首页 > 分享 > 不同播期冬小麦小花发育特性与同化物代谢的相关性

不同播期冬小麦小花发育特性与同化物代谢的相关性

花匠小妙招
2025-06-14 23:35

摘要:

目的

推迟播期能够维持单位面积粒数已在前期研究中得到证实，本研究进一步探讨不同播种期冬小麦小花发育特征及分化、退化的差异性，分析植株同化物积累、分配与小花发育和结实的关系，旨在为提高小麦穗粒数以及丰富小麦高产栽培理论提供参考。

方法

试验于2014—2015年和2015—2016年连续两个小麦生长季，在山东省泰安市岱岳区大汶口镇东武村山东农业大学试验田 (35°57′N，117°3′E) 进行，以泰农18和济麦22为试验材料，设置9月24日、10月1日、10月8日、10月15日和10月22日共5个播期处理，观察记载小花发育过程中分化和退化数量动态以及最终结实粒数，测定开花期穗部和茎秆的干物质和氮素积累量，计算小花退化阶段整株和穗部的干物质和氮素积累速率。

结果

播期推迟条件下，单位面积可孕花数和单位面积结实粒数获得了维持，单位面积最大分化小花数明显降低，小花存活率显著提高。推迟播期明显减少了小花退化数量，小花退化速率明显降低。在小花退化阶段，推迟播期加速了整株和穗部的干物质和氮素积累，从而为开花期穗干重的维持以及干重穗茎比和氮素穗茎比的提高提供了保证。相关分析表明，开花期单位面积可孕花数分别与穗干物重、干物重和氮积累量穗茎比、整株和穗的干物质和氮的积累速率以及小花存活率均呈极显著正相关。推迟播期明显减少了有效茎蘖和无效分蘖所产生的退化小花数，尤其无效分蘖产生的退化小花数下降幅度更加明显。提高主茎在群体中同化物所占比例能够有效维持单位面积可孕花数和减少退化小花数，降低了小花死亡消耗。此外，推迟播期提高了可孕花累积生长度日生产效率，降低了退化小花累积生长度日生产效率。

结论

推迟播期明显降低了小花的退化，从而提高了小花存活率，开花期单位面积可孕花数的维持与植株同化物代谢密切相关。适期晚播提高了主茎在群体中的比重，有利于减少无效消耗和提高资源利用效率。

关键词: 冬小麦 / 播期 / 小花发育 / 同化物代谢

Abstract:

Objectives

Our previous work confirmed that grain number per unit area could been maintained through delaying sowing date. This study further investigated the development characteristics of florets and compared their differences in differentiation and degradation under different sowing dates, and analyzed the relationship between photosynthate accumulation and distribution and floret development and grain setting. The purpose was to provide references for wheat high-yield cultivation.

Methods

Field experiments were conducted during the 2014−2015 and 2015−2016 at the Experimental Station of Dongwu Village (35°57′N, 117°3′E), Shandong Province, China. Two wheat cultivars of Tainong18 and Jimai22 were planted as the experimental materials. Winter wheat was sowed on September 24, October 1, 8, 15 and 22, respectively. Binocular microscope was used to record the dynamics of differentiation and degeneration during the floret development. The final grains number, the dry matter and nitrogen accumulation of spikes and stems at anthesis were investigated, and the accumulation rate of dry matter and nitrogen in the whole plants and spikes during the floret degeneration stage were calculated.

Results

Among the five sowing date treatments, the number of fertile florets and final grains per unit areas were maintained same, but the maximum number of differentiated florets per unit area was significantly reduced, and the survival rate of florets was significantly increased. When the sowing date was delayed, the degenerated rate of florets was significantly reduced. The accumulation of dry matter and nitrogen in the whole plants and spikes were significantly accelerated during the floret degeneration stage, thus ensure the constant spike dry weight and the ratios of dry matter and nitrogen between spikes and stems at anthesis. The number of fertile florets at anthesis exhibited significant and positive correlations with spike dry matter weight, the ratios of dry matter weight and nitrogen accumulation of spike/stem, and the accumulation rate of dry matter and nitrogen in the whole plants and spikes and survival rate of fertile florets, respectively. The number of degenerated florets per square meter produced by effective stems and ineffective tillers significantly reduced as a response to delay sowing date, and the number of degenerated florets per square meter produced by ineffective tillers was even more significantly decreased. Enhanced the proportion of the main shoots in the population of photosynthate can be in favor to maintain the number of fertile florets and decrease the number of degenerated florets, and reduce the depletion of florets death. In addition, the growing degree day production efficiency of fertile florets increased and the growing degree day production efficiency of degenerated florets reduced as a result of delaying the sowing date.

