节点文献
冬麦春播条件下小麦生育特性及其产量形成的生理机制
Character of Growth and Devlopment and Physiological Mechanism of Yield Formation of Winter Wheat Varieties Sowed in Spring
【作者】 徐澜;
【导师】 高志强;
【作者基本信息】 山西农业大学 , 作物学, 2015, 博士
【摘要】 本试验于2013、2014连续两年在山西省农科院玉米研究所试验基地进行。采用大田种植、小区试验与盆栽试验相结合的方法,研究了22个南引冬麦品种在忻定盆地不同播期处理下植株农艺性状、幼穗分化、光合特性及干物质积累、叶绿素荧光参数、主茎内源激素动态变化及籽粒产量等,同时研究了不同品种光合特性、主茎内源激素等与籽粒产量及其构成的关系,明确适宜忻定盆地春播的优良小麦品种资源的生育特点及其产量形成机制,为混播区优良春小麦品种的引进、选择及指标评价提供参考,为西北冬春麦混播区发展高效优质小麦生产提供品种资源与理论依据。主要试验结果如下:1、冬麦春播对小麦幼穗分化及生长发育的影响冬麦春播条件下,其中20个来自长江中下游麦区、西南冬麦区的冬小麦品种(春性或弱春性)在忻定盆地3月上、中、下旬春播,均可以正常拔节、抽穗、成熟;南引半冬性小麦品种皖麦68表现为不拔节,迟抽穗,严重影响籽粒产量,不适宜忻定盆地春播;播期对不同小麦品种的影响不同,在忻定盆地表现良好的冬麦品种,如扬麦13、渝麦12、扬麦20、宁麦13、川麦42较适宜早播,随播期推迟生长发育受到影响,籽粒产量呈下降趋势。播期对小麦穗分化影响显著,随着播期推迟,气温升高,穗分化所需时间变短,Ⅰ播期(早播)、Ⅱ播期(适播)、Ⅲ播期(晚播)完成穗分化所需时间分别为40~45d、36~42d、33~41d;不同品种其幼穗分化进程在分化时间出现的早晚,完成幼穗分化所需天数等方面均不同;总体来看,植株外部形态特征表现出与穗分化进程的一致性,小麦叶片数为3.5~4.8叶时,根条数为8~9,幼穗分化为二棱期;叶片数大于5.6叶时,根条数大于10,幼穗分化开始进入雌雄蕊原基分化期。半冬性品种黔麦18的幼穗分化进程较缓慢,单棱期持续时间较长,当进入二棱期时,相应的叶片数和根条数较多;进入雌雄蕊原基分化期,外部植株叶片数、根条数增加不明显:皖麦68在盆栽试验中,不同播期株高都处于较低水平,生长发育缓慢,穗分化进程未随着植株叶片数和根条数的增加而进行,始终停留在单棱期,植株矮小,分蘖较多,不适应忻定盆地春播,这与大田试验结果完全一致。小麦穗分化受品种自身遗传特性与外界生态条件的共同影响,在不同播期中,不同品种的春小麦的幼穗分化进程不同,所需时间不同,其外部形态特征表现出一定的差异。渝麦12、扬麦13在早播条件下穗分化较其他品种迅速,仅需40d、41d。株高是小麦的重要外部形态指标之一,株高过高容易发生倒伏,植株过矮使冠层叶片拥挤,中下部透光不良,均影响产量。大多数品种表现为晚播处理的株高低于早播、适播,且早播、适播间差异不明显。扬麦13、扬麦20、宁2038、川麦42等获得较高产量的品种,其株高均在75-80 cm之间,而渝麦9的株高(90.7cm)过高,渝麦7仅有50.7cm,该二品种的籽粒产量均较低。生育时期与幼穗分化及形态特征相统一,并且与产量形成关系密切。渝麦12、渝麦10、扬麦13较早进入抽穗期,比对照宁2038、新春30早1~2天,比其他品种早3-5天;扬麦13最早进入散粉期,其生长发育较快,生殖生长阶段占总生育期比例较大,亦最早成熟,成熟期比其他品种早4-7天,这样可以有效避开极端天气对籽粒产量的影响,早播处理下与其他品种相比,其优势更加明显。渝麦12成熟也较早,与扬麦13具有同样的优势。2、冬麦春播对小麦旗叶光合、荧光参数及产量的影响小麦产量形成与其光合特性关系密切。多数品种的叶绿素含量、光合指标均表现为早播>适播或晚播;旗叶Pn较高的品种是扬麦13 (8.9 umolco2.m-2.s-1)、云麦42 (5.6 umolco2.m-2.s-1)、渝麦12 (4.6umolco2.m-2.s-1)(开花期):扬麦13 (16.7 umolco2.m-2.s-1)=川麦42 (16.5umolco2.m-2.s-1)(灌浆期);旗叶蒸腾速率、气孔导度和胞间C02浓度、旗叶水分利用率与Pn表现基本一致,其中WUE在品种间差异显著。大多数旗叶叶绿素荧光参数品种间差异显著,且相关性极显著。不同播期品种间Chl(叶绿素含量)变异系数较小(0.1217-0.1481),PIABS(吸收光能为基础的性能指数)变异系数较大(0.3293-0.3954),二者与产量偏相关系数均较大(0.7004~0.8135)。早播条件下,籽粒产量与PIABS(灌浆期、开花期)、灌浆期Chl呈显著正效应,与灌浆期Vi呈显著负效应,且产量的81.07%-82.78%(2013、2014)可由该三因素的变异决定;不同小麦品种表现不同播期效应,且两年变化趋势一致:扬麦13(春性、中早熟)旗叶Chl及绝大多数荧光参数和产量均显著高于其它品种(p≤0.05),且适宜早播。早播条件下,灌浆期旗叶叶绿素含量、PIABS可作为选择高光效小麦资源的重要评价指标。早播条件下,小麦单株干物质积累均符合logistic模型,拔节前干物质积累较缓慢,孕穗后增长速率显著加快。开花期至乳熟期干物质积累显著高于其他品种的是渝麦12和渝麦9,尤其是渝麦12在干物质积累过程中增长速度呈递增趋势,其最终单株干物质积累量为最高,并显著高于其他品种;大多数南引品种的早播增产效应较明显,且播期间差异显著,两年试验结果基本一致。(2013年)不同播期处理下籽粒平均产量分别为4393.50kg.hm-2(ES:早播),3871.99 kg.hm2(NS:适播),3623.50 kg.hm-2(LS:晚播);其中渝麦12、扬麦13的产量显著高于其他品种,ES、NS、LS播期下分别为5876.90 kg.hm2~5351.55 kg.hm-2、4987.05 kg.hm2~4609.20 kg.hm-2、4014.15 kg.hm2~4058.70 kg.hm2;但个别品种对播期的响应不同,宁2038的产量表现为NS> LS> ES,即适播条件下其产量最高;渝麦9则为LS>NS>ES,晚播时其产量较高,越早播产量反而越低;大粒品种云麦42则为ES>LS>NS,早播产量最高,但晚播的产量又高于适播处理。南引冬小麦品种在产量构成因素中,成穗数和穗粒数占有一定优势,如扬麦13(ES)的成穗数(618.0×104hm-2)显著高于其他品种,渝麦12的穗粒数(两年在35.1粒-40.6粒之间)均显著高于其他处理。可见,冬麦春播条件下,提高小麦产量应主攻成穗数和穗粒数,促进其有效小穗的形成,在此基础上,加强灌浆后期栽培管理调控,促进籽粒灌浆速度与强度,以此增加粒重。渝麦12、扬麦13、川麦42等具有较高光合效率,利于干物质积累,产量较高。