind of flower fostered through selection generation after generation and intraspecific hybridization and cultivation. This flower has special flower forms, abundant flower color, and long florescence. In developed countries such as America, Europe, Japan, etc., the related industrialization production technology of different varieties has been formed. In our country, the planting history of the cyclamen is short and the industrialized culture just starts. Because of lack of corresponding technical research, the breeds introduced abroad cannot realize the localization in the aspects of seed, culturemedium, and fertilizer. The problems such as low commodity rate, high production cost and lack of management skills of the high-quality cyclamen generally exist. In order to accelerate the sustainable development of cyclamen industry of our country, this research, based on the technological status quo of domestic and international industrialization production of cyclamen, investigates the nutrient absorption rule of cyclamen under the cultivation conditions, different fertilizer mode impact on the growth and development and quality, and calcium influence on the development of flowers and the formation of high-quality flowers. The main results are summarized as follows:
Ⅰ. Growth and development characteristics and nutrient absorption rules of Cyclamen cultivated under facility conditions
Aiming at the problems of blind fertilizer application, low fertilizer utilization ratio, high production cost, obstruction of growth and development, difficulty of the improvement of output and quality, etc., the growth and development characteristics and nutrient absorption rules of the main cultivated cyclamen breeds in our country have been studied and the following results are obtained:
1. The study on the growth rules of three cultivars (S3, M1, NP3) of cyclamen of our country shows that, under the testing conditions, the growth and development of three cultivar breeds are slow before the over-summer period (August 21) due to the influence of high temperature stress; The number of leaves accounts for 30～40% of the total number, and the dry matter weight only accounts for 9.8% - 15.5% of the total number. However, after the over-summer period, cyclamen resumes to grow rapidly, leaf emergence accelerates, the number of leaves rapidly adds, and the accumulation of dry matter accelerates, but there are differences on the repair capacity among the breeds, wherein M1＞NP3＞S3, and M1 firstly starts into the full florescence, S3 is the last and NP3 is in the middle; when reaching the period of commodity (January 12), the maximum number of leaf for S3 is 83, for M1 is 84, and for NP3 is 81, all of which exceed the requirements of leaves of choice cyclamen; the maximum of dry mater accumulation for S3 is 49.9g·plant~(-1), for M1 is 43.2 g·plant~(-1), and for NP3 is 42.2 g·plant~(-1); and the total number of flowers for S3 is 23, for M1 is 31, for NP3 is 29.
2. The determination of the nutrient requirements in the whole breeding time of the three cultivars of cyclamen shows that, under the cultivation conditions, the requirements of nitrogen, phosphorus and potassium for S3 are respectively 1257.25mg·plant~(-1), 218.79mg·plant~(-1), 1171.69mg·plant~(-1), the ratio between them is 5.7:1:5.4, and the total amount of the three elements is 2647.73 mg·plant~(-1); the requirements of nitrogen, phosphorus and potassium for M1 are respectively 1214.14mg·plant~(-1), 222.13mg·plant~(-1), 1292.66mg·plant~(-1), the ratio between them is 5.8:1:5.7, and the total amount of the three elements is 2170.831 mg·plant~(-1); while the requirements of nitrogen, phosphorus and potassium for NP3 are respectively 1003.78mg·plant~(-1), 174.231mg·plant~(-1), 992.82mg·plant~(-1), the ratio between them is 5.5:1:5.8, and the total amount of the three elements is 2728.93 mg·plant~(-1); comparing the required amount of the three elements for the three breeds, the result is M1＞S3＞NP3. The requirements of calcium, magnesium for S3 are respectively 184.50 mg·plant~(-1), 85:40mg·plant~(-1); for M1 are respectively. 203.51 mg·plant~(-1), 97.05mg·plant~(-1); while for NP3 are respectively 273.21 mg·plant~(-1), 127.14mg·plant~(-1); comparing the total amount of calcium and magnesium for the three breeds, the result is NP3＞M1＞S3. The requirements of microelement for cyclamen are also different between different breeds. For the total amount, M1 is greater than S3, NP3.
3. Investigation on the nutrient, absorption rules of cyclamen in different cultivation period shows that, under the testing conditions, the absorption amount of nitrogen, phosphorus and potassium before over-summer period is relatively less, accounting for 5.3～7.9%, 7.0～14.4%, 4.8～7.7% of the total absorption amount of nitrogen phosphorus and potassium respectively. But there are differences among the varieties. The absorption accumulation amount of M1 and NP3 is greater than S3 during the early period; the absorption accumulation rate is relatively less for S3 and slowly increases for M1 and NP3; the absorption of nitrogen, phosphorus and potassium of cyclamen greatly increases after over-summer period, and the rules of absorption and accumulation keeps pace with the growth and development of cyclamen. The absorption trends of calcium, magnesium and microelements are similar to the absorption of nitrogen, phosphorus and potassium.
