土壤质地与供氮水平对小麦产量和品质的影响及其生理基础
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摘要
在3种不同质地土壤条件下,对优质强筋小麦品种山农12不同粒位籽粒淀粉积累、淀粉粒度分布、籽粒内源激素水平及淀粉合成相关酶活性变化进行了研究,明确了不同粒位籽粒淀粉粒度分布及淀粉形成的酶学特征。同时对小麦不同粒位籽粒在蛋白质形成、HMW-GS表达量、GMP含量及粒度分布等方面的差异进行了研究,分析了HMW-GS积累与GMP粒度分布的关系。研究了在3种质地土壤上施氮量对小麦产量和品质的影响及其生理基础。主要研究结果如下:
1小麦不同粒位籽粒蛋白质积累差异
小麦不同粒位籽粒蛋白质含量和积累量均表现为2位粒>1位粒>4位粒>3位粒。不同粒位籽粒蛋白质各组分含量亦存在显著差异,上位粒(3、4位籽粒)球蛋白含量显著高于下位粒(1、2位籽粒),而醇溶蛋白和谷蛋白含量均显著低于上位粒,清蛋白含量上位粒和下位粒之间无显著差异。
本研究表明,小麦强势粒和弱势粒HMW-GS均在花后14 d开始形成,强势粒HMW-GS含量明显高于弱势粒,表明强势粒具有较强的HMW-GS积累能力。花后小麦HMW-GS各亚基含量变化与总亚基含量变化趋势一致。各亚基含量表现为5亚基>10亚基>14亚基>15亚基,强势粒各亚基含量在整个籽粒发育进程中亦明显高于弱势粒。在小麦籽粒发育进程中,GMP含量呈先上升后下降再上升的趋势,强势粒GMP含量均明显高于弱势粒。
本研究表明,小麦强、弱势籽粒GMP粒度分布趋势类似,粒径范围0.37~245μm之间。小麦GMP数目主要由<10μm颗粒组成(约占99.8%以上),体积分布的47.57%~69%集中在10-100μm,而表面积分布大部分(66.92%~83.23%)由<10μm颗粒组成。强势粒10-100μm和>100μm颗粒数目比例均显著高于弱势粒;而强势粒>100μm颗粒所占体积比例亦显著高于弱势粒。说明强势粒含有较多的大粒径GMP颗粒。这可能是因为强势粒GMP发育时间早,更多的小粒径GMP颗粒发育成为大粒径GMP颗粒。HMW-GS积累和GMP粒度分布的差异可能是小麦强、弱势籽粒品质差异的重要原因。
2小麦不同粒位籽粒淀粉积累差异
成熟期小麦不同粒位籽粒直链淀粉含量随粒位升高而增加;支链淀粉含量和总淀粉含量亦表现为3、4位粒高于1、2位粒。但由于受粒重影响,直链淀粉积累量、支链淀粉积累量和总淀粉积累量均表现为2位粒>1位粒>3位粒>4位粒。不同粒位的直/支比表现为4位粒>3位粒>2位粒>1位粒,这可能与淀粉组分合成差异有关。本研究表明,花后7~21 d,弱势粒的蔗糖含量显著高于强势粒,而此时期恰与淀粉积累速率高峰出现时间相吻合。弱势粒之所以淀粉积累量低、积累速率慢,与淀粉合成能力低有关,而淀粉合成所需同化物的供给并不是主要限制因子。本研究表明,小麦籽粒淀粉合成的关键酶AGPase、UGPase、SSS和GBSS活性与强势粒淀粉积累速率均呈显著或极显著正相关,强势粒上述酶活性均高于弱势粒,进一步表明淀粉合成相关酶活性较高、淀粉合成能力强是强势粒淀粉积累量高的重要原因。Logistic方程模拟也表明,强势粒淀粉积累起始势(C0)高,活跃持续期长,平均积累速率(Rmean)高,最终淀粉积累量高。
3土壤质地对小麦籽粒品质形成的影响
本试验条件下,3种质地土壤上小麦HMW-GS含量和GMP含量均表现为粘壤土>中壤土>砂壤土,表明粘壤土有利于小麦HMW-GS积累,有利于改善小麦品质。3种质地土壤比较,粘壤土小麦具有较高的沉降值和湿面筋含量,面团形成时间和稳定时间均较高,表明粘壤土小麦具有较好的面团流变学特性,面团品质好。3种质地土壤上小麦GMP颗粒体积分布为双峰曲线。GMP<10μm和10-100μm颗粒所占体积比例均表现为砂壤土>中壤土>粘壤土,而>100μm颗粒对体积的贡献表现为粘壤土>中壤土>砂壤土。3种质地土壤上小麦GMP颗粒数目分布为单峰曲线,粘壤土小麦强势粒中GMP>100μm颗粒数目比例较高。3种质地土壤上小麦GMP颗粒表面积分布为双峰曲线,粘壤土小麦GMP<10μm颗粒所占表面积显著低于砂壤土和中壤土,而10-100μm和>100μm颗粒所占表面积均显著高于砂壤土和中壤土。可见,粘壤土有利于小麦GMP的发育,大体积GMP颗粒所占体积均高于中壤土和砂壤土,有利于改善小麦品质。此外,小麦强势粒与弱势粒HMW-GS积累在砂壤土上差异较大,而在中壤土和粘壤土上二者差异较小;与强势粒相比,弱势粒HMW-GS积累不同质地土壤上差异较大。表明弱势粒HMW-GS积累更易受土壤条件的影响,生产上应注重对弱势粒进行调控以改善籽粒品质。
本研究结果表明,3种质地土壤上小麦粒重比较,中壤土最高,粘壤土次之,砂壤土最低。砂壤土小麦灌浆起始势较高,但最大积累积累速率和平均积累速率均较低,最终粒重低;中壤土小麦具有较高的灌浆起始势和积累速率,因而最终粒重较高。可见,中壤土水气热状况适宜,肥力相对较高,有利于维持小麦较高的灌浆速率和灌浆持续期,有利于获得高产。
砂壤土上小麦支链淀粉含量最高,直/支最低,总淀粉含量最高,但因其粒重较小,总淀粉积累量要小于中壤土。粘壤土上小麦直链淀粉含量较高,直/支较高,总淀粉含量最低,总淀粉积累量也最低。表明砂壤土有利于小麦淀粉的积累,但产量较低。不同土壤质地间水气状况和通气性不同,造成籽粒内源激素水平和淀粉合成相关酶活性的差异可能是不同质地土壤上淀粉组分积累差异的重要原因。不同质地土壤小麦淀粉粒度分布特征存在明显差异。砂壤土小麦B型淀粉粒较多,而A型淀粉粒较少;粘壤土小麦恰好相反。表明粘壤土有利于小麦A型淀粉粒体积比例的增加。3种质地土壤上小麦淀粉数目各粒径范围内无显著差异。3种质地土壤比较,粘壤土小麦A型淀粉粒占总表面积的比例较高。可以看出,粘壤土有利于小麦A型淀粉粒体积和表面积比例的增加,这可能是因为粘壤土促进了小麦淀粉粒发育的缘故,这也与粘壤土小麦直链淀粉含量较高相吻合。
4土壤质地影响小麦品质的生理基础
3种质地土壤上施氮均不同程度提高了小麦穗数、穗粒数和粒重,进而提高了小麦产量。与N0相比,N1和N2处理分别增产15.77%和23.22%(砂壤土);10.98%和18.00%(中壤土);5.96%和19.34%(粘壤土)。3种质地土壤比较,砂壤土小麦产量最低,但施氮增产幅度最大。小麦穗粒数3种质地土壤无显著差异,而穗数和粒重均以中壤土最高。中壤土通气性较好,且保水性和保肥性较强,有利于小麦穗数的增加和粒重的提高,最终获得较高的籽粒产量。