3、冬麦春播对小麦苗穗期植株抗寒性的影响小麦穗分化阶段对低温胁迫较为敏感,对于从长江中下游、西南冬麦区引进的优良冬小麦品种,在忻定盆地春播条件下,其穗分化阶段耐、抗寒性强弱是引种适应性的重要考量因素之一。冬麦春播条件下,经历穗分化阶段低温胁迫后,各小麦品种相对电导率、可溶性蛋白含量、可溶性糖含量不同程度地升高,总叶绿素含量程度不同地下降。扬麦13、渝麦12的叶片相对电导率在低温前后的变化幅度较小(40.7%-24.6%、52.7%-38.2%),低温胁迫后其可溶蛋白含量较高(16.78-19.78mg g-1FW、15.76~18.77mg g-1FW),叶片可溶性糖含量在遭遇低温胁后增幅较大,分别达285%-235%、265.6%-218%(CKl的增幅为222%-143%)。低温胁迫下,品种抗寒性强于对照(宁2038:CKl、新春30:CKz)的依次为:扬麦13、渝麦12、宁麦13、皖麦68、渝麦9;籽粒产量数据方差分析同样表明,渝麦12、扬麦13、宁麦13产量较高,且与其它品种差异显著(p=0.05)(2013)。2014年品种抗寒性综合评价值居前四位的是扬麦13、宁麦13、皖麦68、渝麦12;其产量方差分析表明,扬麦13、宁麦13、渝麦12显著(p=0.05)高于其它品种。扬麦13主茎叶片GA、不同程度地升高,总叶绿素含量明显下降。通过主成分分析和抗寒性度量值(D值)排序,对品种幼穗分化期间的耐、抗寒性进行初步评价:渝麦10、扬麦20、云麦42抗寒性较差;扬麦13、渝麦12、宁麦13表现较强抗寒性且年际稳定性好(D值分别为0.6656-0.6596,0.4934-0.4959,0.4718-0.5835;而CK1、CK2分别为0.3682-0.3978,0.3285-0.3304),其两年籽粒产量亦显著(p=0.05)高于其它品种,可作为混播区引种材料进一步研究。
【Abstract】 Total of 22 winter wheat varieties were imported from the middle and lower reaches of the Yangtze river region in China. Those varieties were sowed in spring in Xinding basin area of Shanxi province (Shanxi Academy of Agricultural Sciences, Maize Research Institute).The field and pot trials were performed for two years (2013-2014). The traits and physiological characteristics included plant agronomic traits, spike differentiation, photosynthetic characteristics and dry matter accumulation, chlorophyll fluorescence parameters, dynamic changes of endogenous hormones in the main stem and grain yield were tested. In order to make clear the formation mechanism of growth characteristics and yield of wheat varieties suitable for Xinding basin spring sowing, the relationship between light photosynthetic characteristics, endogenous hormones of the main stem and the grain yield were studied. To provide theoretical base for selection of high efficiency and evaluation index in wheat introduction and selection for the northwest of mixed sowing area, the field were carried out. The main results are as follows:1> Effect on wheat spike differentiation and growth of spring sowing of winter wheat20 cultivars (spring or weak spring) imported from the middle and lower reaches of the Yangtze river region in China were sowed in March, spring in Xinding basin area of Shanxi province, can be normal jointing, heading, mature; South bring semi winter wheat variety Wanmai 68 showed not to pull out section, delayed heading, seriously affecting grain yield, not suitable for Xinding basin in spring sowing period; different effects on different wheat varieties, good performance of a variety of Winter Wheat in Xinding basin, such as Yu wheat 12, Yangmai 13, Yangmai 20, Ningmai 13, Chuanmai 42 suitable early sowing, sowing time delayed affected the growth and development, grain yield decreased.The spike differentiation processes were obviously influenced by sowing dates, the time of young spike differentiation became shorter with sowing delay, the days completed spike differentiation were different in I, n, III:40-45d,36-42d,33-41d; The spike differentiation