Conclusions

Delayed the sowing date could increase the survival of florets through inhibiting the floret degeneration. The maintenance of the number of fertile florets is closely related to the photosynthate metabolism of plant. Delaying the sowing date could increase the proportion of the main stem in the plant population, which is beneficial to reduce the invalid consumption and improve the utilization efficiency of resource.

图 1 不同播期条件下冬小麦小花分化、退化与累积生长度日 (GDD) 的关系

Figure 1. Relationships between floret differentiation and degeneration and growing degree day (GDD) of winter wheat in different sowing dates

图 2 播期 (累积生长度日，GDD) 对冬小麦小花分化和退化速率的影响

Figure 2. Effects of sowing date (growing degree day, GDD) on the differentiated rates and degenerated rates of florets of winter wheat

图 3 开花期同化物积累和分配与单位面积退化小花数的相关分析

Figure 3. Correlation analysis between photosynthate accumulation and distribution and the number of degenerated florets per square meter at anthesis

表 1 播期对冬小麦单位面积穗数、穗粒数、粒重和产量的影响

Table 1 Effects of sowing date on spikes per square meter, grains per spike, grain weight and yield

品种
Cultivar 播期 (m-d)
Sowing date 穗数 (No./m2)
Spikes number 穗粒数 (No./spike)
Grains per spike 粒重 (mg)
Grain weight 产量 (t/hm2)
Yield T18 09-24 (S1) 710.17 a 38.97 c 37.41 a 9.19 a 10-01 (S2) 699.23 ab 39.98 c 37.59 a 9.32 a 10-08 (S3) 684.38 b 41.89 b 37.82 a 9.48 a 10-15 (S4) 667.89 c 43.30 ab 38.09 a 9.58 a 10-22 (S5) 635.67 d 43.74 a 38.23 a 9.42 a J22 09-24 (S1) 736.67 a 34.50 b 39.30 a 9.19 a 10-01 (S2) 714.45 ab 35.90 b 39.66 a 9.38 a 10-08 (S3) 691.85 bc 37.37 a 39.74 a 9.62 a 10-15 (S4) 674.23 cd 38.18 a 40.08 a 9.54 a 10-22 (S5) 642.45 d 38.57 a 40.29 a 9.41 a 注（Note）：数据为两生长季平均值；同列数据后的不同字母表示同一品种不同播期间差异显著 (P < 0.05)。 Values were the mean of the two growing seasons; Values followed by different letters in the same column mean significant difference among sowing dates for the same cultivar (P < 0.05).

表 2 小麦品种T18和J22达到出苗期、拔节期、孕穗期、开花期和成熟期的日期 (y-m-d)

Table 2 The date for emergence, jointing, booting, anthesis and maturity for wheat cultivars of T18 and J22