本研究发现,3种质地土壤上开花期植株营养器官氮素积累量和花前贮存氮素运转量均随施氮量增加而增加,3种质地土壤上都以N2处理最大。同时,施氮显著降低了花前贮存氮素运转率,施氮处理间无显著差异。3种质地土壤比较,砂壤土上施氮对小麦花前氮素运转量的影响较大(平均增加56.26%),而中壤土(平均增加27.43%)和粘壤土(平均增加3.58%)各处理间差异较小,说明砂壤土施氮对小麦氮素运转有更显著的调控效应。3种质地土壤上适量施氮均能提高旗叶内肽酶和羧肽酶活性,促进旗叶可溶性蛋白质的降解,但施氮量过多不利于内肽酶活性的提高。与砂壤土相比,中壤土和粘壤土小麦灌浆后期旗叶内肽酶和羧肽酶活性下降速率较慢。酶活性高且高值持续时间长,有利于保持较高的氮素转移效率,从而提高籽粒蛋白质含量。
本研究表明,小麦开花期0-20cm土层根系鲜重砂壤土最低,粘壤土次之,中壤土最高;20-40cm土层砂壤土最高。这可能是因为砂壤土比较疏松,有利于根系的下扎,增大了下层根系的比例。开花10d后各土层根系鲜重均表现为粘壤土>中壤土>砂壤土。这可能是因为粘壤土有利于小麦根系生长并延缓其衰老的缘故。3种质地土壤比较,0-20cm土层小麦根系活力均表现为中壤土>粘壤土>砂壤土;20-40cm土层均表现为粘壤土>中壤土>砂壤土。表明中壤土和粘壤土有利于延缓小麦根系的衰老。
3种质地土壤上施氮均明显提高了根系SOD活性,有利于及时清除活性氧。3种质地土壤比较,中壤土和粘壤土小麦根系生育后期仍能保持较高的SOD活性,根系MDA含量低,表明中壤土和粘壤土小麦根系膜脂过氧化程度低,根系衰老慢。3种质地土壤上施氮均能明显提高小麦旗叶SOD、POD、CAT活性,降低MDA含量,表明施氮有利于提高小麦旗叶活性氧清除能力,延缓衰老。3种质地土壤比较,砂壤土小麦旗叶SOD活性前期升高快,后期迅速下降,MDA含量增加快,小麦膜脂过氧化现象严重。砂壤土上施氮旗叶MDA含量降低幅度较大,表明砂壤土小麦旗叶活性氧代谢更易受到氮肥的调控。因此,在砂壤土上种植小麦可以通过合理施氮来延缓衰老,以获得优质高产。
The high quality wheat cultivar shannong 12 was used in this experiment under three different soil texture conditions, in order to study starch accumulation characteristics, starch particle size distribution, endogenous hormone levels and enzyme activities related to starch synthesis of different grain position. The protein formation, HMW-GS expression, GMP content and size distribution of GMP particle were studied at the same time, and the relationship between HMW-GS accumulation and GMP size distribution was analyzed. The effects of nitrogen rate on yield and quality under different soil textures and its physiological basis were studied in the experiment. Key findings are as follows:
1 Protein accumulation differences at different grain positions
The protein content and accumulation amount of wheat grain at different position both showed second position grain> first position grain> forth position grain> third position grain. The content of protein components of wheat at different grain positions were different significantly. The upper grains (3,4-Grains) have higher globulin content and lower gliadin content and glutenin content than lower grains (1,2-Grains).There are no significant difference of albumin content between upper grains and lower grains.
This study shows that the HMW-GS appeared at 14 days after anthesis of wheat both in superior grain and inferior grain.The HMW-GS content of superior grain was higher than that in superior grain, which indicated that the accumulation ability of HMW-GS was stronger in superior grain. After anthesis, each HMW-GS content changed in the same trend as total HMW-GS content. The content of 5 subunits> 10 subunits> 14 subunits> 15 subunits, each subunits content in superior grain was also higher than inferior grain. The GMP content raised first and then fell down in the process of grain filling.The GMP content in superior gain was also higher than inferior grain. Lower HMW-GS accumulation amount and GMP content maybe one of the reasons for poor quality of inferior grain.