process is different of different cultivars; At the same time, plant morphological characteristics showed consistency with spike differentiation progress. When numbers of wheat leaves reached 3.5-4.8, root numbers was 8-9, the spike differentiation got into double ridge stage; When the number was over 5.6 leaf, root number was than 10, it entered the spike differentiation of pistil and stamen. The spike differentiation process of Qian Mai 18 was slower, single ridge stage lasted longer time; When entering double ridge stage, the number of leaves and root number were increased obviously; During the spike differentiation of pistil and stamen, the external leaf number and root number increased remarkably. Wanmai 68 in different sowing date, plant height were low, slow growth, spike differentiation process were not increased with the leaf number and root number, and always stayed in the single ridge stage, tiller was more, could not adapt to the Xinding basin spring sowing, the same result was obtained from field test. Genetic characteristics of wheat varieties and ecological conditions were influenced the spike differentiation; Different varieties of spring wheat spike differentiation was different in different sowing period, its external morphology showed some differences. Compared to other, spike differentiation of Yumai 12 and Yangmai 13 was fast in early sowing date, only 40d and 41d.Plant height is one of the important indicators of the external morphology of wheat, high too prone to lodging, plant canopy leaves too short so crowded, the lower light transmission was bad, affecting production. Plant height of most varieties in late sowing treatment were lower than early or normal sowing dates, and there was no significant difference between ES and NS. Yangmai 13, Yangmai 20, Ning 2038, Chuanmai 42 varieties such as access to high yield, plant height were between 75-80 cm, but Yumai 9 was high(90.7cm), Yu Mai 7 was only 50.7cm, grain yield of the two species were lower. Panicle initiation, growth stages and morphological characteristics of plant were closed ties with production formed. Yumai 12, Yumai 10, Yangmai 13 early reached heading stage for 1-2 days with control N2038, for 3-5 days earlier than other varieties; Yangmai 13 first entered shedding period, their growth and development rapid reproductive growth stage of a larger proportion of total fertility, but also the earliest maturity, which could effectively avoid the impact of local rainfall and hot wind on grain yield; mature 4-7 days earlier than other varieties, which can effectively avoid the extreme impact of weather on grain yield, compared with other species, their advantage is more obvious under the early sowing treatment. Yumai 12 matured earlier, and it had same advantages as Yangmai 13.2、Effect on photosynthetic characteristics, chlorophyll fluorescence kinetics parameters flag leaf and yield of spring sowing of winter wheatWheat yield