【作者】 李晶；

【导师】 祖伟；

【作者基本信息】 东北农业大学 ， 作物栽培学与耕作学， 2009， 博士

【摘要】 综合中国各地推广应用小黑麦的栽培技术可知,大部分引种地区充分利用了小黑麦耐贫瘠、抗性好的特点,但有关小黑麦的具体栽培调控措施仍未引起足够的重视。本研究于2006~2008年度,以八倍体东农5305和六倍体东农96026为材料,设施氮水平(N)和栽培密度(D)两个因子,施氮量0 kg/hm2(N0)、75kg/hm2(N75)、150kg/hm2(N150)、225kg/hm2(N225);密度300万株/hm2(D300)、450万株/hm2(D450)、600万株/hm2(D600)。分析了不同密度和施氮水平下小黑麦生长发育、光合特性、产量形成、子粒品质和氮代谢相关酶活性的变化规律,明确了密度和施氮水平对不同倍性小黑麦产量和子粒品质形成的生理基础,为深化和拓展小黑麦子粒高产、品质改善和指导不同倍性小黑麦优化栽培提供了理论基础和技术途径。1适当施肥降低密度,提高小黑麦茎蘖成穗率,促进子粒产量的提高。合理的基本苗数的确定,可以控制拔节前无效分蘖的发生,降低高峰茎蘖与最终穗数的比值,提高茎蘖成穗率来实现高产。分蘖至成熟各时期每公顷总茎数随密度增加而增加,但成熟期成穗率东农5305随密度增加而降低,东农96026成穗率随密度增加先增加后下降。各氮素营养水平条件下生育期间总茎数及分蘖成穗率均高于对照N0处理;同一密度条件下两个品种茎蘖成穗率均随施氮量增加呈先上升后下降趋势,其中东农5305以D300N75处理最高,东农96026以D450N150处理最高,此时产量也最高。2不同倍性小黑麦要实现高产,应侧重不同主攻方向。对于分蘖能力较高的东农5305应适当降低基本苗,少施氮肥,主攻穗粒数和粒重,东农96026在控制适宜群体基础上,适量施氮,协调产量构成三因素,实现二者高产。两品种穗数随密度增大而增大,而穗粒数和粒重与此相反;氮肥的不断增加,两品种公顷穗数和穗粒数有先增加后下降趋势,两品种粒重随氮肥的施加下降。表明增加密度有效穗数显著增加,对穗粒数和粒重起到负作用,密度过高增加的穗数并不能抵消穗粒数和千粒重的降低,产量不升反降;施加氮肥有效穗数和穗粒数的增加弥补了粒重的降低,利于产量提高;密肥对穗粒数的互作影响较粒重明显。3适宜的密度及施氮水平有利于小黑麦群体光合能力改善。生育前期东农5305中密度处理略高于其它密度处理,东农96026低密度处理叶绿素含量较其它处理高,随生育进程的推迟,高密度处理较低密度处理下降幅度大,叶绿素含量随密度的增高而呈下降趋势,花后28d东农5305低密度处理叶绿素含量显著高于高密度处理;随施氮水平提高叶绿素含量有先增加后降低的趋势,光合速率表现出与叶绿素一致的规律。适宜的密度及施氮一方面提高了光能转换效率和PSⅡ的潜在活性,另一方面抑制了光能的非光化学耗散,过量施氮无助于光合性能的提高。东农5305低密度处理各生育时期光化学淬灭系数均高于中密度和高密度处理,东农96026各施氮水平下中密度处理各生育时期光化学淬灭最高,两品种各施氮处理光化学淬灭系数均较对照N0高,东农5035施氮75kg/hm2时旗叶获得最高的电子传递活性,东农96026施氮150kg/hm2时出现最高的光化学淬灭;东农5305、东农96026各施氮水平下的低密度处理、中密度处理各生育时期非光化学淬灭最低,东农5305施氮75kg/hm2时非光化学淬灭最小,东农96026施氮150kg/hm2时出现最小的非光化学淬灭。4降低密度和保证氮肥供应可以提高小黑麦子粒营养品质。降低密度和保证氮肥供应提高了子粒蛋白质含量。成熟期东农5305子粒蛋白含量随密度增加而降低,低密300万株/hm2蛋白含量最高;而东农96026则在中密450万株/hm2水平下蛋白含量最高。两品种在各密度水平上,各施氮处理子粒总蛋白含量均高于N0处理,施氮量由150kg/hm2增加到225kg/hm2,子粒蛋白含量下降。降低密度增加了小黑麦子粒清蛋白和球蛋白含量,密度过高醇溶蛋白和谷蛋白含量也有降低趋势;四种蛋白组分含量随氮肥增加先增加后降低,适当增施氮肥能有效增加蛋白组分含量,醇溶蛋白和谷蛋白含量受氮肥的调控作用较清蛋白和球蛋白含量明显,密度和氮肥互作效应较氮肥作用显著。东农5305和东农96026各施氮处理氨基酸含量均明显高于对照未施氮处理,超过N150水平的N225处理氨基酸含量反而降低。同一氮素水平下,随密度增加,氨基酸总量有下降趋势,密度间氨基酸含量的差异较施氮处理小。东农5305和东农96026两品种成熟期子粒赖氨酸含量与蛋白质含量均呈极显著正相关,两品种成熟期氨基酸含量与蛋白质含量呈极显著正相关。5适当加大密度有利于改善小黑麦子粒烘烤品质,氮肥效应因小黑麦倍性而异。东农5305随群体密度的增加,子粒容重呈增加趋势,东农96026变化不显著,施氮显著增加了子粒容重;中高密度湿面筋含量高于低密度水平,施用氮肥湿面筋含量有增加趋势,影响不显著;两品种在高密度下降落值有所降低,施加氮肥东农5305降落值有所增加,而东农96026有下降变化。施氮和增加密度有利于提高子粒容重和湿面筋含量;沉降值大小受栽培措施影响甚微,主要因品种而有所不同;中密度下可以获得高的降落值,氮肥对降落值的改善作用因小黑麦倍性而不同,八倍体施加氮肥有改善作用,而对六倍体则起到负作用。6适当施肥降低密度保证了小黑麦子粒高蛋白含量形成的生理基础。增施氮肥促进了花后叶片的氮素同化,降低密度蛋白质的降解能力被削弱。增加施氮量旗叶NR和GS活性提高,内肽酶和氨肽酶活性降低;旗叶NR和GS活性随密度的加大而下降,低密度D300处理下活性最高,内肽酶和氨肽酶活性随密度升高而增加;花后各时期,旗叶可溶性蛋白质含量与硝酸还原酶及谷氨酰胺合成酶活性呈极显著相关,与内肽酶及氨肽酶呈极显著负相关。两品种氮素转运量、转运效率、转运氮贡献率以及N收获指数表现为随氮肥施加先增加后下降,氮肥的影响作用均达极显著水平。两品种子粒蛋白含量和产量均与花后氮素积累的相关最佳,通过提高花前贮存氮素转运量可以实现子粒氮的积累,对子粒蛋白质含量和产量的提高有极显著的促进效应;氮素积累和转运与子粒蛋白质组分含量的相关性表明,氮素的积累和运转是提高蛋白组分含量的关键因素,氮素的积累更利于醇溶蛋白含量的提高,氮运转更利于清蛋白含量的提高,清蛋白和醇溶蛋白含量增加是小黑麦子粒蛋白质含量增加的主要原因。7质谱鉴定结果进一步验证了适当增施氮肥的正向效应。通过MALDI-TOF-MS和生物信息学分析找到7个表达差异蛋白,即与蛋白质合成和信号传导相关的蛋白、与清除自由基有关的酶、与叶绿体的发育和代谢有关蛋白、与植物碳代谢有关的酶、与氮素同化有关的酶。这些蛋白表达量显著增加,进一步说明氮素对光合的影响是通过对光合有关酶的活化来调节植物的光合作用;氮素对于植物衰老的延缓可能与其增强氧自由基清除密切相关;氮素不仅在维持光合性能方面起作用,而且增强了信息传导提高了植物体抗性;氮素的增加直接促进了氮素代谢和同化;氮素对蛋白质合成也有积极影响。氮素对植物生长发育、物质代谢和信号传导影响全面,从蛋白点的鉴定分析来看,功能蛋白是其影响的重点。更多还原

【Abstract】 From the cultivation techniques of triticale around China, we know most species Introduction region adequately utilizes the characteristics of triticale with strong resistance,However, the concrete control measures about triticale has not been fully explored and explained. The case study was carried out in 2006~2008, two triticale cultivars octaploid Dongnong5305 and sextuploid Dongnong96026 were selected. There were two factors in the experiment: nitrogen application (N) and density treatment (D).Four nitrogen applications were as follow: 0kg/hm2(N0), 75kg/hm2 (N75), 150kg/hm2 (N150), 225kg/hm2 (N225). Three density treatments were namely