This study showed that, the GMP size distribution of wheat was similar in superior grain and inferior grain, the particle diameter ranged for 0.37~245μm. The number of GMP particle mainly composed by <10μm particle (accounting for 99.8%), volume distribution percentage focus on 10-100μm (47.57%~69%), surface area mainly composed by <10μm particle (66.92%~83.23%). The number percentage of 10~100μm and >100μm particle of GMP in superior grain were higher than inferior grain.The volume percentage of >100μm particle of GMP in superior grain were also higher than inferior grain. So superior grain contains more large size GMP particles. This maybe because more small size particles became large size particles in superior grain. The difference of HMW-GS accumulation and GMP size distribution maybe one of the reasons of quality difference between superior grain and inferior grain.
2 Starch accumulation differences at different grain positions
The amylose content of wheat grain at mature increased with the moving up of grain positions,amylopectin content and total starch content of third position grain and forth position grain were higher than that of first position grain and second position grain. The accumulation of amylose and amylopectin and total starch showed second position grain>first position grain>third position grain>forth position grain because of the weight impact. The amylose/amylopectin ratio showed firth position >third position grain>second position grain>first position grain, which may be related to the difference of starch component synthesis.
This study showed that, 7-21days after anthesis is the vigorous period of sucrose consumption, which matched with time of starch accumulation rate.The sucrose content in inferior grain was higher than that in superior grain in this period. It is indicated that the substrate of starch synthesis should not be the limiting factor for starch accumulation in inferior grain, which related to starch synthesis efficiency. This study showed that, the activities of SS, AGPase, UGPase, SSS, and GBSS changed in the pattern of a single-peak curve during grain filling. The related enzyme activities in superior grain, which had higher starch accumulation, were higher than those in inferior grain. The simulation with Logistic equation showed that the accumulation rate and initial potential were higher, the accumulation duration was longer, so the starch final amount was higher in superior grain.