品种
Cultivar 年份
Year 播期
Sowing date 出苗期
Emergence 拔节期
Jointing 孕穗期
Booting 开花期
Anthesis 成熟期
Maturity T18 2014—2015 2014-09-24 2014-09-30 2015-03-29 2015-04-22 2015-05-03 2015-06-10 2014-10-01 2014-10-08 2015-03-29 2015-04-22 2015-05-03 2015-06-10 2014-10-08 2014-10-15 2015-03-29 2015-04-22 2015-05-03 2015-06-10 2014-10-15 2014-10-23 2015-03-30 2015-04-22 2015-05-03 2015-06-10 2014-10-22 2014-10-31 2015-03-30 2015-04-22 2015-05-03 2015-06-10 2015—2016 2015-09-24 2015-10-01 2016-03-26 2016-04-19 2016-04-30 2016-06-08 2015-10-01 2015-10-09 2016-03-26 2016-04-19 2016-04-30 2016-06-08 2015-10-08 2015-10-16 2016-03-26 2016-04-19 2016-04-30 2016-06-08 2015-10-15 2015-10-24 2015-03-27 2016-04-19 2016-04-30 2016-06-08 2015-10-22 2015-10-31 2016-03-27 2016-04-19 2016-04-30 2016-06-08 J22 2014—2015 2014-09-24 2014-10-01 2015-03-28 2015-04-21 2015-05-01 2015-06-10 2014-10-01 2014-10-09 2015-03-28 2015-04-21 2015-05-01 2015-06-10 2014-10-08 2014-10-16 2015-03-28 2015-04-21 2015-05-01 2015-06-10 2014-10-15 2014-10-24 2015-03-29 2015-04-21 2015-05-01 2015-06-10 2014-10-22 2014-10-31 2015-03-29 2015-04-21 2015-05-01 2015-06-10 2015—2016 2015-09-24 2015-10-01 2016-03-26 2016-04-18 2016-04-29 2016-06-08 2015-10-01 2015-10-09 2016-03-26 2016-04-18 2016-04-29 2016-06-08 2015-10-08 2015-10-16 2016-03-26 2016-04-18 2016-04-29 2016-06-08 2015-10-15 2015-10-24 2016-03-27 2016-04-18 2016-04-29 2016-06-08 2015-10-22 2015-10-31 2016-03-27 2016-04-18 2016-04-29 2016-06-08

表 3 播期对冬小麦穗分化和结实特性的影响

Table 3 Effects of sowing date on the characteristics of floret differentiation and seed setting of winter wheat

品种
Cultivar 播期 (m-d)
Sowing date MDF
(× 104 No./m2) FF
(× 104 No./m2) GN
(× 104 No./m2) SF
(%) SRF
(%) SRFF
(%) T18 09-24 (S1) 14.51 a 3.17 a 2.89 a 21.88 d 19.95 d 91.27 a 10-01 (S2) 13.82 a 3.22 a 2.92 a 23.34 d 21.19 c 90.84 a 10-08 (S3) 12.87 b 3.32 a 3.00 a 25.79 c 23.34 b 90.56 a 10-15 (S4) 11.95 c 3.37 a 3.05 a 28.26 b 25.53 ab 90.40 a 10-22 (S5) 10.97 d 3.29 a 2.96 a 30.04 a 27.08 a 90.23 a J22 09-24 (S1) 14.45 a 2.58 a 2.38 a 17.85 d 16.47 d 92.27 a 10-01 (S2) 13.90 a 2.62 a 2.41 a 18.81 d 17.30 d 91.95 a 10-08 (S3) 13.16 b 2.64 a 2.42 a 20.07 c 18.38 c 91.58 a 10-15 (S4) 12.18 c 2.68 a 2.45 a 21.97 b 20.09 b 91.45 a 10-22 (S5) 10.55 d 2.63 a 2.41 a 24.96 a 22.81 a 91.37 a 注（Note）：MDF—最大分化小花数 Maximum number of differentiated florets；FF—可孕花数 Numbers of fertile florets；GN—结实粒数 Grain number；SF—小花存活率 Survival rate of florets；SRF—小花结实率 Setting rate of florets；SRFF—可孕花结实率 Setting rate of fertile florets；数据为两生长季平均值 Values were the mean of the two growing seasons. 同列数据后不同小写字母表示同一品种不同播期间差异显著 (P < 0.05) Values followed by different small letters mean significant difference among the sowing dates for the same cultivar (P < 0.05).