Ⅱ. Research on the growth and nutrient absorption and utilization under different fertilization modes
Three different fertilization modes are established based on the nutrient absorption characteristics and production requirements of cyclamen cultivated in the different facilities. The following results have been obtained on the three different breeds:
1. Based on the nutrient absorption rules and growth of testing conditions, three fertilizer supply modes are established. Compared with fertilizer supply model, Mode 2 and mode 3 can obviously decrease the high temperature stress during the over-summer period, resume the growth more quickly, and increase leaves earlier and more; accumulates and increases dry matters rapidly, splits up more bud, blooms early and neatly with less flowers below leaves, and quality flower increases. Among them, mode 3 is superior to mode 2; for the three varieties, the influence effects of M1, NP3 are superior to S3.
2. Analysis of different fertilization supply modes effects on the nutrient absorption shows that, different fertilizer supply modes have similar effects on the nitrogen absorptions; that the impacts of mode 1 is greater than mode 3, and mode 3 is greater than mode 2. For mode 1, too much nitrogen absorption is not good for the development of cyclamen. However, the effects on element of phosphorus are inconsistent among different varieties: for S3, mode 2＞mode 3＞mode 1, for NP3, mode 1＞mode 3＞mode 2, but for M1, there is no obvious difference on the phosphorus absorption under the three modes. Among them, for NP3, the surplus absorption in phosphorus under mode 1 influences the development and growth and bloom quality of cyclamen. As to the impacts on potassium, calcium, magnesium, mode 2 and mode 3 are higher than mode 1. For the impact on absorption of the microelements, there are differences among the modes and among the breeds. But as a whole, the fertilizer supply mode 2 is more favorable to the absorption of microelements.
3. Calculation on the nutrient utilization ratio under different fertilizer supply modes obviously shows that, compared with mode 1, mode 2 and mode 3 have obviously improved the utilization ratio of the nitrogen phosphorus and potassium nutrients, among them the utilization ratio of nitrogen has been increased by 1.9～2.5 times, utilization ratio of phosphorus by 3.5～4.6 times, while utilization ratio of potassium has been increased by 2.2～3.3 times. But for the impact on nutrient utilization ratio of different fertile modes, there are some differences among the breeds.
4. Under the facilities culture conditions in the southeast of China, high-temperature stress makes a great influence on the growth of development of cyclamen, which leads to the reduction of output and bad quality. It is an important restricting factor for the production of high-quality cyclamen. How to clarify the stress physiology and improve the capacity of resisting high-temperatures stress is the question that cyclamen producers care about very much. Through the study on biochemical and physiological index on relevant cell membrane steady and soluble sugar contents of root activity, SOD enzyme activity of the leaf, MDA content, it shows that, the capability of resisting high-temperature stress of mode 2 and mode 3 is greater than that of the regular mode 1, which can be also demonstrated by the fact of the characteristics of leaf and bud growth conditions of cyclamen during the over-summer period, the rapid growth and quick stepping into the prosperous growth period in advance after over-summer period, and forming of large high-quality flowers.
Considering the above descriptions and the production practice, for S3, there is no obvious difference in growth and development under the fertilizer supply between mode 2 and mode 3, and it is simple for fertilizing and application in mode 2, so mode 2 can be chosen for fertilizing in the production; the growth conditions of NP3, and M1 are obviously better than mode 2. Not only the leaf growth and bud splitting up is quickly after over-summer period, but also the products can come into the markets in advance with excellent quality, so mode 3 can be chosen for fertilizing in the production.
Ⅲ. Research on the mechanism of floral organs development and high-quality flower formation influenced by calcium nutrition
1. Under the water culture condition, the absorption of calcium is closely related to the floral organs development. When the concentration of calcium is Ommol L~(-1), 2.5 mmol L~(-1) and 5 mmol L~(-1) respectively in nutrient solution, the absorption rates of calcium are steadily around -0.6mg plant~(-1) d~(-1), 5mg plant~(-1) d~(-1) and 15 mg plant~(-1) d~(-1) respectively; the corresponding contents of calcium in the floral organs are respectively 3.372g kg~(-1)d.wt, 6.341 g kg~(-1) d.wt and 8.697 g kg~(-1) d.wt; the corresponding bloom quantity are 3, 8 and 12 respectively; the average height of pedicel is 20.2cm, 24.7cm and 28.3cm respectively. Therefore, the increase of calcium concentration within a certain scope can improve the quality of blooming. The improvement of nitrogen concentration will obviously inhibit the absorption of calcium. The average absorption rate of calcium is 2.5 mg plant~(-1) d~(-1), the content of calcium in floral organs is 3.784 g kg~(-1)d.wt, and the height of pedicel is even 19.8% lower than that under the condition of calcium deficiency. The increase of boron concentration in the nutrient solution has no obvious influence on the absorption of calcium, but it accelerates the quantity of blooming, and the ratio between calcium and boron in the floral organs is close to the results of the group treated with 5mmol L~(-1) calcium.
2. The result of SEM scan shows that under the acalcerosis treatment and high nitrogen treatment for the floral organs containing relatively lower calcium, the vascular tissue of scape is small with irregular cell arrangement and indistinct difference of medullary ray, xylem and phloem. However, under the treatment for other flower organs containing higher calcium, the cell in the vascular tissue of scape which is regularly arranged, is clear, and there is distinct difference in medullary ray, xylem and phloem. Moreover, annular and thickening vessel element can be observed, of which the treatment for 5mmol L~(-1) calcium is the most obvious. Therefore, one of the reasons that calcium has influence on the growth of flower organs is that it has influence on the growth of the scape cell.