3 Effects of different soil textures on quality formation of wheat
In this experiment, HMW-GS content and GMP content of wheat under different soil textures showed clay soil> loam soil > sandy soil, which indicated that clay soil was favorable for HMW-GS accumulation and will help to improve wheat quality. The sedimentation volume, wet gulten content and dough development time and dough stability time were highest of wheat from clay soil. That is to say, wheat in clay soil has good dough quality. The results showed that the volume distribution of GMP particle of wheat was two peak curve. The percentage of <10μm and 10~100μm GMP particle volume distribution in sandy soil was highest, while in clay soil was lowest. However, the percentage of >100μm GMP particle volume distribution in sandy soil was lowest, while in clay soil was highest. The number distribution of GMP particle of wheat was single peak curve. Clay soil wheat has higher number percentage of >100μm GMP particle. The surface area distribution of GMP particle of wheat was two peak curve. The surface area percentage of <10μm GMP particle was lower, while 10~100μm and >100μm GMP particle was higher in clay soil. So clay soil was favourble for GMP development and improving of wheat quality. In addition, the difference of HMW-GS content between superior grain and inferior grain was larger under sandy soil, while smaller under loam soil and clay soil. It indicated that the HMW-GS accumulation in inferior grain maybe easily affected by soil conditions. More attention should be paid to the regulation of inferior grain to improve grain quality in wheat production.
The results showed that the grain weight of loam soil wheat was larger than sandy soil wheat and loam soil wheat. Sandy soil wheat has higher amylopectin content and higher starch content than loam soil wheat and clay soil wheat but lower starch accumulation amount because of lower grain weight. Clay soil wheat has higher amylose content and lower total starch content. Sandy soil is favorable for starch accumulation, but the starch yield was lower. The starch granule distribution were different among wheat under different texture soils. Wheat in sandy soil has more B-type starch granule and less A-type starch granule. There is no significant difference of starch granule number distribution of wheat among different soil textures. It indicated that clay soil was farvourble for the increase of volume and surface area percentage of A-type granule, because clay soil promoted the development of starch granule possibly, which also consistent with higher amylose of clay soil wheat.
4 Physiological basis for effects of different soil texture on wheat quality
Nitrogen application increased spike number, kernel number and grain weight under three soil textures, thus increased the yield of wheat. Compared with N0 treatment, the yield increased 15.77% in N1 treatment and 23.22% in N2 treatment respectively under sandy soil, and 10.98% and 18.00% respectively under loam soil, and 5.96% and 19.34% respectively under loam soil. The yield was lowest in sandy soil, but there were largest yield increase of nitrogen application among three soil textures. There was no significant difference of kernel number among three soil textures, but the spike number and grain weight were highest in loam soil. Loam soil has better conditions of air, water and fertilizer, was propitious to increase spike number and kernel weight.
This study found that nitrogen accumulation and translocation increased with increasing of nitrogen rate. The nitrogen translation rate was decreased by nitrogen application, but there is no difference among nitrogen treatment. Effect of nitrogen application on nitrogen translocation was higher (an average increase of 56.26%) in sandy soil, while that was lower in loam soil (an average increase of 27.43%) and clay soil (an average increase of 3.58%). Appropriate nitrogen application increased endopeptidases activities and carboxypetidase activities and promoted the decomposition of soluble protein. The endopeptidases activities and carboxypetidase activities of wheat flag leaf in loam soil and clay soil were higher and could keep a long period, which help to keep higher nitrogen translation rate and increase protein content of wheat grain.
This study showed that sandy soil wheat has highest root fresh weight in 0-20 cm soil layer while lowest in 20-40 cm soil layer at anthesis. This may be due to loose texture of sandy soil, which increased the proportion of the lower roots. The root fresh weight showed clay soil>loam soil >sandy soil 10 days after the flowering. Because clay soil promoted root growth and delayed the senescence of wheat. The root vigor in 0-20 cm layer soil showed loam soil >clay soil>sandy soil, and clay soil >loam soil>sandy soil in 20-40cm layer soil. It indicated that loam soil and clay soil could help to delay senescence of wheat root.