表 4 不同播期小麦开花期不同植株器官单位面积干物质积累量和积累速率

Table 4 Accumulation and accumulative rate of dry matter of different plant organs at anthesis stage under different sowing dates

品种
Cultivar 播期 (m-d)
Sowing date 积累量 Accumulation (g/m2) 积累速率 Accumulative rate [g/(m2∙d)] 穗 Spike 茎 Stem 穗茎比 Spike/stem 穗 Spike 整株 Whole plant T18 09-24 (S1) 240.23 a 717.64 a 0.33 b 13.46 b 22.85 d 10-01 (S2) 244.37 a 710.39 ab 0.34 b 14.34 b 25.95 cd 10-08 (S3) 247.82 a 701.87 ab 0.35 ab 14.96 ab 27.75 bc 10-15 (S4) 250.94 a 695.18 b 0.36 ab 15.50 ab 29.92 ab 10-22 (S5) 252.43 a 674.83 b 0.37 a 15.95 a 31.34 a J22 09-24 (S1) 205.41 a 746.81 a 0.28 b 11.70 b 24.64 d 10-01 (S2) 208.72 a 738.19 ab 0.28 b 12.17 b 26.76 cd 10-08 (S3) 210.35 a 725.79 ab 0.29 ab 12.54 ab 28.82 bc 10-15 (S4) 211.58 a 713.07 b 0.30 ab 12.91 ab 30.29 ab 10-22 (S5) 212.31 a 688.82 c 0.31 a 13.30 a 33.34 a 注（Note）：数据为两生长季平均值 Values were the mean of the two growing seasons. 同列数据后不同小写字母表示同一品种不同播期间差异显著 (P < 0.05) Values followed by different small letters mean significant difference among the sowing dates for the same cultivar (P < 0.05).

表 5 播期对开花期不同植株器官单位面积氮素积累量和积累速率的影响

Table 5 Effects of sowing date on the accumulation and accumulative rate of nitrogen of different plant organs at anthesis

品种
Cultivar 播期 (m-d)
Sowing date 氮积累 N accumulation (g/m2) 氮积累速率 N accumulative rate [g/ (m2∙d) ] 穗 Spike 茎 Stem 穗茎比 Spike/stem 穗 Spike 整株 Whole plant T18 09-24 (S1) 5.36 a 12.43 a 0.43 b 0.20 b 0.28 d 10-01 (S2) 5.10 ab 11.92 a 0.43 b 0.21 b 0.30 cd 10-08 (S3) 5.07 bc 11.39 ab 0.45 ab 0.22 ab 0.32 bc 10-15 (S4) 4.88 c 10.71 b 0.46 a 0.22 ab 0.35 ab 10-22 (S5) 4.59 d 10.00 b 0.46 a 0.23 a 0.37 a J22 09-24 (S1) 5.25 a 13.36 a 0.37 b 0.22 b 0.28 c 10-01 (S2) 5.16 a 13.01 ab 0.38 b 0.23 ab 0.31 c 10-08 (S3) 4.85 b 12.31 b 0.39 ab 0.24 ab 0.34 b 10-15 (S4) 4.80 b 12.01 bc 0.40 a 0.24 ab 0.36 b 10-22 (S5) 4.54 c 11.25 c 0.41 a 0.25 a 0.40 a 注（Note）：数据为两生长季平均值 Values were the mean of the two growing seasons. 同列数据后不同小写字母表示同一品种不同播期间差异显著 (P < 0.05) Values followed by different small letters mean significant difference among the sowing dates for the same cultivar (P < 0.05).

表 6 开花期单位面积可孕花数与同化物积累和分配、干物质和氮积累速率的相关分析

Table 6 Correlation between fertile florets number and photosynthetic accumulation and distribution and accumulative rate of dry matter and nitrogen at anthesis stage

