摘要
马铃薯是世界第四大粮食作物,也是是新世纪最有发展前景的高产经济作物之一,在我国粮食、蔬菜、饲料和加工业生产上占有重要的地位。但是目前我国马铃薯整体生产水平较低,单位产量不高,先进的实用栽培技术和基础栽培理论比较欠缺。为此开展马铃薯高产栽培技术及高产栽培理论研究,充分发挥马铃薯的生产潜力,节约生产资源,降低生产成本,对推动我国马铃薯再上台阶,提升市场竞争力,对保证我国粮食安全,有效提高农业综合经济效益和增加农民收入,具有重要的意义。本研究选用鲜食品种、高淀粉和低淀粉加工型品种为研究材料,立足于改善促进块茎发育的营养条件,从块茎发育中的激素行为和酶行为的动态变化入手,探讨营养条件对马铃薯块茎发育过程影响的机理,阐明营养条件影响马铃薯产量和品质的内在机制,并从光合作用以及叶绿素荧光特性方面探讨营养条件对马铃薯光能利用的影响机制。同时研究春、秋马铃薯对氮肥的需求特性,为马铃薯的高产栽培,合理的营养运筹,充分发挥马铃薯品种的遗传潜力提供理论依据和参考。研究结果表明:
1.过低、过高的营养供给,降低了块茎发育前期细胞分裂素(ZT)的含量,中期细胞生长素(IAA)的含量以及促进块茎发育的脱落酸(ABA)、茉莉酸(JAs)含量,增加了各时期促进匍匐茎发生的赤霉素(GA_3)含量,导致在块茎形成和膨大的关键时期,GA_3/ABA和GA_3/JAs比值失衡,抑制了块茎的正常发育。而合理营养水平供给,保持了前期较高水平的ZT含量,中期较高的ABA、IAA、JAs含量和较低的GA_3含量,保证了块茎发育各时期对重要激素的需要,GA_3/ABA和GA_3/JAs在块茎发育各时期也保持在合理的水平。
2.马铃薯块茎中的多胺(polyamines)以腐胺(put)含量最多,亚精胺(spd)次之,精胺(spm)最少,三种多胺在块茎发育初期、中期含量高,呈降低—升高—降低的变化趋势,但对不同肥力水平的响应有差异,中等肥力水平处理在块茎发育中期高于低肥力和高肥力水平,高肥力水平使put含量降低,峰值出现时间提前,而使spa和spm峰值出现时间延后。put/spd呈现随肥力水平的提高,峰值不断降低,峰值出现时间提前的趋势。茉莉酸(JAs)与多胺(polyamines)的相关性较大,其中与腐胺(put)为正相关,与精胺(spm)、亚精胺(spa)显著负相关,spm与spd极显著正相关,GA_3与三种多胺均为负相关,但均未达显著水平,JAs与GA_3的相关性也很小。
3.马铃薯的重要光合参数净光合速率(Pn)、气孔导度(Gs)、胞间CO_2浓度(Ci)均随着施氮量的增加出现先增长后降低的单峰变化趋势。在对光量子通量密度(PFD)以及CO_2的响应中,马铃薯净光合速率(Pn)也随着施氮量的增加出现先增加后降低的趋势。从氮营养对马铃薯需光特性、CO_2需求特性以及叶绿素荧光特性的影响看,最大净光合速率(Am)、表观量子效率(Ψ)、CO_2饱和点(CSP)等参数也随着施氮量的增加呈现先升高后下降的变化趋势,光化学猝灭系数(qP)、实际光化学效率(φPSⅡ)、电子传递速率(ETR)也存在同样的变化规律。
4.不同类型的马铃薯品种在光能利用上表现出一定的差异。高淀粉品种QS2的气孔导度(Gs)、蒸腾速率(Trmmol)、光化学猝灭系数(qP)、非光化学猝灭系数(qN)、在同样施氮条件下高于低淀粉品种C117,且光能利用率(LUE)和净光合速率(Pn)比低淀粉品种C117更具优势。但是低淀粉品种C117在各施氮条件下的水分利用率(WUE)、光饱和点(LSP)、光补偿点(LCP)、潜在光化学效率(Fv/Fm)和实际光化学效率(φPSⅡ)比高淀粉品种QS2高,且C117在同水平施氮条件,不同浓度CO_2下的最大净光合速率(Am)、CO_2补偿点(CCP)高于QS2。
5.不同施氮量下块茎总淀粉、支链淀粉和直链淀粉的含量呈“S”曲线变化,符合Logistic方程Y=K/(1+ae~(-bT)),但是不同施氮量对各淀粉积累强度,达到最大积累强度的时间存在差异,品种之间的影响也不尽一致。对于低淀粉品种C117,随着施氮量的增加,直链淀粉的最大积累强度呈逐渐下降,支链淀粉呈不断上升,总淀粉呈先上升后又下降的变化趋势。对于高淀粉品种QS2,直链淀粉的最大积累强度在中间施肥水平较低,而在低、高施肥水平时较高,支链淀粉、总淀粉随着施氮量的增加呈先升后降的变化趋势。
6.随着生育期的推进,各马铃薯块茎中不同施氮水平下的ADPGPPase、SSS、GBSS、SBE呈现先逐渐增加,达到最大值后又开始下降的单峰曲线变化趋势,各施氮水平均在出苗后54—61d左右达到最大值。随着施氮量的增加,ADPGPPase、SSS也出现先升后降的单峰曲线变化趋势,而对于GBSS和SBE的影响,品种间表现差异。低淀粉品种C117的ADPG-PPase、SSS的含量在各施氮水平下均高于高淀粉品种QS2,具有较高的ADPG-PPase、SSS活性。在氮营养影响条件下,SSS活性与淀粉总含量和支链淀粉的含量极显著正相关(r总淀粉=0.86~(**),r支链淀粉=0.83~(**)),而与直链淀粉含量相关不大。
7.氮肥运筹结果说明,底肥和追肥对春马铃薯产量影响很大,合理追肥的增产作用明显,对于秋马铃薯,底肥的增产作用显著,追肥的增产作用不明显。因此春马铃薯的氮肥运筹应在重视底肥的基础上,特别重视追肥的使用,而对秋马铃薯的氮肥运筹则要特别重视底肥的使用,追肥要视具体情况慎重使用,做到少量、早施,以取得最大的追肥效果。
Potato ranges the forth of food crops in the world while it is increasing importance in Chinese food,vegetable,feed and processing.However the unit yield and production level are low and the cultivation technique is lacking today than those of developed countries.So it is very significant to study the high-yield culture techniques in order to improve the potential of production and increase the economic benefit.It is also important to promote the market competitive power in the world,to assure the food safety in our country and to raise the comprehensive economic benefits in agriculture,so it can push the