D300 (3million basic seedling per hectare), D450 (4.5million basic seedling per hectare), D600 (6million basic seedling per hectare).The study analyzed the changing law of triticale growing development, photosynthetic characteristics, grain yield and quality, enzymatic activity about nitrogen metabolism under different density and nitrogen application. Besides, physiological basis of grain yield and quality in different ploidy triticale has been discussed to provide the theory base and technical approach for improving triticale grain yield and quality and giving directions to cultivate different ploidy triticale.1. Fertilizing properly and reducing the density can increase the percentage of earbearing tiller and grain yield of triticale.Determining the appropriate number of basic seedlings can control the nonbearing tillering being produced before jointing stage and reduce the ratio of peak tillering to final spike number and increase percentage of earbearing tiller to attain the high output. The total number of stems per hectare increased with density increasing from the tillering to the maturity, but the percentage of earbearing tiller of Dongnong5305 decreased with density increasing in mature period, while the percentage of earbearing tiller of Dongnong96026 first increased then declined. The total number of triticale stems and the percentage of earbearing tiller during growth periods were higher than that of control treatment N0.In the same density, percentage of earbearing tiller of the two varieties both presented increased then downward with the nitrogen increased., thereinto the tiptop of the two respectively presented to: D300N75 treatment of Dongnong5305, D450N150treatment of Dongnong96026, and This is when the highest yield attained.2. The main direction should be angled In order to attain good harvest in different ploidy triticale. Dongnong5305 which has the higher tiller capability should appropriately reduce the number of basic seedlings and the level of nitrogen and increase grains/ear and kernel weight. Based on the suitable group controlled, Dongnong96026 should apply nitrogen fertilizer properly and coordinate yield component to realize the high production. The spike number of the two varieties increased with the density increasing, however, grains/ear and kernel weight with opposite effect. With the increasing of nitrogen fertilizer application, the spikes per hectare and the number of grains per spike of the two varieties presented increased then downward, and grain weight reduced. The result showed that the number of productive ear increased significantly with the density increasing which produced negative effects to grains/ear and kernel weight. The density was so high that the added spikes could not counteract the reducing of the number of grains per spike and 1000-grain weight, the yield did not rise but lower. The increasing of the number of productive ear and grains/ear made up for the lower grain weigh that beneficial to yield improvement. Density and fertilizer has produced an obvious effect on grains/ear compare to grain weight.3 Suitable density and nitrogen fertilizer level were favorable for improving the photosynthetic capacity of triticale.In early growth stages, the density treatment in Dongnong5305 was slightly higher than the other treatments. Low density treatment in Dongnong96026 has higher comparing to the other treatments. With the growing process postponed, the decline rate at high density treatment was bigger. Chlorophyll content reduced with density increasing, 28d after flowering Dongnong5305 at low density was significantly higher than high density. With the increased