指标Indices X1 X2 X3 X4 X5 X6 X7 X8 X9 X10 X11 X12 X1 1 X2 –0.295 1 X3 0.910** –0.611** 1 X4 0.036 0.670** –0.312 1 X5 –0.604** 0.821** –0.856** 0.740** 1 X6 0.935** –0.539* 0.984** –0.279 –0.829** 1 X7 0.750** –0.574** 0.67** –0.939** –0.677** 0.240 1 X8 0.826** –0.698** 0.969** –0.445 –0.913** 0.938** 0.410 1 X9 –0.016 –0.884** 0.289 –0.748** –0.657** 0.626** 0.688** 0.436 1 X10 0.408 –0.549* 0.139 –0.722** –0.352 0.590** 0.680** 0.025 0.842** 1 X11 0.757** –0.546* 0.910** –0.386 –0.744** 0.915** 0.567** 0.931** 0.561* 0.499* 1 X12 0.996** –0.284 0.900** –0.041 –0.600** 0.934** 0.766** 0.814** 0.713** 0.723** 0.854** 1 注（Note）：X1—穗干重 Pike dry weight；X2—茎干重 Stem dry weight；X3—干重穗茎比 Dry weight ratio of spike to stem；X4—穗氮素积累量 Spike N accumulation；X5—茎氮积累量 Stem N accumulation；X6—氮积累量穗茎比 N accumulation ratio of spike to stem；X7—整株干物质积累速率 Accumulative rate of dry matter in the whole plant；X8—穗干物质积累速率 Accumulative rate of dry matter in spike；X9—整株氮积累速率 Accumulative rate of N in the whole plant；X10—穗氮积累速率 Accumulative rate of N in spike；X11—小花存活率 Survival rate of fertile florets；X12—单位面积可孕花数 Number of fertile florets per square meter；*—P < 0.05；**—P < 0.01.

表 7 播期对小麦单位面积无效分蘖和有效茎蘖退化小花数以及可孕小花和退化小花GDD生产效率的影响

Table 7 Effect of sowing dates on the number of degenerated florets in ineffective tillers (IT) and effective main shoots and tillers (EMT) and the GDD production efficiency of fertile florets (FF) and degenerated florets (DF) of winter wheat

品种
Cultivar 播期
Sowing date
(m-d) 退化小花数 (× 104 No./m2)
Number of degenerated floret GDD生产效率 [No./(m2·℃ d)]
Production efficiency of GDD 无效分蘖 IT 有效茎蘖 EMT 可孕小花 FF 退化小花 DF T18 09-24 (S1) 8.93 a 5.09 a 20.70 c 70.80 a 10-01 (S2) 8.30 a 4.78 a 22.78 c 71.63 a 10-08 (S3) 7.25 b 4.39 b 25.00 b 68.60 ab 10-15 (S4) 6.29 c 4.02 b 26.66 ab 64.92 bc 10-22 (S5) 5.20 d 3.66 c 28.48 a 60.47 c J22 09-24 (S1) 8.41 a 6.31 a 14.70 c 74.40 a 10-01 (S2) 8.06 ab 5.94 ab 16.19 b 76.67 a 10-08 (S3) 7.42 b 5.51 b 17.82 b 74.70 a 10-15 (S4) 6.41 c 5.10 c 19.23 ab 70.92 a 10-22 (S5) 4.43 d 4.58 c 20.28 a 61.44 b 注（Note）：数据为两生长季平均值 Values were the mean of the two growing seasons. 同列数据后不同小写字母表示同一品种不同播期间差异显著 (P < 0.05) Values followed by different small letters mean significant difference among the sowing dates for the same cultivar (P < 0.05). [1] 吕晓康, 温晓霞, 廖允成, 刘杨. 外源多胺对小麦小花退化的调控机制[J]. 作物学报, 2016, 42(9): 1391–1401.

Lv X K, Wen X X, Liao Y C, Liu Y. Effect of exogenous polyamines on mechanism of floret degeneration in wheat[J]. Acta Agronomica Sinica, 2016, 42(9): 1391–1401.

[2] 冯伟森, 张学品, 吴少辉, 等. 不同播期对洛旱7号小麦幼穗分化及产量的影响[J]. 河南农业科学, 2011, 40(10): 32–34. DOI: 10.3969/j.issn.1004-3268.2011.10.009

Feng W S, Zhang X P, Wu S H, et al. Effects of different sowing dates on spike characteristics and yield of Luohan 7[J]. Journal of Henan Agricultural Sciences, 2011, 40(10): 32–34. DOI: 10.3969/j.issn.1004-3268.2011.10.009

[3] 李存东, 曹卫星. 小麦阶段发育生理生态特征评述[J]. 南京农业大学学报, 1997, 20(2): 17–21.