Nitrogen application increased SOD activities of wheat root, and was conducive to remove reactive oxygen species timely. The SOD activities of wheat root in sandy soil were higher than that in loam soil and clay soil. Nitrogen application increased SOD, POD, CAT activities and decreased MDA content of wheat flag leaf, which showed that nitrogen application was conducive to raise wheat active oxygen scavenging ability. The effect of nitrogen application on MDA content of wheat flag leaf was larger in sandy soil. So the active oxygen metaboism of flag leaf in sandy soil wheat was easy to be regulated by nitrogen fertilizer. Therefore, the wheat cultivation in sandy soil can be regulated by reasonable nitrogen application to delay senescence, to obtain high quality and high yield.
1小麦不同粒位籽粒蛋白质积累差异
小麦不同粒位籽粒蛋白质含量和积累量均表现为2位粒>1位粒>4位粒>3位粒。不同粒位籽粒蛋白质各组分含量亦存在显著差异,上位粒(3、4位籽粒)球蛋白含量显著高于下位粒(1、2位籽粒),而醇溶蛋白和谷蛋白含量均显著低于上位粒,清蛋白含量上位粒和下位粒之间无显著差异。
本研究表明,小麦强势粒和弱势粒HMW-GS均在花后14 d开始形成,强势粒HMW-GS含量明显高于弱势粒,表明强势粒具有较强的HMW-GS积累能力。花后小麦HMW-GS各亚基含量变化与总亚基含量变化趋势一致。各亚基含量表现为5亚基>10亚基>14亚基>15亚基,强势粒各亚基含量在整个籽粒发育进程中亦明显高于弱势粒。在小麦籽粒发育进程中,GMP含量呈先上升后下降再上升的趋势,强势粒GMP含量均明显高于弱势粒。
本研究表明,小麦强、弱势籽粒GMP粒度分布趋势类似,粒径范围0.37~245μm之间。小麦GMP数目主要由<10μm颗粒组成(约占99.8%以上),体积分布的47.57%~69%集中在10-100μm,而表面积分布大部分(66.92%~83.23%)由<10μm颗粒组成。强势粒10-100μm和>100μm颗粒数目比例均显著高于弱势粒;而强势粒>100μm颗粒所占体积比例亦显著高于弱势粒。说明强势粒含有较多的大粒径GMP颗粒。这可能是因为强势粒GMP发育时间早,更多的小粒径GMP颗粒发育成为大粒径GMP颗粒。HMW-GS积累和GMP粒度分布的差异可能是小麦强、弱势籽粒品质差异的重要原因。
2小麦不同粒位籽粒淀粉积累差异
成熟期小麦不同粒位籽粒直链淀粉含量随粒位升高而增加;支链淀粉含量和总淀粉含量亦表现为3、4位粒高于1、2位粒。但由于受粒重影响,直链淀粉积累量、支链淀粉积累量和总淀粉积累量均表现为2位粒>1位粒>3位粒>4位粒。不同粒位的直/支比表现为4位粒>3位粒>2位粒>1位粒,这可能与淀粉组分合成差异有关。本研究表明,花后7~21 d,弱势粒的蔗糖含量显著高于强势粒,而此时期恰与淀粉积累速率高峰出现时间相吻合。弱势粒之所以淀粉积累量低、积累速率慢,与淀粉合成能力低有关,而淀粉合成所需同化物的供给并不是主要限制因子。本研究表明,小麦籽粒淀粉合成的关键酶AGPase、UGPase、SSS和GBSS活性与强势粒淀粉积累速率均呈显著或极显著正相关,强势粒上述酶活性均高于弱势粒,进一步表明淀粉合成相关酶活性较高、淀粉合成能力强是强势粒淀粉积累量高的重要原因。Logistic方程模拟也表明,强势粒淀粉积累起始势(C0)高,活跃持续期长,平均积累速率(Rmean)高,最终淀粉积累量高。
3土壤质地对小麦籽粒品质形成的影响
本试验条件下,3种质地土壤上小麦HMW-GS含量和GMP含量均表现为粘壤土>中壤土>砂壤土,表明粘壤土有利于小麦HMW-GS积累,有利于改善小麦品质。3种质地土壤比较,粘壤土小麦具有较高的沉降值和湿面筋含量,面团形成时间和稳定时间均较高,表明粘壤土小麦具有较好的面团流变学特性,面团品质好。3种质地土壤上小麦GMP颗粒体积分布为双峰曲线。GMP<10μm和10-100μm颗粒所占体积比例均表现为砂壤土>中壤土>粘壤土,而>100μm颗粒对体积的贡献表现为粘壤土>中壤土>砂壤土。3种质地土壤上小麦GMP颗粒数目分布为单峰曲线,粘壤土小麦强势粒中GMP>100μm颗粒数目比例较高。3种质地土壤上小麦GMP颗粒表面积分布为双峰曲线,粘壤土小麦GMP<10μm颗粒所占表面积显著低于砂壤土和中壤土,而10-100μm和>100μm颗粒所占表面积均显著高于砂壤土和中壤土。可见,粘壤土有利于小麦GMP的发育,大体积GMP颗粒所占体积均高于中壤土和砂壤土,有利于改善小麦品质。此外,小麦强势粒与弱势粒HMW-GS积累在砂壤土上差异较大,而在中壤土和粘壤土上二者差异较小;与强势粒相比,弱势粒HMW-GS积累不同质地土壤上差异较大。表明弱势粒HMW-GS积累更易受土壤条件的影响,生产上应注重对弱势粒进行调控以改善籽粒品质。
本研究结果表明,3种质地土壤上小麦粒重比较,中壤土最高,粘壤土次之,砂壤土最低。砂壤土小麦灌浆起始势较高,但最大积累积累速率和平均积累速率均较低,最终粒重低;中壤土小麦具有较高的灌浆起始势和积累速率,因而最终粒重较高。可见,中壤土水气热状况适宜,肥力相对较高,有利于维持小麦较高的灌浆速率和灌浆持续期,有利于获得高产。
砂壤土上小麦支链淀粉含量最高,直/支最低,总淀粉含量最高,但因其粒重较小,总淀粉积累量要小于中壤土。粘壤土上小麦直链淀粉含量较高,直/支较高,总淀粉含量最低,总淀粉积累量也最低。表明砂壤土有利于小麦淀粉的积累,但产量较低。不同土壤质地间水气状况和通气性不同,造成籽粒内源激素水平和淀粉合成相关酶活性的差异可能是不同质地土壤上淀粉组分积累差异的重要原因。不同质地土壤小麦淀粉粒度分布特征存在明显差异。砂壤土小麦B型淀粉粒较多,而A型淀粉粒较少;粘壤土小麦恰好相反。表明粘壤土有利于小麦A型淀粉粒体积比例的增加。3种质地土壤上小麦淀粉数目各粒径范围内无显著差异。3种质地土壤比较,粘壤土小麦A型淀粉粒占总表面积的比例较高。可以看出,粘壤土有利于小麦A型淀粉粒体积和表面积比例的增加,这可能是因为粘壤土促进了小麦淀粉粒发育的缘故,这也与粘壤土小麦直链淀粉含量较高相吻合。
