一氧化氮对小麦产量和蛋白质品质及茎蘖位叶片衰老进程的调控
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摘要
一氧化氮（NO）是植物体内重要的信号分子，与许多生理进程密切相关，包括逆境调节与衰老控制。为了明确NO对小麦生长调节的生理机制，本研究采用以下两种方法：1.喷施外源NO对干旱胁迫下小麦叶片光合及叶绿素荧光特性及籽粒蛋白组分、GMP含量和粒度分布的影响。2.内源NO的氮肥调节及其含量变化对小麦不同茎蘖位衰老的影响。试验1在雨养条件下进行，在小麦生育主要时期（拔节期、抽穗期和开花期）进行不同浓度NO喷施处理；试验2选用两个不同穗型的高产冬小麦品种济南17（多穗型）和泰农18（大穗型）作为研究对象，设定不同的施氮水平，研究从拔节期到开花期不同茎蘖位小麦主要功能叶片内硝酸还原酶（NR）活性和NO含量的变化及衰老的发生。主要研究结果如下：
1外源一氧化氮对干旱胁迫下冬小麦叶片光合特性及籽粒蛋白质组成的调节
1.1外源一氧化氮对干旱胁迫下小麦叶片光合及叶绿素荧光特性的调控
不同时期外源NO处理显著提高干旱胁迫下叶片的光合速率，随着NO处理浓度的增加，光合速率表现出先升高、后降低的趋势，低浓度NO显著提高干旱胁迫下叶片的光合速率并使其一直保持较高水平；叶片的蒸腾速率（Tr）的影响因时期、品种和处理浓度不同而存在差异，总的来说，随着NO处理浓度的增加，表现为先升后降的趋势，但是，外源NO对不同小麦品种类型的影响存在差异，可以显著提高多穗型品种的蒸腾速率，而对大穗型品种作用不明显。低浓度NO对气孔导度表现为促进作用，高浓度则抑制，适量NO浓度处理对维持干旱胁迫下小麦叶片气孔开放具有积极意义。外源NO处理提高了干旱胁迫下叶片的瞬时水分利用效率，并在C2（0.4mmol L-1）浓度下达最大值，不同NO处理时期间影响效果不同，拔节期>抽穗期>开花期。外源NO提高叶片了叶绿素含量，但是对类胡萝卜素含量无显著影响。NO能提高小麦最大光化学效率（Fv/Fm），以拔节期喷施效果最显著。喷施NO处理增加了光合电子传递能量占总吸收光能的比例，提高了光能利用率。
1.2外源一氧化氮对干旱胁迫下不同基因型小麦籽粒蛋白组分的调控
干旱胁迫条件下，通过对4个不同筋型小麦籽粒蛋白质组成和谷蛋白大聚合体含量及粒度分布的研究显示，外源NO处理提高了小麦籽粒中的谷蛋白比例，而对清蛋白、球蛋白和醇溶蛋白的影响存在品种间差异，说明小麦籽粒谷蛋白的合成与积累受NO的调节。在外源NO作用下，更多的贮存蛋白以谷蛋白形式存在。NO处理可以增加强筋型品种的清蛋白含量，降低醇溶蛋白含量，对球蛋白调节效果不明显。外源NO增加了中筋型品种的球蛋白含量，降低清蛋白含量，醇溶蛋白则对NO调节表现不敏感。
2内源一氧化氮对冬小麦茎蘖位叶片衰老的调节
2.1内源NO含量对小麦不同茎蘖位叶片衰老进程的调节
NO含量的改变可以引起衰老。在主茎和一级分蘖中，叶片内NO含量在抽穗期达到最大值，然后在开花期迅速下降，与其叶片DNA片段化的发生时间一致。从拔节期到抽穗期，二级分蘖叶片中NO含量一直处于较高水平，二级分蘖叶片DNA降解发生时间最早。表明NO确实与叶片的衰老密切相关，而且NO含量的骤然改变可能是引起衰老的主要原因。
2.2施N水平对小麦叶片内源NO产生和NR活性的影响
拔节期和挑旗期，不同蘖位叶片内NO含量和NR活性都呈显著正相关关系，增施氮肥提高NR活性及NO含量，表明NO的产生主要依赖于叶片内的NR途径，并且氮肥对二者具有调节作用。随着生育时期进程，叶片内NR活性与NO的产生的相关性减弱，甚至在抽穗期二者无相关性，说明对于孕穗期后NO的产生，叶片内合成作用减弱。不同氮肥处理条件下，NO含量和NR活性都存在着饱和性，二者都是先升高，在达到最大值之后再降低。在抽穗期，叶片内NO含量和NR活性达较高水平。
2.3茎蘖位衰老进程对产量和蛋白质含量的影响
施氮量低于N-240处理时，增施氮肥可以获得更多的有效穗数。N-240处理条件下，济南17和泰农18的每个植株分别可以产生最多的分蘖数3.84个和1.94个，继续增施氮肥至N-360，穗数降低，济南17为3.19个分蘖，泰农18为1.69个。两品种在施氮量N-240处理时，产量最高，且产量的形成与一级分蘖的成穗率密切相关，后者受氮肥施入量的影响。N-360处理条件下，一级分蘖叶片的衰老较晚，直到开花期才开始，但是成熟后单位面积有效穗数少于N-240处理，说明在开花期有很多无效分蘖死亡，这些无效分蘖在消耗了大量养分后死亡，导致产量相对N-240处理下有所降低。施氮量影响籽粒中蛋白质含量，增施氮肥增加籽粒中的蛋白质含量。与主茎相比，一级分蘖籽粒中蛋白质含量显著减少，增加氮肥施入量，则减少主茎与一级分蘖间的含量差异。表明，通过控制氮肥施入量调节一级分蘖有效穗数及其籽粒品质的途径是可行的。
Nitric oxide (NO) is a key signaling molecule in different physiological processes ofplants, including adapting stress and senescence. The objective of this study was to determinethe effects of NO in wheat growing period. Field-experiments were conducted in2009-2011growing seasons used exogenous and endogenous two kinds of methods to certain:1.Theeffect of exogenous NO on photosynthetic characteristics, chlorophyll fluorescence in leavesand protein components, GMP content and particle size distribution in grains under droughtstress.2. The effect of endogenous NO on senescence of tillers in leaves in wheat. The firstexperiment was carried out in water stress condition which depending on raining entirely,with different concentrations on NO at different spraying time (jointing stage, heading stageand anthesis) was conducted. The second experiment was carried out under different nitrogenlevels using two wheat cultivars with different panicle-type. The main results were as follows:
1. The adjustment of exogenous nitric oxide to leaves photosynthetic characteristicsand grain protein composition under drought stress in winter wheat
1.1The adjustment of exogenous nitric oxide to leaves photosynthetic characteristics andChlorophyll fluorescence properties under drought stress in winter wheat
Exogenous NO improved the leaf photosynthetic rate under drought stress significantly,and with concentration increasing, photosynthetic rate rose at first and then declined. Lowconcentration NO increased leaf photosynthetic rate significantly and made it keep higherlevel. Transpiration rate was influenced by period, variety and NO concentration. In generallyspeaking, with increasing of concentration, it first increased and then trends dropped.However, exogenous treatment was not same at different wheat varieties. Low NOconcentration promoted the stomata conductance and high concentration was suppressed. Ithad a positive meaning for NO to maintain suitable stomata opening wheat under droughtstress. NO improved instantaneous water use efficiency of leaves under drought stress, andC2(0.4mmol L-1) improved significantly. Different effects showed as different periodprocessing, jointing stage>heading stage>anthesis. NO improved the chlorophyll content buthad no significantly effect on carotenoid content. It increased Fv/Fm most significantly spraying at jointing stage. Because of increasing the ratio of photosynthetic electron transferenergy to total absorption of light energy, the utilization rate of light energy improved.
1.2The adjustment of exogenous nitric oxide to protein composition under drought stress inwinter wheat
Under drought stress condition, we researched different wheat types for grain protein,GMP content and particle size distribution. It showed that, exogenous NO treatment improvedproportion in glutenin, but to albumin, globulin and gliadin, it influenced differently betweenthe varieties. More storage protein in the cereal was the form of glutenin with the NOspraying. NO increased the strong gluten varieties of albumin content type and reducedgliadin content but had no obviously effect on globulin. It increased in the weak glutencultivars type of globulin content, declined albumin content and was not sensitive to gliadin.In conclusion, it was known that NO adjusted the ratio of glutenin and gliadin to influence thegluten type of wheat. While for the structure protein, it regulated albumin or globulinbasically to suit the function of their metabolic activity.