Li C D, Cao W X. A review on eco-physiological characterization of wheat phasic development[J]. Journal of Nanjing Agricultural University, 2011, 40(10): 32–34.

[4] 郑春风, 朱慧杰, 朱云集, 等. 冬小麦小花发育及结实特性对叶面喷硼的响应[J]. 植物营养与肥料学报, 2016, 22(2): 550–556.

Zheng C F, Zhu H J, Zhu Y J, et al. Responses of floret development and grain setting characteristics of winter wheat to foliar spray boron[J]. Journal of Plant Nutrition and Fertilizer, 2016, 22(2): 550–556.

[5] 朱云集, 崔金梅, 王晨阳, 等. 小麦不同生育时期施氮对穗花发育和产量的影响[J]. 中国农业科学, 2002, 35(11): 1325–1329. DOI: 10.3321/j.issn:0578-1752.2002.11.005

Zhu Y J, Cui J M, Wang C Y, et al. Effects of nitrogen application at different wheat growth stages on floret development and grain yield of winter wheat[J]. Scientia Agricultura Sinica, 2002, 35(11): 1325–1329. DOI: 10.3321/j.issn:0578-1752.2002.11.005

[6]

Serrago A R, Miralles D J, Slafer G A. Floret fertility in wheat as affected by photoperiod during stem elongation and removal of spikelets at booting[J]. European Journal of Agronomy, 2008, 28: 301–308. DOI: 10.1016/j.eja.2007.08.004

[7]

González F G, Terrile I I, Falcón M O. Spike fertility and duration of stem elongation as promising traits to improve potential grain number (and yield): variation in modern argentinean wheats[J]. Crop Science, 2011, 51: 1693–1702. DOI: 10.2135/cropsci2010.08.0447

[8] 王志敏, 王树安, 苏宝林. 小麦穗粒数的调节Ⅱ开花前遮光对穗碳水化合物代谢和内源激素水平的影响[J]. 华北农学报, 1997, 12(4): 42–47. DOI: 10.3321/j.issn:1000-7091.1997.04.009

Wang Z M, Wang S A, Su B L. Regulation of grain number in wheatⅡ effects of shading on carbohydrate metabolism and hormone levels in spikes before anthesis[J]. Acta Agriculturae Boreali-Sinica, 1997, 12(4): 42–47. DOI: 10.3321/j.issn:1000-7091.1997.04.009

[9]

Slafer G A, Abeledo L G, Miralles D J, et al. Photoperiod sensitivity during stem elongation as an avenue to raise potential yield in wheat[J]. Euphytica, 2001, 119: 191–197.

[10]

Bancal P. Positive contribution of stem growth to grain number per spike in wheat[J]. Field Crops Research, 2008, 105(1–2): 27–39.

[11]

Miralles D J, Katz S D, Colloca A, et al. Floret development in near isogenic wheat lines differing in plant height[J]. Field Crops Research, 1998, 59: 21–30. DOI: 10.1016/S0378-4290(98)00103-8

[12] 王兆龙, 曹卫星, 戴廷波, 周琴. 不同穗型小麦品种小花发育与结实特性研究[J]. 南京农业大学学报, 2000, 23(4) : 9–12.

Wang Z L, Cao W X, Dai T B, Zhou Q. Characteristics of floret development and grain set in three wheat genotypes of different spike sizes[J]. Journal of Nanjing Agricultural University, 2000, 23(4): 9–12.

[13] 文祥朋, 任伟, 孙克刚, 等. 不同穗型小麦小花发育过程中幼穗内同化物分配与穗粒数的关系[J]. 江西农业学报, 2017, 29(8): 11–14.

Wen X P, Ren W, Sun K G, et al. Relationship between photosynthate distribution in young ear at floret development stage and grain number per ear of different ear-type wheat varieties[J]. Acta Agriculturae Jiangxi, 2017, 29(8): 11–14.

[14] 于振文. 不同密度条件对冬小麦小花发育的影响[J]. 作物学报, 1984, 10(3): 185–194.

Yu Z W. Influence of different density conditions on floret development of winter wheat[J]. Acta Agronomica Sinica, 1984, 10(3): 185–194.