4土壤质地影响小麦品质的生理基础
3种质地土壤上施氮均不同程度提高了小麦穗数、穗粒数和粒重,进而提高了小麦产量。与N0相比,N1和N2处理分别增产15.77%和23.22%(砂壤土);10.98%和18.00%(中壤土);5.96%和19.34%(粘壤土)。3种质地土壤比较,砂壤土小麦产量最低,但施氮增产幅度最大。小麦穗粒数3种质地土壤无显著差异,而穗数和粒重均以中壤土最高。中壤土通气性较好,且保水性和保肥性较强,有利于小麦穗数的增加和粒重的提高,最终获得较高的籽粒产量。
本研究发现,3种质地土壤上开花期植株营养器官氮素积累量和花前贮存氮素运转量均随施氮量增加而增加,3种质地土壤上都以N2处理最大。同时,施氮显著降低了花前贮存氮素运转率,施氮处理间无显著差异。3种质地土壤比较,砂壤土上施氮对小麦花前氮素运转量的影响较大(平均增加56.26%),而中壤土(平均增加27.43%)和粘壤土(平均增加3.58%)各处理间差异较小,说明砂壤土施氮对小麦氮素运转有更显著的调控效应。3种质地土壤上适量施氮均能提高旗叶内肽酶和羧肽酶活性,促进旗叶可溶性蛋白质的降解,但施氮量过多不利于内肽酶活性的提高。与砂壤土相比,中壤土和粘壤土小麦灌浆后期旗叶内肽酶和羧肽酶活性下降速率较慢。酶活性高且高值持续时间长,有利于保持较高的氮素转移效率,从而提高籽粒蛋白质含量。
本研究表明,小麦开花期0-20cm土层根系鲜重砂壤土最低,粘壤土次之,中壤土最高;20-40cm土层砂壤土最高。这可能是因为砂壤土比较疏松,有利于根系的下扎,增大了下层根系的比例。开花10d后各土层根系鲜重均表现为粘壤土>中壤土>砂壤土。这可能是因为粘壤土有利于小麦根系生长并延缓其衰老的缘故。3种质地土壤比较,0-20cm土层小麦根系活力均表现为中壤土>粘壤土>砂壤土;20-40cm土层均表现为粘壤土>中壤土>砂壤土。表明中壤土和粘壤土有利于延缓小麦根系的衰老。
3种质地土壤上施氮均明显提高了根系SOD活性,有利于及时清除活性氧。3种质地土壤比较,中壤土和粘壤土小麦根系生育后期仍能保持较高的SOD活性,根系MDA含量低,表明中壤土和粘壤土小麦根系膜脂过氧化程度低,根系衰老慢。3种质地土壤上施氮均能明显提高小麦旗叶SOD、POD、CAT活性,降低MDA含量,表明施氮有利于提高小麦旗叶活性氧清除能力,延缓衰老。3种质地土壤比较,砂壤土小麦旗叶SOD活性前期升高快,后期迅速下降,MDA含量增加快,小麦膜脂过氧化现象严重。砂壤土上施氮旗叶MDA含量降低幅度较大,表明砂壤土小麦旗叶活性氧代谢更易受到氮肥的调控。因此,在砂壤土上种植小麦可以通过合理施氮来延缓衰老,以获得优质高产。
The high quality wheat cultivar shannong 12 was used in this experiment under three different soil texture conditions, in order to study starch accumulation characteristics, starch particle size distribution, endogenous hormone levels and enzyme activities related to starch synthesis of different grain position. The protein formation, HMW-GS expression, GMP content and size distribution of GMP particle were studied at the same time, and the relationship between HMW-GS accumulation and GMP size distribution was analyzed. The effects of nitrogen rate on yield and quality under different soil textures and its physiological basis were studied in the experiment. Key findings are as follows:
1 Protein accumulation differences at different grain positions
The protein content and accumulation amount of wheat grain at different position both showed second position grain> first position grain> forth position grain> third position grain. The content of protein components of wheat at different grain positions were different significantly. The upper grains (3,4-Grains) have higher globulin content and lower gliadin content and glutenin content than lower grains (1,2-Grains).There are no significant difference of albumin content between upper grains and lower grains.