2The effect of endogenous NO on senescence of tillers in leaves in wheat
2.1The adjustment of endogenous nitric oxide to senescence of tillers in leaves in wheat
The changes of NO content influenced senescence. NO content in main stem andprimary tiller was produced plentifully at heading and then fallen sharply at anthesis. It was atheading that the cell of main stem and primary tiller leaves going active apoptosis began toproduced the DNA fragmentation (except for the primary tiller under N3treatment). Inaddition, the shift induced by NO content changes accompanying with the aging of leaves.NO content in secondary tiller were much higher than main stem and primary tiller duringjointing to heading, and the earliest DNA degradation happened at secondary tiller sincebooting stage, when NO was found to accumulate rapidly. The present results had shown thatthe aging of leaves could be significantly influenced by NO, and it was also suggested thatsenescence were more sensitive relation to the marked changes of NO content than NOcontent itself.
2.2The adjustment of N supplying to both endogenous nitric oxide content and NR activity inwheat leaves
An apparent positive correlation existed in both cultivars leaves between NO content andNR activity among main stem, primary tiller and secondary tiller at early stage. Higher Napplications produced higher NR activity and higher NO content. With the progress ofgrowing period, it was weakened for the relationship between NR activity and NO content in leaves, or even with no correlation at heading stage. And with N application levels increasing,both NR activity and NO content reached the maximum and then decreased. There wassaturability for both the NR activity and the NO content in vivo as the N applicationincreasing.
2.3The adjustment of endogenous nitric oxide to both tillers senescence and yield and proteincontent
There was an increase in the survival of productive tillers with increase of nitrogen leveluntil N2treatment. Number of live tillers per plant in wheat increased to a peak of3.84(or1.94) for JN17(or TN18) at240-N treatment as more N application supplied. The maximumgrain yield reached at240-N application as the tillers increasing. Thus, grain production wasclosely related to the number of ears producing by primary tiller, which had much more effectupon by N-fertilizer. The aging happened later for primary tiller than main stem under thesame N level. Unlike the other N treatment, the360-N treatment resulted in later aging forprimary tiller till anthesis, however, the largest numbers of ears were not obtained under theN-360application level at maturity. Numbers of tillers died at anthesis. Too late dying fornumbers of ineffective primary tiller had consumed lots of nutrients which caused thereduction of grain yield comparing with240-N application. Nitrogen level influenced kernelprotein content. Compared with main stem, it was decreased by15.8to2.4%for primary tillerin JN17, and by15.7to3.7%in TN18, respectively. Supplied with more N-fertilizer, itbecame more less for the difference between main stem and primary tiller. Overall, these dataindicated that N-fertilizer regulated the ears number of primary tiller and reduced thedifference of protein content between main stem and primary tiller, in other words, it was aneffective ways to regulate the number and quality of primary tiller by N application in order tobalance the yield and uniformity of ears.
1外源一氧化氮对干旱胁迫下冬小麦叶片光合特性及籽粒蛋白质组成的调节
1.1外源一氧化氮对干旱胁迫下小麦叶片光合及叶绿素荧光特性的调控
不同时期外源NO处理显著提高干旱胁迫下叶片的光合速率，随着NO处理浓度的增加，光合速率表现出先升高、后降低的趋势，低浓度NO显著提高干旱胁迫下叶片的光合速率并使其一直保持较高水平；叶片的蒸腾速率（Tr）的影响因时期、品种和处理浓度不同而存在差异，总的来说，随着NO处理浓度的增加，表现为先升后降的趋势，但是，外源NO对不同小麦品种类型的影响存在差异，可以显著提高多穗型品种的蒸腾速率，而对大穗型品种作用不明显。低浓度NO对气孔导度表现为促进作用，高浓度则抑制，适量NO浓度处理对维持干旱胁迫下小麦叶片气孔开放具有积极意义。外源NO处理提高了干旱胁迫下叶片的瞬时水分利用效率，并在C2（0.4mmol L-1）浓度下达最大值，不同NO处理时期间影响效果不同，拔节期>抽穗期>开花期。外源NO提高叶片了叶绿素含量，但是对类胡萝卜素含量无显著影响。NO能提高小麦最大光化学效率（Fv/Fm），以拔节期喷施效果最显著。喷施NO处理增加了光合电子传递能量占总吸收光能的比例，提高了光能利用率。