formation was closely related to its photosynthetic characteristics. Chlorophyll content, photosynthetic performance indicators for most varieties were ES> NS or LS; Pn flag leaf varieties were higher,such as Yangmai 13 (8.9 umolco2.m-2.s-1), Yunmai 42 (5.6 umolco2.m-2.s-1), Yumai 12 (4.6umolco2.m-2.s-1) (flowering stage); Yangmai 13 (16.7 umolco2.m-2.s-1), Chuanmai 42 (16.5umolco2.m-2.s-1) (filling stage); Tr, Gs, Ci, WUE on flag leaf were agreed with Pn, which WUE significant differences among varieties.Significantly correlations were observed in the most chlorophyll fluorescence kinetics parameters of different cultivars, except for the chlorophyll content. The variation coefficients of chlorophyll content and PIBS were respectively lower (0.12-0.17) and higher (0.32~0.39), and partial correlation coefficients of them with yield in 2013、2014 were 0.70~0.81 under two sowing dates. Path analysis and multiple stepwise regression analysis indicate that chlorophyll content (filling stages) and PIBS (flowering and filling stages) has an obvious positive effect on grain weight, but Vi in flag leaves at filling stages has a negative effect in early sowing. So, the three factors were determiners of grain yield (account for 81.07%~82.78%). The chlorophyll content, Pn, most chlorophyll fluorescence kinetics parameters and grain yield of Yangmai 13 were significantly difference at p<0.05, and early sowing was suited to it. By multiple statistics, the chlorophyll content (filling stages), PIABS (flowering and filling stages) were considered as important indices in the selection of high photosynthetic efficiency in wheat under early sowing dates.Under the conditions of early sowing, plant dry matter accumulation of wheat are in line with logistic model, the dry matter accumulation before jointing more slowly, after booting the growth rate significantly accelerated. Flowering to milky stage of dry matter accumulation was significantly higher than other varieties of wheat is Yumai 12 and Yu mai 9, especially in the dry matter accumulation Yumai 12 during growth rate showed an increasing trend, the final amount of dry matter accumulation per plant maximum, and significantly higher than other varieties; yield effect early sowing varieties most Southern cited more obvious during the broadcast and the difference is remarkable, two-year test results are basically the same. (2013) Different Sowing average grain yield was 4393.50kg.hm-2 (ES),3871.99 kg.hm-2 (NS),3623.50 kg.hm-2 (LS); The yield of Yumai 12 and Yangmai 13 were significantly higher than other varieties, respectively were 5876.90 kg.hm-2~5351.55 kg.hm"2,4987.05 kg.hm-2~4609.20kg.hm-2,4014.15 kg.hm-2~ 4058.70 kg.hm"2 under three sowing dates; but the response of individual species to sowing dates were different:yield of N2038 was NS> LS> ES; yield of Yumai 9 was LS> NS> ES; but large grain varieties Yunmai 42 was ES> LS> NS. Southern cited winter wheat varieties