[15] 屈会娟, 李金才, 沈学善. 播种密度对冬小麦不同穗位与粒位结实粒数和粒重的影响[J]. 作物学报, 2009, 35(10): 1875–1883.

Qu H J, Li J C, Shen X S. Effects of plant density on grain number and grain weight at different spikelets and grain positions in winter wheat cultivars[J]. Acta Agronomica Sinica, 2009, 35(10): 1875–1883.

[16]

Demotes-Mainard S, Jeuffroy M H. Effects of nitrogen and radiation on dry matter and nitrogen accumulation in the spike of winter wheat[J]. Field Crops Research, 2004, 87(23): 221–233.

[17]

Ferrante A, Savin A, Slafer G A. Floret development and grain setting differences between modern durum wheat under contrasting nitrogen availability[J]. Journal of Experimental Botany, 2013, 64: 169–184. DOI: 10.1093/jxb/ers320

[18]

González F G, Slafer G A, Miralles D J. Floret development and spike growth as affected by photoperiod during stem elongation in wheat[J]. Field Crops Research, 2003, 81: 29–38. DOI: 10.1016/S0378-4290(02)00196-X

[19]

Dreccer M F, Wockner K B, Palta J A, et al. More fertile florets and grains per spike can be achieved at higher temperature in wheat lines with high spike biomass and sugar content at booting[J]. Functional Plant Biology, 2014, 41: 482–495. DOI: 10.1071/FP13232

[20]

Dai X L, Wang Y C, Dong S X, et al. Delayed sowing can increase lodging resistance while maintaining grain yield and nitrogen use efficiency in winter wheat[J]. The Crop Journal, 2017, 5: 541–552. DOI: 10.1016/j.cj.2017.05.003

[21]

Waddington S R, Cartwright P M, Wall P C. A quantitative scale of spike initial and pistil development in barley and wheat[J]. Annals of Botany, 1983, 51: 119–130. DOI: 10.1093/oxfordjournals.aob.a086434

[22]

Ferrante A, Savin R, Slafer G A. Floret development and grain setting differences between modern durum wheat under contrasting nitrogen availability[J]. Journal of experimental botany, 2012, 64(1): 169–184.

[23]

Zheng C, Zhu Y, Zhu H, et al. Floret development and grain setting characteristics in winter wheat in response to pre-anthesis applications of 6-benzylaminopurine and boron[J]. Field Crops Research, 2014, 169: 70–76. DOI: 10.1016/j.fcr.2014.09.005

[24] 张小燕, 宋哲民. 关中小麦品种产量构成因素的相互关系和产量育种目标[J]. 西北植物学报, 1996, 16(1): 81–87. DOI: 10.3321/j.issn:1000-4025.1996.01.015

Zhang X Y, Song Z M. Relationship of yield and its components and breeding targets for wheat cultivars in Shaanxi Central Plain[J]. Acta Botanica Boreali-occidentalis Sinica, 1996, 16(1): 81–87. DOI: 10.3321/j.issn:1000-4025.1996.01.015

[25] 朱慧杰, 郑春风, 张国钊, 等. 栽培管理模式对冬小麦小花发育与结实特性的影响[J]. 麦类作物学报, 2014, 34(10): 1383–1389. DOI: 10.7606/j.issn.1009-1041.2014.10.011

Zhu H J, Zheng C F, Zhang G Z, et al. Effects of cultivation modes on the characteristics of floret development and grain set of winter wheat[J]. Journal of Triticeae Crops, 2014, 34(10): 1383–1389. DOI: 10.7606/j.issn.1009-1041.2014.10.011

[26] 沙之敏, 边秀举, 郑伟, 等. 最佳养分管理对华北冬小麦养分吸收和利用的影响[J]. 植物营养与肥料学报, 2010(5): 1049–1055.

Sha Z M, Bian X J, Zheng W, et al. Effects of optimum nutrient management on nutrient uptake and utilization of winter wheat in North China Plain[J]. Journal of Plant Nutrition and Fertilizer, 2010(5): 1049–1055.