1.2外源一氧化氮对干旱胁迫下不同基因型小麦籽粒蛋白组分的调控
干旱胁迫条件下，通过对4个不同筋型小麦籽粒蛋白质组成和谷蛋白大聚合体含量及粒度分布的研究显示，外源NO处理提高了小麦籽粒中的谷蛋白比例，而对清蛋白、球蛋白和醇溶蛋白的影响存在品种间差异，说明小麦籽粒谷蛋白的合成与积累受NO的调节。在外源NO作用下，更多的贮存蛋白以谷蛋白形式存在。NO处理可以增加强筋型品种的清蛋白含量，降低醇溶蛋白含量，对球蛋白调节效果不明显。外源NO增加了中筋型品种的球蛋白含量，降低清蛋白含量，醇溶蛋白则对NO调节表现不敏感。
2内源一氧化氮对冬小麦茎蘖位叶片衰老的调节
2.1内源NO含量对小麦不同茎蘖位叶片衰老进程的调节
NO含量的改变可以引起衰老。在主茎和一级分蘖中，叶片内NO含量在抽穗期达到最大值，然后在开花期迅速下降，与其叶片DNA片段化的发生时间一致。从拔节期到抽穗期，二级分蘖叶片中NO含量一直处于较高水平，二级分蘖叶片DNA降解发生时间最早。表明NO确实与叶片的衰老密切相关，而且NO含量的骤然改变可能是引起衰老的主要原因。
2.2施N水平对小麦叶片内源NO产生和NR活性的影响
拔节期和挑旗期，不同蘖位叶片内NO含量和NR活性都呈显著正相关关系，增施氮肥提高NR活性及NO含量，表明NO的产生主要依赖于叶片内的NR途径，并且氮肥对二者具有调节作用。随着生育时期进程，叶片内NR活性与NO的产生的相关性减弱，甚至在抽穗期二者无相关性，说明对于孕穗期后NO的产生，叶片内合成作用减弱。不同氮肥处理条件下，NO含量和NR活性都存在着饱和性，二者都是先升高，在达到最大值之后再降低。在抽穗期，叶片内NO含量和NR活性达较高水平。
2.3茎蘖位衰老进程对产量和蛋白质含量的影响
施氮量低于N-240处理时，增施氮肥可以获得更多的有效穗数。N-240处理条件下，济南17和泰农18的每个植株分别可以产生最多的分蘖数3.84个和1.94个，继续增施氮肥至N-360，穗数降低，济南17为3.19个分蘖，泰农18为1.69个。两品种在施氮量N-240处理时，产量最高，且产量的形成与一级分蘖的成穗率密切相关，后者受氮肥施入量的影响。N-360处理条件下，一级分蘖叶片的衰老较晚，直到开花期才开始，但是成熟后单位面积有效穗数少于N-240处理，说明在开花期有很多无效分蘖死亡，这些无效分蘖在消耗了大量养分后死亡，导致产量相对N-240处理下有所降低。施氮量影响籽粒中蛋白质含量，增施氮肥增加籽粒中的蛋白质含量。与主茎相比，一级分蘖籽粒中蛋白质含量显著减少，增加氮肥施入量，则减少主茎与一级分蘖间的含量差异。表明，通过控制氮肥施入量调节一级分蘖有效穗数及其籽粒品质的途径是可行的。
Nitric oxide (NO) is a key signaling molecule in different physiological processes ofplants, including adapting stress and senescence. The objective of this study was to determinethe effects of NO in wheat growing period. Field-experiments were conducted in2009-2011growing seasons used exogenous and endogenous two kinds of methods to certain:1.Theeffect of exogenous NO on photosynthetic characteristics, chlorophyll fluorescence in leavesand protein components, GMP content and particle size distribution in grains under droughtstress.2. The effect of endogenous NO on senescence of tillers in leaves in wheat. The firstexperiment was carried out in water stress condition which depending on raining entirely,with different concentrations on NO at different spraying time (jointing stage, heading stageand anthesis) was conducted. The second experiment was carried out under different nitrogenlevels using two wheat cultivars with different panicle-type. The main results were as follows:
1. The adjustment of exogenous nitric oxide to leaves photosynthetic characteristicsand grain protein composition under drought stress in winter wheat
1.1The adjustment of exogenous nitric oxide to leaves photosynthetic characteristics andChlorophyll fluorescence properties under drought stress in winter wheat
Exogenous NO improved the leaf photosynthetic rate under drought stress significantly,and with concentration increasing, photosynthetic rate rose at first and then declined. Lowconcentration NO increased leaf photosynthetic rate significantly and made it keep higherlevel. Transpiration rate was influenced by period, variety and NO concentration. In generallyspeaking, with increasing of concentration, it first increased and then trends dropped.However, exogenous treatment was not same at different wheat varieties. Low NOconcentration promoted the stomata conductance and high concentration was suppressed. Ithad a positive meaning for NO to maintain suitable stomata opening wheat under droughtstress. NO improved instantaneous water use efficiency of leaves under drought stress, andC2(0.4mmol L-1) improved significantly. Different effects showed as different periodprocessing, jointing stage>heading stage>anthesis. NO improved the chlorophyll content buthad no significantly effect on carotenoid content. It increased Fv/Fm most significantly spraying at jointing stage. Because of increasing the ratio of photosynthetic electron transferenergy to total absorption of light energy, the utilization rate of light energy improved.