processing industry of potato enter a new stage in the country.In order to reveal the mechanism how nutrients promote tuber development under different nutritional levels,the flesh-edible potato,high yielding variety,high-starch and low-starch content potato have been used as experiment materias.The first objective of this study was to reveal the changes of endogenous hormones,polyamines(PAs),yield and quality in the process of tuber development in potato(Solanum tuberosum L.)under different nutrition.The second objective was to analyse the the dynamic accumulation of starch,amlose and amylopectin and to determine the patters of the actives of key enzyme involved in starch synthesis and to find the relationship between the accumulation of starch and key enzyme in the process of buberization uder different levels of nitrogen.The third objective was to discuss the characteristic of photosynthesis and chlorophyll fluorescenceS under different nutritional conditions in order to reveal the mechanism how nutrition influence the photosynthesis of potato.The expected results would help understand the mechanism how nutrients affect the tuber development and obtain a new technique of high yield.The main results are as follows:
1.In tuber development,under- or over-supply of nutrients reduced ZT content at initial stage and IAA,ABA and JAs content at middle stage,but increased GA_3 content in the whole process,which caused GA_3/ABA and GA_3 /JAs to be off balance in the key stage of tuberization.Appropriate nutrient supply kept higher ZT content at initial stage and higher ABA,JAs and IAA content and lower GA_3 content at middle period,which could satisfy the demand of important hormones in every stage of tuberization and keep GA_3/ABA and GA_3/JAs at reasonable levels during tuber formation.
2.The contents of three kinds of PAs(put,spm and spd)have two peak values at early and middle stages in the tuber development respectively,comforming with the trend of decrease-increase-decrease.However the contents had different respondenc to different fertility levels.In middle stage of tuberization PAs contents under moderate fertility level were higher than that under low and high fertility levels.High fertility made the put content decrease,the peak time of put advance,and the peak time of spm and spd postpone. With the increase of fertility the peak value of put/spd reduced continuously and the peak time advanced.Under low fertility level JAs content was low in all stage.And in the middle stage the JAs content under moderate level was higher than those under low and high fertility levels.The GA_3/JAs under low and high fertility levels increased firstly and then decreased.The change trend was reverse to that under the moderate fertility level.JAs content had significant positive correlation with the content of put,and significant netive correlation with the contents of spd and spm.