[27]

Kuiper D. Sink strength: established and regulated by plant growth regulators[J]. Plant, Cell & Environment, 1993, 16, 1025–1026.

[28]

Roitsch T, Ehneß R, Goetz M, et al. Regulation and function of extracellular invertase from higher plants in relation to assimilate partitioning, stress responses and sugar signaling[J]. Function Plant Biology, 2000, 27, 815–825. DOI: 10.1071/PP00001

[29] 马鑫, 代兴龙, 王晓婧, 等. 冬小麦高产高效群体的年际间稳产性能[J]. 应用生态学报, 2017, 28(12): 3926–3934.

Ma X, Dai X L, Wang X J, et al. Stability of a winter wheat population with high yield and high resource use efficiency[J]. Chinese Journal of Applied Ecology, 2017, 28(12): 3926–3934.

[30] 赵广才. 小麦优势蘖利用超高产栽培技术研究[J]. 中国农业科技导报, 2007, 9(2): 44–48. DOI: 10.3969/j.issn.1008-0864.2007.02.008

Zhao G C. Studies on super-high yield cultivation technique using superior tiller in wheat[J]. Review of China Agricultural Science and Technology, 2007, 9(2): 44–48. DOI: 10.3969/j.issn.1008-0864.2007.02.008

[31] 宋欣欣, 贺德先. 小麦生育后期主茎和分蘖次生根对籽粒产量和品质的影响[J]. 麦类作物学报, 2011, 31(2): 281–285

Song X X, He D X. Effect of culm and tiller nodal roots at late growing period on grain yield and quality of wheat (Triticum aestivum L.)[J]. Journal of Triticeae Crops, 2011, 31(2): 281–285

[32]

Guo Z F, Schnurbusch T. Variation of floret fertility in hexaploid wheat revealed by tiller removal[J]. Journal of Experimental Botany, 2015, 66, 5945–5958. DOI: 10.1093/jxb/erv303

[33] 王晓宇, 冯伟, 郭天财, 等. 两种穗型小麦品种分蘖衰亡进程中茎蘖碳氮代谢的差异[J]. 西北农业学报, 2010(11): 38–42. DOI: 10.3969/j.issn.1004-1389.2010.11.008

Wang X Y, Feng W, Guo T C, et al. Difference of carbon and nitrogen metabolism in leaves between main caulis and tillers during tiller senescence of two spike-type winter wheat[J]. Acta Agriculturae Boreali-occidentalis Sinica, 2010(11): 38–42. DOI: 10.3969/j.issn.1004-1389.2010.11.008

[34] 徐丽娜, 冯伟, 盛坤, 等. 小麦兰考矮早八茎蘖幼穗激素差异及其密度调控效应[J]. 中国农业科学, 2011, 44(6): 1283–1291. DOI: 10.3864/j.issn.0578-1752.2011.06.025

Xu L N, Feng W, Shen K, et al. Difference of endogenous hormones in young spike between main stem and tillers and its effects under different densities in Lankao Aizao 8[J]. Scientia Agricultura Sinica, 2011, 44(6): 1283–1291. DOI: 10.3864/j.issn.0578-1752.2011.06.025

[35] 蔡大同, 王义炳, 茆泽圣, 等. 不同生态条件下播期和氮肥对优质小麦产量和品质性状的影响[J]. 植物营养与肥料学报, 1994, 1: 74–82. DOI: 10.11674/zwyf.1994.0110

Cai D T, Wang Y B, Mao Z S, et al. The effect of sowing date and nitrogen application under different ecosystem on the yield and quality of the superior wheat variety[J]. Journal of Plant Nutrition and Fertilizer, 1994, 1: 74–82. DOI: 10.11674/zwyf.1994.0110

[36] 张敏, 王岩岩, 蔡瑞国, 等. 播期推迟对冬小麦产量形成和籽粒品质的调控效应[J]. 麦类作物学报, 2013, 33(2): 325–330.

Zhang M, Wang Y Y, Cai R G, et al. Regulating effect of delayed sowing date on yield formation and grain quality of winter wheat[J]. Journal of Triticeae Crops, 2013, 33(2): 325–330.