nitrogen rates, the overall trend of chlorophyll content was first increased then declined, photosynthetic rate showed consistent rule with chlorophyll.Proper density and nitrogen application improved light conversion efficiency and the potential activity of PSⅡon the one hand, on the other hand, inhibited the non-photochemical dissipation, excessive nitrogen was not conducive to improving photosynthetic characteristics. Dongnong5305 photochemical quenching coefficient at low-density was higher than the medium-density and high-density treatment in growth periods.Dongnong96026 photochemical quenching at the medium-density was highest under different levels of nitrogen fertility treatment in growth periods .The two varieties photochemical quenches coefficients which under various nitrogen treatments was higher than contrast N0,Dongnong5035 flag leaf got the highest electronic transfer activity with N level 75kg/hm2. Dongnong96026 showed the highest photochemical quenching with N level 150kg/hm2. The lowest non-photochemical quenching of the two varieties that under various nitrogen treatments in growth periods respectively presented to: low-density treatment of Dongnong5305, medium-density treatment of Dongnong96026. The smallest non-photochemical quenching of the two varieties were respectively: N level 75kg/hm2 of Dongnong5305, N level 150kg/hm2 of Dongnong96026.4 Reducing the density and proper nitrogen application could improve grain quality of triticale. Reducing the density and proper nitrogen application increased grain protein content. Grain protein content of Dongnong5305 brought down with the density increasing during maturity stage, grain protein under 3 million basic seedling per hectare was highest; while grain protein of Dongnong96026 under 4.5 million basic seedling per hectare was highest. Grain total protein content of each nitrogen application treatments were higher than N0 treatment under different density treatment of two varieties. Grain protein content reduced with nitrogen application increased from 150kg/hm2to225kg/hm2. Reducing the density increased grain albumin and globulin content while increasing the density decreased gliadin and glutenin content; four protein components content in grains increased then decreased with nitrogen application increased, increasing nitrogen application properly could effectively increase protein components content, regulative function of nitrogen on gliadin and glutenin content was more obvious than on albumin and globulin content, interaction effects of density and nitrogen fertilizer was better than effect of nitrogen fertilizer.Amino acids content of each nitrogen application treatments were higher than N0 treatment in two varieties. Amino acids content of N225was lower than that of N150. Amino acids content reduced with the density increased under the same nitrogen level,difference of amino acids content among nitrogen treatment was more obvious than that among density treatment. There was significantly positive correlation between lysine content and protein content during maturity stage in two varieties, the correlated relationship between amino acids content and protein content showed the same trend.5 Increasing the density was favorable to improve grain baking quality of triticale, effect of nitrogen varied from the ploidy of triticale.Grain volume wight of Dongnong5305 showed an increasing trend with the density increased, but the change of Dongnong96026 was not significant, nitrogen application significantly increased grain volume wight; wet gluten content of middle and high density were higher than that of low density, wet gluten content showed