杂种小麦及亲本旗叶老化过程中RubisCO特性的研究

小麦（ＴｒｉｔｉｃｕｍａｅｓｔｉｖｕｍＬ．）旗叶的ＲｕＢＰｃａｓｅ活性、含量及ＲｕＢＰｏａｓｅ活性在旗叶全展或全展后１０ｄ达最大值,以后逐渐下降。与亲本相比,供试杂种小麦“麦优４号”在旗叶一生中尤其老化后期上述参数皆表现明显的杂种优势。旗叶ＲｕＢＰｃａｓｅ比活性在叶绿素缓降期保持平稳,在叶绿素速降期逐渐下降。供试杂种小麦较亲本具有较高的ＲｕＢＰ羧化酶和加氧酶活性,表明杂种小麦不仅具有较强的光合羧化作用,而且叶片光合作用过程中的光呼吸也较强。结果与旗叶ＲｕｂｉｓＣＯ亲合ＣＯ２和Ｏ２的动力学常数的测定结果相符。     

叶部激素变化与小麦的整体衰老

旗叶中的叶绿素含量在抽穗—开花期达其最大值 ,然后迅速下降 ;丙二醛的含量逐渐上升 ;SOD活性逐渐下降。脱落酸的含量在灌浆的初期逐渐升高 ,中期达其最高值 ;玉米素在开花期达其峰值后迅速下降 ,旗叶的衰老与叶片中二激素间的浓度变化、平衡有关 ;根系活力在拔节—孕穗期达其高峰值 ,以后逐渐下降 ;根系活力在生育中、后期的下降与叶片中玉米素的下降、脱落酸的上升有直接关系 ,根系的早衰可能是引发小麦地上部衰老的重要原因之一。     

黑暗诱导衰老对不同年代冬小麦品种旗叶光系统Ⅱ功能的影响

小麦品种的更新换代是小麦产量不断提高的重要因素,阐明小麦品种演替过程中不同生理特性的变化对新品种选育具有重要参考价值.旗叶衰老速率快慢是影响小麦产量水平的关键因素,目前对于不同小麦品种衰老过程中旗叶光系统Ⅱ功能的变化规律尚不清楚.本试验选用1941-2014年间河南地区不同时期种植的31个品种,通过黑暗诱导离体叶片衰老,测定旗叶叶绿素荧光诱导动力学参数、叶绿素相对含量的变化,分析了光系统Ⅱ功能的变化规律.结果表明:品种演替过程中旗叶的叶绿素含量逐渐提高,衰老过程中近代品种叶绿素的降解速率低于较早年代品种;旗叶衰老过程中,近代品种荧光诱导动力学曲线的J点上升幅度小于I点;品种更替过程中光系统Ⅱ最大光化学效率和单位面积有活性反应中心数目逐渐增加,但是近代品种降低速率低于较早年代品种.叶绿素含量的变化与未衰老叶片中F_v/F_m没有显著相关性,但是随着衰老程度增加,相关性逐渐增大,且趋势线斜率逐渐提高;光系统Ⅱ单位面积有活性反应中心数目与品种育成时间呈显著正相关,且随着衰老程度增加,相关程度和趋势线斜率均显著提高.综上,小麦品种演替过程中,旗叶叶绿素含量逐渐升高,降解速率逐渐减缓,光合电子传递过程中Q_A到Q_B电子传递的抗衰老能力得到改善,从而减缓了衰老过程中光系统Ⅱ最大光化学效率和有活性反应中心的衰减速率,同时,叶绿素含量的提高和旗叶光系统Ⅱ抗衰老能力的增强也是品种更替过程中产量逐渐提高的重要因素.     

Influence of reproductive organs on plant senescence in rice and wheat

The chlorophyll and protein contents of the flag, second and third leaves gradually decreased during the reproductive development of rice (Oryza sativa L. cv. Rasi) and wheat (Triticum aestivum L. cv. Sonalika) plants, whereas proline accumulation increased up to the grain maturation stage and slightly decreased thereafter. In rice plant, the rate of decrease in chlorophyll and protein and increase in proline level were higher in the flag leaf than in the second leaf. It was opposite in wheat plant. The export of [³²P]-phosphate from leaves to grains gradually increased reaching a maximal stage at the grain development stage, and then declined. The export of this radioisotope was greater in rice than in wheat. Removal of panicle at the anthesis and grainfilling stages delayed leaf senescence of rice plant, while in wheat the ponicle removal at any stage did not have a marked effect on delaying leaf senescence. The contents of chlorophyll and protein of glumes were higher in wheat than in rice. The variation of such source-sink relationship might be one of the possible reasons for the above effect on leaf senescence.     