1.2The adjustment of exogenous nitric oxide to protein composition under drought stress inwinter wheat
Under drought stress condition, we researched different wheat types for grain protein,GMP content and particle size distribution. It showed that, exogenous NO treatment improvedproportion in glutenin, but to albumin, globulin and gliadin, it influenced differently betweenthe varieties. More storage protein in the cereal was the form of glutenin with the NOspraying. NO increased the strong gluten varieties of albumin content type and reducedgliadin content but had no obviously effect on globulin. It increased in the weak glutencultivars type of globulin content, declined albumin content and was not sensitive to gliadin.In conclusion, it was known that NO adjusted the ratio of glutenin and gliadin to influence thegluten type of wheat. While for the structure protein, it regulated albumin or globulinbasically to suit the function of their metabolic activity.
2The effect of endogenous NO on senescence of tillers in leaves in wheat
2.1The adjustment of endogenous nitric oxide to senescence of tillers in leaves in wheat
The changes of NO content influenced senescence. NO content in main stem andprimary tiller was produced plentifully at heading and then fallen sharply at anthesis. It was atheading that the cell of main stem and primary tiller leaves going active apoptosis began toproduced the DNA fragmentation (except for the primary tiller under N3treatment). Inaddition, the shift induced by NO content changes accompanying with the aging of leaves.NO content in secondary tiller were much higher than main stem and primary tiller duringjointing to heading, and the earliest DNA degradation happened at secondary tiller sincebooting stage, when NO was found to accumulate rapidly. The present results had shown thatthe aging of leaves could be significantly influenced by NO, and it was also suggested thatsenescence were more sensitive relation to the marked changes of NO content than NOcontent itself.
2.2The adjustment of N supplying to both endogenous nitric oxide content and NR activity inwheat leaves
An apparent positive correlation existed in both cultivars leaves between NO content andNR activity among main stem, primary tiller and secondary tiller at early stage. Higher Napplications produced higher NR activity and higher NO content. With the progress ofgrowing period, it was weakened for the relationship between NR activity and NO content in leaves, or even with no correlation at heading stage. And with N application levels increasing,both NR activity and NO content reached the maximum and then decreased. There wassaturability for both the NR activity and the NO content in vivo as the N applicationincreasing.
2.3The adjustment of endogenous nitric oxide to both tillers senescence and yield and proteincontent
There was an increase in the survival of productive tillers with increase of nitrogen leveluntil N2treatment. Number of live tillers per plant in wheat increased to a peak of3.84(or1.94) for JN17(or TN18) at240-N treatment as more N application supplied. The maximumgrain yield reached at240-N application as the tillers increasing. Thus, grain production wasclosely related to the number of ears producing by primary tiller, which had much more effectupon by N-fertilizer. The aging happened later for primary tiller than main stem under thesame N level. Unlike the other N treatment, the360-N treatment resulted in later aging forprimary tiller till anthesis, however, the largest numbers of ears were not obtained under theN-360application level at maturity. Numbers of tillers died at anthesis. Too late dying fornumbers of ineffective primary tiller had consumed lots of nutrients which caused thereduction of grain yield comparing with240-N application. Nitrogen level influenced kernelprotein content. Compared with main stem, it was decreased by15.8to2.4%for primary tillerin JN17, and by15.7to3.7%in TN18, respectively. Supplied with more N-fertilizer, itbecame more less for the difference between main stem and primary tiller. Overall, these dataindicated that N-fertilizer regulated the ears number of primary tiller and reduced thedifference of protein content between main stem and primary tiller, in other words, it was aneffective ways to regulate the number and quality of primary tiller by N application in order tobalance the yield and uniformity of ears.
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