an indistinctive increasing trend with nitrogen application increased; falling number reduced under high density in two varieties, falling number of Dongnong5305 increased while that of Dongnong96026 decreased when nitrogen application increased. Increasing nitrogen application and density was favorable to enhance grain volume wight and wet gluten content; effect of cultivation measure on sedimentation number was little and varied from varieties; it could obtain higher falling number under middle density, effect of nitrogen on falling number varied from ploidy, increasing nitrogen application had a positive effect on octoploid but a negative effect on sextuploid.6 Reducing the density and proper nitrogen application guaranteed physiological basis for formation of high grain protein content.Increasing nitrogen application promoted nitrogen assimilation of leaf after flowering, reducing the density weakened degrading ability of protein. Increasing nitrogen application enhanced the activity of NR and GS and decreased the activity of endopeptidase and aminopeptidase in flag leaf; the activity of NR and GS in flag leaf decreased with density increased, the activity of NR and GS under D300 treatment was highest, the activity of endopeptidase and aminopeptidase increased with density decreased; soluble protein content in flag leaf had significant positive correlation with the activity of NR and GS, and had significant negative correlation with the activity of endopeptidase and aminopeptidase after flowering. N translocation amount、N translocation rate、translocation N contribution rate and N harvest index increased then reduced with nitrogen application increased in two varieties, effect of nitrogen had reached significant level. Grain protein content and yield showed a significant positive with N accumulation after flowering, enhancing storage N translocation amount before flowering could accumulate N of grain and significantly enhance grain protein content and yield; the correlation between N accumulation and translocation and grain protein components content showed that N accumulation and translocation was the key factor that enhanced protein components content, N accumulation was favorable to increase gliadin content, N translocation was favorable to increase albumin content, albumin and gliadin content was the important reason for increasing grain protein content.7 The result of identified by mass spectra verified the positive effect of increasing nitrogen application.7 Differential expression proteomes were found by MALDI-TOF-MS and bioinformatics analysis, these were proteome related with synthesis and signal transduction of protein, enzyme related with. Elimination of the free radical, proteome related with chloroplast growth and metabolism, enzyme related with carbon metabolism, enzyme related with nitrogen assimilation.The increasing of proteomes expression amount further showed that the effect of nitrogen on photosynthesis is based on the activation of photosynthesis relevant enzymes to adjust the plant photosynthesis, nitrogen could delay senescence of plant might be closely related with strengthening elimination of oxygen free radical; nitrogen not only played role in keeping photosynthesis performance but also strengthened information conduction and improved the resistance of plant; increasing of nitrogen directly promoted nitrogen metabolize and assimilate; nitrogen had positive impact on protein synthesis; Nitrogen overally affected on growth、substance metabolism and signal conduction of plant, identify and analysis on protein spots showed that the emphasis of its influence was functional proteins.更多还原