小麦旗叶Rubisco周转与籽粒含氮量的关系

随着旗叶的衰老,Rubisco含量逐渐减少.延缓小麦旗叶的衰老进程(抽穗期施氮肥),可增加旗叶Rubisco的含量,提高籽粒的全氮含量.在小麦旗叶全展后28d内,Rubisco的15N丰度处于较高水平,表明仍有Rubisco的重新合成;而在28d以后,Rubisco的15N丰度处于低水平,表明无Rubisco的重新合成.但这时籽粒的15N丰度却上升.旗叶全展后14d内Rubisco的15N丰度高于旗叶中全氮的丰度,说明此时期Rubisco重新合成的速率高于其它蛋白质;旗叶衰老过程中Rubisco的15N丰度的净转移高于全氮,Rubi-sco净N转移也高于全氮,表明Rubisco向籽粒中转移的氮素多于其它蛋白质,对籽粒含氮量的影响最大.     

播种量和施氮量对垄沟覆膜栽培冬小麦花后生理性状的影响

为协调冬小麦个体与群体间的关系,充分发挥旱作条件下垄沟栽培优势,以冬小麦品种小偃22为材料,采用二元二次正交旋转组合设计,通过田间试验研究了垄下集中施肥、垄上覆膜、膜际种植模式下播种量和施氮量对冬小麦花后生理性状的影响.结果表明： 花后叶面积指数、旗叶叶绿素含量和净光合速率均随施氮量的增加而增加.灌浆前中期叶面积指数随播种量的增加呈先增后稳的趋势;灌浆后期叶面积指数随播种量的增加而降低.随播种量的增加,旗叶的叶绿素含量和净光合速率降低,单株产量呈先减少后增加的趋势.适宜的播种量可以协调个体与群体间的矛盾,而适量增施氮肥有利于花后小麦生理性状的改善和产量的提高.在供试条件下,小偃22在播种量112.5 kg·hm-2与施氮量180～222 kg N·hm-2配置时,个体与群体的关系比较协调,花后叶面积指数较高,群体结构适宜,而且旗叶叶绿素含量、净光合速率和单茎产量较高,能获得高产.     

播种量和施氮量对垄沟覆膜栽培冬小麦花后生理性状的影响

为协调冬小麦个体与群体间的关系,充分发挥旱作条件下垄沟栽培优势,以冬小麦品种小偃22为材料,采用二元二次正交旋转组合设计,通过田间试验研究了垄下集中施肥、垄上覆膜、膜际种植模式下播种量和施氮量对冬小麦花后生理性状的影响.结果表明:花后叶面积指数、旗叶叶绿素含量和净光合速率均随施氮量的增加而增加.灌浆前中期叶面积指数随播种量的增加呈先增后稳的趋势;灌浆后期叶面积指数随播种量的增加而降低.随播种量的增加,旗叶的叶绿素含量和净光合速率降低,单株产量呈先减少后增加的趋势.适宜的播种量可以协调个体与群体间的矛盾,而适量增施氮肥有利于花后小麦生理性状的改善和产量的提高.在供试条件下,小偃22在播种量112.5 kg hm-2与施氮量180 ～222 kg N·hm-2配置时,个体与群体的关系比较协调,花后叶面积指数较高,群体结构适宜,而且旗叶叶绿素含量、净光合速率和单茎产量较高,能获得高产.     