yield components, spike and grain number to gain some advantages, such as Yangmai 13 (ES) of a spike (618.0×104hm-2) was significantly higher than other varieties, Yumai 12 grains (two years between 35.1~40.6) were significantly higher than other treatments. Visible under winter wheat sowing conditions to improve wheat production should be the main spike and grain number, promote the formation of effective spikelets, on this basis, to strengthen the cultivation and management of post-filling regulation to promote grain filling speed and strength, in order to increased grain weight. Yumai 12, Yangmai 13, Chuanmai 42, etc. have higher photosynthetic efficiency, dry matter accumulation, higher yield.3、Effect on cold tolerance of plants during heading stage of spring sowing of winter wheatLow temperature stress is one of the important factors resulting in the yield reduction of wheat. Spike differentiation stage is more sensitive to low temperature stress, for excellent winter wheat varieties from the Yangtze River, southwest of the introduction of the winter wheat region, Xinding Basin in spring conditions, the panicle initiation stage of resistance, cold resistance strength is Adaptability of One of the important factors to consider. The current study was conducted to investigate the low-temperature response and cold tolerance of winter wheat varieties sowed in spring during spike differentiation stage, the most sensitive period of wheat to temperature. The traits and physiological characteristics tested under low temperature stress included grain yield, total content of chlorophyll photosynthesis, osmotic adjustment, membrane system, ion leakage rate, content of soluble sugar and soluble protein. The cold tolerance level of those wheat varieties were assessed integrative by statistical tools such as ANOVA analysis of grain yields and Principal Component Analysis. Low temperature stress led to increase of wheat leaf ion leakage rate, soluble sugar and protein contents, but reduction of chlorophyll. The folds of changes were different for different varieties tested. According to Principal Component Analysis and D value, Yumai 10, Yangmai 20, and Yunmai 42 were classed as cold sensitive wheat varieties. Yangmai 13,Yumai 12, and Ningmai 13 were classed as stronger cold-resistant wheat genotypes, and the phenotype showed stable through two-year field trials. The D values of each variety calculated for two years were 0.6656~ 0.6596,0.4934-0.4959, and 0.4718-0.5835 for the three cold resistance varieties, respectively while D values for CK1 and CK2 were 0.3682~0.3978,and 0.3285~0.3304, respectively. The grain yield of the cold resistant wheat varieties were significantly higher than other varieties tested(p<0.05). Therefore, Yangmai 13,Yumai 12 and Ningmai 13 could be imported from low latitude areas, and used as the cold tolerant wheat varieties for planting in North Plain of China. However, more studies are needed.