This study shows that the HMW-GS appeared at 14 days after anthesis of wheat both in superior grain and inferior grain.The HMW-GS content of superior grain was higher than that in superior grain, which indicated that the accumulation ability of HMW-GS was stronger in superior grain. After anthesis, each HMW-GS content changed in the same trend as total HMW-GS content. The content of 5 subunits> 10 subunits> 14 subunits> 15 subunits, each subunits content in superior grain was also higher than inferior grain. The GMP content raised first and then fell down in the process of grain filling.The GMP content in superior gain was also higher than inferior grain. Lower HMW-GS accumulation amount and GMP content maybe one of the reasons for poor quality of inferior grain.
This study showed that, the GMP size distribution of wheat was similar in superior grain and inferior grain, the particle diameter ranged for 0.37~245μm. The number of GMP particle mainly composed by <10μm particle (accounting for 99.8%), volume distribution percentage focus on 10-100μm (47.57%~69%), surface area mainly composed by <10μm particle (66.92%~83.23%). The number percentage of 10~100μm and >100μm particle of GMP in superior grain were higher than inferior grain.The volume percentage of >100μm particle of GMP in superior grain were also higher than inferior grain. So superior grain contains more large size GMP particles. This maybe because more small size particles became large size particles in superior grain. The difference of HMW-GS accumulation and GMP size distribution maybe one of the reasons of quality difference between superior grain and inferior grain.
2 Starch accumulation differences at different grain positions
The amylose content of wheat grain at mature increased with the moving up of grain positions,amylopectin content and total starch content of third position grain and forth position grain were higher than that of first position grain and second position grain. The accumulation of amylose and amylopectin and total starch showed second position grain>first position grain>third position grain>forth position grain because of the weight impact. The amylose/amylopectin ratio showed firth position >third position grain>second position grain>first position grain, which may be related to the difference of starch component synthesis.
This study showed that, 7-21days after anthesis is the vigorous period of sucrose consumption, which matched with time of starch accumulation rate.The sucrose content in inferior grain was higher than that in superior grain in this period. It is indicated that the substrate of starch synthesis should not be the limiting factor for starch accumulation in inferior grain, which related to starch synthesis efficiency. This study showed that, the activities of SS, AGPase, UGPase, SSS, and GBSS changed in the pattern of a single-peak curve during grain filling. The related enzyme activities in superior grain, which had higher starch accumulation, were higher than those in inferior grain. The simulation with Logistic equation showed that the accumulation rate and initial potential were higher, the accumulation duration was longer, so the starch final amount was higher in superior grain.