Ven、K型小麦雄性不育系春性F1杂种的优势表现及与叶绿素含量和同工酶谱的关系

利用Ven、K型小麦雄性不育系与北方春性普通小麦杂交,研究了Ven、K型春性F1杂种的优势表现及其与叶绿素含量、同工酶谱的关系。结果表明：Ven型和K型杂种F1存在显著的个体杂种优质,在主要农艺性状中尤其以单株穗数、千粒重、单株生产力的杂种优势明显,其中单株穗数对单株生产力的贡献最大;相关分析表明：单株生产力与单株穗数呈正相关,相关系数为0.7122;与每穗粒数和千粒重的相关系数分别为－0.1183和0.4941。灌浆期旗叶、倒二叶叶绿素含量与千粒重的相关性最大,呈正相关。杂种F1优势强的组合其种子和苗期的过氧化物同工酶和酯酶的酶谱一般呈双亲互补型或“杂种型”酶带,该特征可作为杂种优势预测的指标。     

RuBP carboxylase activity in wheat following ear emergence and evaluation of the role of ear

Ribulose bisphosphate carboxylase activity and chlorophyll content were measured in the leaves and reproductive parts of two wheat cultivars after ear emergence. The chlorophyll content of the flag leaf was mostly higher than that of the awns and glumes. Awns had the highest chlorophyll content among reproductive parts. Light transmission to the lower leaves was higher in the dwarf cultivar Moti than in the medium tall cultivar Kalyansona. RuBP carboxylase activity in Kalyansona leaves was higher than in Moti leaves. In postanthesis stages there was no difference in RuBP carboxylase activity in the flag leaf and lower leaves between cultivars. Awns had the maximum activity of RuBP carboxylase followed by glumes and grains among ear parts. The relative capacity for photosynthesis in the ear parts was several times higher than in the flag leaf on a unit chlorophyll basis. It is suggested that in a crop canopy in the field, the spike(ear) may have a greater importance in grain development than has been previously estimated.     

Responses of Penultimate and Flag Leaves of Wheat to Different Nitrogen Supply

Uniculm wheat (Triticum aestivum L.) was grown to maturity at four concentrations of nitrogen corresponding to 3 (N1), 6 (N2), 9 (N3) and 12 (N4) g m^–2. Penultimate and flag leaves were examined throughout the ontogeny. Sub-optimal concentrations of N resulted in sharp decline in both area and dry mass of the leaves. Decline in leaf area was due to fewer mesophyll cells. Net photosynthetic rate (P_N) increased up to full expansion, remained constant for about a week and then declined. P_N, nitrogen, ribulose-1,5-bis-phosphate carboxylase/oxygenase (RuBPCO) amount and activity, chlorophyll and soluble protein contents were similar at all the N concentrations. Both amount and activity of RuBPCO in the flag leaf were about two fold higher as compared to penultimate leaf, but P_N was similar. This indicates the presence of an excess amount of RuBPCO in the flag leaf.     

三种栽培模式下不同基因型冬小麦旗叶衰老代谢比较

为了探索冬小麦在不同栽培模式下功能叶片衰老代谢的生理机制,以cp02(213)、cp99(1)和陕农512为材料,比较研究了常规栽培、覆草栽培、地膜覆盖3种栽培模式下小麦旗叶衰老代谢特性.结果表明,覆草栽培叶面积、旗叶功能期、叶绿素含量、叶片保护酶活性(SOD、POD、CAT)显著高于常规栽培,膜脂过氧化程度较低,叶片衰老速度缓慢,代谢强度旺盛,有利于籽粒灌浆和光合产物的积累.灌浆前期,地膜覆盖叶面积、旗叶功能期、叶绿素含量、叶片保护性酶活性(SOD、POD、CAT)显著高于常规栽培,膜脂过氧化程度低于常规栽培;灌浆后期,叶绿素含量急剧下降,叶片衰老速度加快,膜脂过氧化程度加剧.参试品种(系)中陕农512叶片衰老速度缓慢,保绿性好.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

小麦旗叶自然衰老过程中清除活性氧能力的变化

野生一粒小麦(Triticum boeoticum Boiss)、栽培小麦“扬麦五号”(T.aestivum L.)的旗叶自然衰老过程中,活性氧清除系统中各部分的清除能力下降是不均衡的。在光合速率高值持续期(叶绿素含量缓降期),SOD的活力略有下降,过氧化氢酶(CAT)活力却迅速下降,同时抗坏血酸过氧化物酶(ASP)活力呈现先上升后下降的趋势,上述SOD、CAT、ASP活力变化的不均衡,最终导致H_2O_2的迅速累积,从而使叶片迅速进入衰老(叶绿素含量速降期),于是SOD活力迅速下降。野生一粒小麦活性氧清除系统中各部分清除能力失衡过快,可能是其早衰的原因之一。H_2O_2的迅速累积与叶片衰老的启动密切相关。     