3.The photosynthetic rate(Pn),stomatal conductance(Gs),intercellular CO_2 concentration(Ci)of potato showed a single-peak trend with the amount of nitrogen increasing.As far as responsing to the light quantum flux density(PFD)and CO_2,the Pn showed the similar trend.To the nitrogen nutrition on the characteristic of light and CO_2 demand of potato,the maximum net photosynthetic rate(Am),the apparent quantum efficiency(Ψ),CO_2 saturation point(CSP)increased at first with raising of nitrogen levels then diecreased with further nitrogen enhancement At the effects of nitrogen on the chlorophyll fluorescence characteristics,the actual photochemical efficiency of PSⅡ(ΦPSⅡ),photochemical fluorescence quenching(qP)and electronics trancelation rate(ETR)showed of the same changes.
4.There were some differences in the use of light energy at different varieties of potatoes.To the high-starch varieties QS2,the stomatal conductance(Gs),transpiration rate(Trmmol),photochemical quenching factor(qP),non-photochemical quenching factor(qN)and CO_2 saturation point(CSP)were higher than the low-starch variety C117 in the same level of nitrogen.Furthermore the light use efficiency(LUE)and net photosynthetic rate(Pn)also had more advantages than the low-starch variety C117. However to the low-starch varieties C117,the efficiency of water(WUE),the light saturation point(LSP),the light compensation point(LCP),the CO_2 concentrations point(CCP),the potential photochemical efficiency(Fv/Fm)and actual photochemical efficiency(ΦPSⅡ)were higher than the high-starch variety QS2 under the different conditions of nitrogen fertilizer.At the same conditions of nitrogen,the maximum net photosynthetic rate(Am)of C117 was higher than the QS2 in different concentration of CO_2.However,the light use efficiency(LUE)and photosynthetic rate(Pn)of high-starch cotent variety was higher than the C117 at NO,N1,N2 levels of nitrogen,expressing a stronger ability of using light.
5.The amylose,amylopectin and starch content(the sum of amylose and amylopectin content)increased in "S" curve fashion in the process of tuber development under different levels of nitrogen,and the curves satisfied the Logistic function of Y=K/(1+ae~(-bT)),but the greatest accumulated intensity and the time reaching the peak exsisted difference at different levels of nitrogen and different varietyis of potato.For the low-starch variety C117,with the increasing of nitrogen nutrient,the greatest accumulation intensity of amylose showed decling tendency,and the amylopectin showed the increasing trend while the starch showed increasing until reach the peak then decreasing.The time of reacing the gretest accumulated intensity of amylose and starch esist the trend of increasing at first and then decreasing,but the amylopectin showed the decreasing constantly.For the high-starch content variety of QS2,the gretest accumulated intensity of amylose were lower with middle level of nitrogen than that of low and high level of nitrogen,but the trend of amylopectin and starch increasing at first and then decreasing with increasing of the nitrogen levels.As a result the amylase,amylopectin and starch content of QS2 were lower than C117,not reading it's potential of stach accumulating as a high-starch variety under the circumustance in Sichuan.
6.In the process of tuber development,all of the ADPGPPase,SSS,GBSS,SBE content increased constantly after reaching the maximal then decreasing until the last, showing the single peak trend at every nitrogen level and reaching a stationary phase of maxismal content at 54-61 days after emergence of seeding.As far as to the effects of nitrogen on key enzyme,the ADPGPPase,SSS showed the same changes of single peak trend,but for the influence of the GBSS and the SBE,different varieties had different results.The content of the ADPG-PPase,SSS of low-starch variety of C117 was higher than the high-starch content variety of QS2 under each nitrogen levels,showing the significant activety.At the influence of nitrogen nourishment on correlation of the enzymes and starch accumulation,the SSS activity was significantly and positively correlated with the starch and amylopectin content(r_(starch)=0.86~(**),r_(amylopectin)=0.83~(**)),but was not significantly and positively correlated with the the amylose content.
7.The result of nitrogen application on potato showed that all of the base fertilizer and the topdressing of nitrogen were important on yields of the springs potato,especially to the topdressing,and the yield increased more obviously than the base fertilizer.But for the autumn potato,the function of base fertilizer was more important than the topdressing of nitrogen on the yield,and the action of topdressing was absent.So in order to obtain the high yield of spring potato,we should think much of topdressing besides the base fertilizer, but for the autumn potato,it should play important role of base fertilizer,and topdressing of nitrogen should be specially cautious.
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