3. TEM ultrastructure shows that under the acalcerosis treatment and high nitrogen treatment, the chloroplast, chondriosome and cell nucleus of the scape cell will be damaged obviously. Under acalcerosis treatment, the thickness of cell wall is exceptionally inhomogeneous and broken, but under high nitrogen treatment, although there is no breakage of cell wall and cell membrane, the cell wall is very thin. Under the treatment of other flower organs with relatively high calcium concentration, the cell structure is normal, and the organelle is complete without any breakage. As the increase by 5mmol L~(-1) of calcium concentration, the cell structure becomes clearer and more complete compared with the treatment group of 2.5mmol L~(-1) calcium.
4. The results for the cytochemistry allocation of calcium shows that, the cell wall and intercellular space in the scape cell are the main distribution places of Ca~(2+), which are also distributing in the cell membrane, nuclear membrane and cytoplasm with a little amount. Under 5mmol L~(-1) calcium treatment, there will be maximum amount of Ca~(2+) precipitation grain, and there will be little Ca~(2+) precipitation grain under the acalcerosis treatment and high nitrogen treatment. The distribution amount of Ca~(2+) precipitation grain in the intercellular space of the new added boron is relatively small, while the amount and volume of Ca~(2+) grain on the cell wall increases. Take a section of the scape cell for EDS energy spectrum analysis, and measure the comparative calcium content of scape cell under different calcium conditions. The measurement result is comparatively consistent with the result of cytochemistry allocation of calcium. Under 5mmol L~(-1) calcium treatment, the Ca/O ratio of high density electronic body in the scape cell is relatively high. The order of the Ca/O ratio under all kinds of treatment should be Ca_0＜+Ca+N＜+Ca＜+Ca+B＜++Ca.
5. Absorption and transportation of Ca~(2+) are closely related to the activity of Ca~(2+)-ATPase. Under Calcium-deficiency treatment, the calcium contents in all organs of cyclamen decrease obviously, calcium concentration in cells also reduces and activities of Ca~(2+) -ATPase in all organs also markedly decease. This may cause disorder of Ca~(2+) messenger system. Increase of calcium concentration from external source can improve calcium contents in all organs and obviously promote the activity of Ca~(2+) -ATPase. Increase of nitrogen supply inhibits the absorption of calcium and also decreases the activity of Ca~(2+) -ATPase.
6. The changes of contents of all components in cell wall of cyclamen under conditions with different calcium supply are analyze and measured. The result indicates that with the increase of calcium content in organs, the cellulose content obviously increases. The cellulose content in cell wall under calcium-deficiency and nitrogen-efficiency is about 1/3 of that under treatment with comparatively high calcium content. However, the contents of pectin and hemicellulose decrease with the increase of calcium concentration. The increase of nitrogen supply makes the proportion of hemicellulose reach 65.6%, With the increase of calcium content in flower organs, the content of aldehydic acid increases in pectin, decreases, in cellulose and changes without regular rules in hemicellulose. The rule of change of total sugar content is similar to that of change of aldehydic acid content. The content of protein increases in hemicellulose, decreases in cellulose and changes a little in pectin. Therefore, calcium supply has direct effect on growth of organs of cyclamen and changes of structure of cell wall.
7. The different calcium condition can obviously affect the cyclamen flower organ endogene hormone content. At the calcium absorption normal treatment, IAA, ZTs and GA3 content are increasing along with the floral elongates to blossoms. At the treatments with lacking the calcium and the calcium absorption blocked, the gibberelin class matter content drops suddenly during the period of floral elongates to blossoms. It may cause the pedicel elongation to be blocked. The change of the IAA content level was opposite within the lacking calcium treatment and the calcium supply sufficient treatment. It also possibly causes the abnormal phenomenon of pedicel curving etc. At the same time, the lower content of growth hormone class matter is possibly the imperfect primary factor of vascular tissue growth during later period of pedicel growth. At the treatment with high calcium content supplying, both the IAA and the ZTs content are higher and the incrassation phenomenon of vessel is obviously. Maybe, there are some corresponding relations.
8. Under different facility cultivation condition, studies on the influence of difficult calcium-supplying methods and different calcium resources on cyclamen-blooming show that, stopping the external supply of calcium has greater impact on the quality of cyclamen-blooming, with the height only about 2cm higher than the leave which is beyond the need of the cyclamen goods. CaEDTA is more effective than Ca(NO_3)_2 to promote cyclamen's absorption of calcium, as well as to improve cyclamen-blooming quality. Due to different calcium-supplying methods, tremendous differences exist in calcium content of nutrition organ of the cyclamen, for example, as to calcium supplied by root method, calcium content of root is higher than corm, which is higher than leaves. However it turns out to be different for calcium sprayed on leaves method, with which the calcium content of leaves is higher than root, and root higher than corm. Nevertheless, there is no obvious difference in calcium content in its floral organs resulted from different calcium-supplying methods.
引文
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