杂交小麦不同发育时期叶绿素蛋白复合体的变化

杂交小麦“９０１”在不同发育时期及不同叶位,叶片色素蛋白复合体含量均高于对照品种陕２２９,尤其表现在“９０１”的旗叶,而陕２２９的倒二叶在籽粒形成后期、灌浆期,略高于“９０１”,倒三叶在籽粒发育后期至成熟期“９０１”的含量要高于陕２２９。叶绿素蛋白复合体分析结果说明,在不同发育时期、不同叶位,“９０１”叶片色素蛋白复合体中含有较高的叶绿素b,尤其表现在籽粒形成后期的旗叶上。２个品种在不同发育时期,不同时叶位之间,色素蛋白复合体组分上没有差异。仅表现在成熟期,陕２２９的倒三叶各色素带含量减少,并且消失１条叶绿素蛋白复合体带,初步认为此带是ＬＨＣＰ的寡聚体之一,而“９０１”比陕２２９晚７d倒二叶上表现出样的现象,表明“９０１”叶片色素蛋白复合体抗衰老能力很强。     

Characterisation of grain yield and nitrogen level in relation to flag leaf senescence of wheat varieties

Three wheat cultivars ( Triticum aestivum L.), Splendeur, Hobbit and Maris Huntsman grown in pots were compared. Especially when compared to Splendeur, the flag leaf senesced most rapidly in Maris Huntsman, which presented the most rapid loss of moisture, chlorophyll and nitrogen. The uptake of exogenous nitrogen during the post-anthesis period was lower in the rapidly than in the slowly senescing variety. A higher concentration of free amino nitrogen in the flag leaf at a given sampling date was associated with a lower percentage decrease of soluble proteins at the following date. Acid proteinase activity in the flag leaf was inversely related to moisture percentage and free amino nitrogen level, but unrelated to the nitrogen loss of the flag leaf. Acid proteinase activity in the flag leaf was directly related to grain nitrogen percentage, but inversely related to grain yield. Grain yield was also directly related to the mean soluble protein content of the flag leaf through senescence.     

冷型小麦旗叶衰老和活性氧代谢特性研究

以典型的冷型小麦和暖型小麦为试验材料,研究了同一环境背景下不同温度型小麦开花后的旗叶衰老和活性氧代谢特性。结果表明,与暖型小麦相比,冷型小麦籽料灌浆期旗叶叶绿素和可溶性蛋白质含量下降缓慢、含量高,整个业粒形成和灌浆期ＭＤＡ积累速度慢、含量低,籽粒灌浆期防御活性氧伤害的关键性保护酶（ＳＯＤ、ＣＡＴ和ＰＯＤ）活性下隆幅度小,灌浆中后期活性水平高。由此认为,小麦旗叶衰老和活性氧代谢特性与其温度型的归属关     

小麦根系活力变化与叶片衰老的研究

对小麦根系活力、叶片叶绿素含量和光合速率变化的田问实验结果表明,根系活力的衰退早且快于叶片叶绿素含量和光合速率的衰退;根系活力和叶片光合速率呈正相关关系(r=0．8006)．拔节期追施氮肥,改善根际营养条件后,叶片的光合速率、根系活力明显提高,叶片中ABA含量降低,ZRs升高,SOD活性增加,MDA的高峰期比对照推迟8～10d,说明根系活力变化与地上部叶片的衰老密切相关,改善根际营养,提高生育前期的根系活力有助于延缓地上部分的衰老．     