3 Effects of different soil textures on quality formation of wheat
In this experiment, HMW-GS content and GMP content of wheat under different soil textures showed clay soil> loam soil > sandy soil, which indicated that clay soil was favorable for HMW-GS accumulation and will help to improve wheat quality. The sedimentation volume, wet gulten content and dough development time and dough stability time were highest of wheat from clay soil. That is to say, wheat in clay soil has good dough quality. The results showed that the volume distribution of GMP particle of wheat was two peak curve. The percentage of <10μm and 10~100μm GMP particle volume distribution in sandy soil was highest, while in clay soil was lowest. However, the percentage of >100μm GMP particle volume distribution in sandy soil was lowest, while in clay soil was highest. The number distribution of GMP particle of wheat was single peak curve. Clay soil wheat has higher number percentage of >100μm GMP particle. The surface area distribution of GMP particle of wheat was two peak curve. The surface area percentage of <10μm GMP particle was lower, while 10~100μm and >100μm GMP particle was higher in clay soil. So clay soil was favourble for GMP development and improving of wheat quality. In addition, the difference of HMW-GS content between superior grain and inferior grain was larger under sandy soil, while smaller under loam soil and clay soil. It indicated that the HMW-GS accumulation in inferior grain maybe easily affected by soil conditions. More attention should be paid to the regulation of inferior grain to improve grain quality in wheat production.
The results showed that the grain weight of loam soil wheat was larger than sandy soil wheat and loam soil wheat. Sandy soil wheat has higher amylopectin content and higher starch content than loam soil wheat and clay soil wheat but lower starch accumulation amount because of lower grain weight. Clay soil wheat has higher amylose content and lower total starch content. Sandy soil is favorable for starch accumulation, but the starch yield was lower. The starch granule distribution were different among wheat under different texture soils. Wheat in sandy soil has more B-type starch granule and less A-type starch granule. There is no significant difference of starch granule number distribution of wheat among different soil textures. It indicated that clay soil was farvourble for the increase of volume and surface area percentage of A-type granule, because clay soil promoted the development of starch granule possibly, which also consistent with higher amylose of clay soil wheat.
4 Physiological basis for effects of different soil texture on wheat quality
Nitrogen application increased spike number, kernel number and grain weight under three soil textures, thus increased the yield of wheat. Compared with N0 treatment, the yield increased 15.77% in N1 treatment and 23.22% in N2 treatment respectively under sandy soil, and 10.98% and 18.00% respectively under loam soil, and 5.96% and 19.34% respectively under loam soil. The yield was lowest in sandy soil, but there were largest yield increase of nitrogen application among three soil textures. There was no significant difference of kernel number among three soil textures, but the spike number and grain weight were highest in loam soil. Loam soil has better conditions of air, water and fertilizer, was propitious to increase spike number and kernel weight.
This study found that nitrogen accumulation and translocation increased with increasing of nitrogen rate. The nitrogen translation rate was decreased by nitrogen application, but there is no difference among nitrogen treatment. Effect of nitrogen application on nitrogen translocation was higher (an average increase of 56.26%) in sandy soil, while that was lower in loam soil (an average increase of 27.43%) and clay soil (an average increase of 3.58%). Appropriate nitrogen application increased endopeptidases activities and carboxypetidase activities and promoted the decomposition of soluble protein. The endopeptidases activities and carboxypetidase activities of wheat flag leaf in loam soil and clay soil were higher and could keep a long period, which help to keep higher nitrogen translation rate and increase protein content of wheat grain.
This study showed that sandy soil wheat has highest root fresh weight in 0-20 cm soil layer while lowest in 20-40 cm soil layer at anthesis. This may be due to loose texture of sandy soil, which increased the proportion of the lower roots. The root fresh weight showed clay soil>loam soil >sandy soil 10 days after the flowering. Because clay soil promoted root growth and delayed the senescence of wheat. The root vigor in 0-20 cm layer soil showed loam soil >clay soil>sandy soil, and clay soil >loam soil>sandy soil in 20-40cm layer soil. It indicated that loam soil and clay soil could help to delay senescence of wheat root.
Nitrogen application increased SOD activities of wheat root, and was conducive to remove reactive oxygen species timely. The SOD activities of wheat root in sandy soil were higher than that in loam soil and clay soil. Nitrogen application increased SOD, POD, CAT activities and decreased MDA content of wheat flag leaf, which showed that nitrogen application was conducive to raise wheat active oxygen scavenging ability. The effect of nitrogen application on MDA content of wheat flag leaf was larger in sandy soil. So the active oxygen metaboism of flag leaf in sandy soil wheat was easy to be regulated by nitrogen fertilizer. Therefore, the wheat cultivation in sandy soil can be regulated by reasonable nitrogen application to delay senescence, to obtain high quality and high yield.
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