施氮量和栽培模式对旱地冬小麦旗叶衰老及其活性氧代谢的影响

以冬小麦小偃22为试验材料,研究3种栽培模式(常规栽培、覆草栽培、地膜覆盖)和3种施氮水平(施纯氮0、120和240 kg/hm2)下旗叶衰老与活性氧代谢特性.结果表明,与常规栽培(CK)相比较,覆草栽培条件下叶绿素含量始终较高(P<0.05),叶片衰老速度缓慢,代谢强度旺盛,有利于籽粒灌浆和光合产物的积累,产量显著增加(P<0.05).在灌浆前期,地膜覆盖条件下叶片叶绿素含量增加(P<0.05),叶片保护性酶活性(POD、CAT)提高,膜脂过氧化程度低;但在灌浆后期,叶绿素含量急剧下降,叶片衰老速度加快,膜脂过氧化程度加剧,产量仍显著增加(P<0.05).施用氮肥在一定范围内可提高旗叶叶绿素含量和保护性酶活性(POD、CAT),降低膜脂过氧化程度;施氮量为120 kg/hm2时,冬小麦旗叶叶绿素含量最高,叶片衰老迟缓,代谢强度降低缓慢,膜脂过氧化程度低,有利于小麦后期生长和籽粒灌浆,在3种栽培模式下产量均最高.     

Characterization of photosynthesis of flag leaves in a wheat hybrid and its parents grown under field conditions

Two wheat cultivars, one with high yield and the other with a high tolerance against oxidative stress, were compared with a hybrid of these two cultivars by investigating their photosynthetic characteristics of flag leaves. From the beginning of flowering to the 17th day, CO(2) assimilation rate (P(max)) was maintained and there were no appreciable differences between the hybrid and its parents. P(max) showed no decrease at noon compared to that in the morning. From the 20th to the 30th day of flowering, P(max) decreased significantly, and this decrease was significantly less in the hybrid than in its parents. The actual photosystem II (PSII) efficiency (Phi(PSII)) and the maximal efficiency of PSII photochemistry (F(v)/F(m)) showed a significant decrease only on the 30th day after anthesis; this decline was much less marked in the hybrid relative to its parents, both in the morning and at noon. A decrease occurred in Phi(PSII) and F(v)/F(m) at noon when compared to that in the morning, but this decrease was less marked in the hybrid than in its parents. Rubisco activity decreased significantly from the 13th day of flowering and was higher in the morning than at noon both in the hybrid and its parents. However, the hybrid always showed a higher value of Rubisco activity. The activities of posphoenolpyruvate carboxylase and pyruvate phosphate dikinase showed similar changes to those in Rubisco activity, particularly from the 20th to 30th day. The results of this study suggest that the higher photosynthetic capacity of the flag leaf in the hybrid can help to accumulate more dry material, and may be the physiological basis for higher yield over its parents.     

二倍体和四倍体杂交兰幼苗对低温胁迫的生理响应差异分析

为探究不同倍性杂交兰（Cymbidium hybrid）对低温胁迫的生理响应差异,对低温胁迫下（昼温10℃、夜温4℃）二倍体杂交兰以及四倍体杂交兰株系T1和T2幼苗叶片中脯氨酸、可溶性糖、可溶性蛋白质、MDA和叶绿素含量,O-·2产生速率以及SOD、POD和CAT活性的动态变化进行了测定和分析;在此基础上,比较了二倍体和四倍体杂交兰的耐低温特性差异。结果显示：随着胁迫时间的延长,二倍体杂交兰和四倍体杂交兰株系T1和T2幼苗叶片中脯氨酸、可溶性糖、可溶性蛋白质和MDA含量,O-·2产生速率以及POD和CAT活性总体呈逐渐增加的趋势;叶绿素含量呈逐渐下降的趋势;SOD活性呈先上升后下降的趋势,并分别在胁迫后第9天和第6天达到峰值。在常温（25℃）下恢复生长3d后,二倍体杂交兰和四倍体杂交兰株系T1和T2幼苗叶片中脯氨酸、可溶性糖、可溶性蛋白质和MDA含量,O-·2产生速率以及SOD、POD和CAT活性均降低,而叶绿素含量升高。总体上,二倍体杂交兰叶片中脯氨酸、可溶性糖和可溶性蛋白质含量以及SOD、POD和CAT活性的增幅明显小于四倍体杂交兰,MDA含量和O-·2产生速率的增幅则大于四倍体杂交兰,而叶绿素含量的降幅则明显大于四倍体杂交兰。综合分析结果表明：在低温胁迫下,四倍体杂交兰叶片中渗透调节物质含量和保护酶活性均较高,其对低温的耐性强于二倍体杂交兰。