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播期和种植密度对油菜产量和茎秆抗倒性的影响

本站小编 Free考研考试/2021-12-26
袁圆,1, 汪波1, 周广生1, 刘芳2, 黄俊生3, 蒯婕,11华中农业大学植物科学技术学院,武汉 430070
2全国农业技术推广服务中心,北京 100125
3建始县畜牧技术推广站,湖北恩施 445300

Effects of Different Sowing Dates and Planting Densities on the Yield and Stem Lodging Resistance of Rapeseed

YUAN Yuan,1, WANG Bo1, ZHOU GuangSheng1, LIU Fang2, HUANG JunSheng3, KUAI Jie,11College of Plant Science and Technology, Huazhong Agricultural University, Wuhan 430070
2National Agricultural Technology Extension Service Center, Beijing 100125
3Animal Husbandry Technology Extension Service Station, Enshi 445300, Hubei

通讯作者: * 蒯婕,E-mail： kuaijie@mail.hzau.edu.cn

责任编辑: 杨鑫浩
收稿日期:2020-06-22接受日期:2020-08-17网络出版日期:2021-04-16
基金资助:国家重点研发计划.2018YFD1000900
湖北省技术创新专项重大项目.2017ABA064


Received:2020-06-22Accepted:2020-08-17Online:2021-04-16
作者简介 About authors
袁圆,E-mail： 2788049152@qq.com。








摘要
【目的】茎秆倒伏是制约我国油菜生产效益提高的重要因素,研究不同播期及密度下油菜茎秆抗倒性变化规律及其生理机制,为油菜高产抗倒栽培提供理论及技术支撑。【方法】本研究以华油杂62和沣油520为材料,设置2个播期（9月25日、10月25日）和4个密度（15×104、30×104、45×104和60×104 株/hm2）裂区试验,测定产量及其构成,茎秆抗折力、倒伏指数、显微结构、主要成分及木质素合成关键酶活性等指标。【结果】（1）9月25日播种（T1）,密度从15×104hm-2增至60×104hm-2,油菜单株产量、单株角果数及每角粒数均下降,小区产量在45×104 hm-2处理达峰值,此时倒伏指数最小,抗倒能力最强,产量及抗倒性协同提高;播期推迟至10月25日（T2）,在任何密度下,小区产量、单株产量、单株角果数及每角粒数均显著降低,但地上部鲜重下降更明显,导致迟播油菜的倒伏指数下降、抗倒性增强。（2）适期播种时,密度增大,株高和茎秆干重均显著降低,倒伏指数呈先降后增的趋势,易倒伏部位从主茎中上部转移至主茎中下部,茎秆维管束长度/髓腔外组织宽度和维管束面积/茎横截面积等指标参数逐渐增加,茎秆木质素和纤维素含量呈先增后降趋势。油菜播期从T1推迟至T2,株高和茎秆干重均显著降低,茎秆木质素、纤维素含量显著下降,但植株地上部鲜重降幅较大,倒伏指数下降,抗倒性增强。逐步回归分析表明木质素是改善输导组织结构、协调倒伏指数及小区产量的关键指标,茎秆木质素含量及群体木质素总量高,可同时获得较强的茎秆抗倒性及较高的小区产量。（3）适期播种,密度从15×104hm-2增至60×104hm-2时,与木质素合成相关的过氧化物酶（POD）、肉桂醇脱氢酶（CAD）、苯丙氨酸解氨酶（PAL）及4-香豆酰-CoA连接酶（4CL）酶活性增强,油菜播期从T1推迟至T2,木质素合成酶POD、CAD、PAL、4CL的活性均显著降低。【结论】不同播期条件下,优化种植密度,可显著提高油菜的群体产量;且播期推迟,可通过进一步增大种植密度弥补单株产量的不足,晚播密植条件下茎秆木质素合成能力增强,木质素含量增加,协调了高产和抗倒的矛盾。
关键词： 油菜;播期;密度;抗倒性;产量

Abstract
【Objective】Stem lodging is an important factor that restricts the increase of rapeseed production efficiency in China. The purpose of this study was to investigate the changing discipline and physiological mechanism of the lodging resistance of rapeseed stem treated with different sowing dates and densities, so as to provide theoretical and technical supports for high-yield and lodging resistance of rapeseed cultivation. 【Method】A split-plot experiment with two canola varieties (Huayouza 62 and Fengyou 520), two sowing dates (September 25, October 25) and four densities (15×104, 30×104, 45×104, and 60×104 plants/hm2)was performed to investigate yield and yield composition, stem mechanical strength, lodging index, microstructure, the main components of stem and key enzyme activities of lignin synthesis. 【Result】(1) At the sowing date of September 25 (T1), yield per plant, effective pods per plant and seeds per pod decreased with the density increased from 15×104 hm-2 to 60×104 hm-2, and the highest yield with the lowest lodging index was observed at density of 45×104 hm-2; Under the delayed the sowing date to October 25 (T2), the population yield, yield per plant, effective pods per plant and seeds per pod decreased significantly compared with those under T1, but the above-ground fresh weight decreased more significantly, resulting in the decrease of lodging index and the increase of lodging resistance under delayed sowing dates; (2)When sowing at the appropriate time, both the plant height and stem dry weight decreased significantly, while the lodging index was with a tendency of increasing firstly and then decreasing with increased densities, and the susceptible lodging part was transferred from the upper part to the base part of rapeseed stem. The parameters such as vascular bundle length/the thickness outside the pith and vascular bundle area/stem cross-sectional area gradually increased, and the content of lignin and cellulose of stem increased first and then decreased. With sowing dates delayed from September 25 to October 25, plant height, stem dry weight, stem lignin and cellulose content decreased significantly, but the above-ground fresh weight decreased more, resulting in the decreased lodging index. Stepwise regression analysis showed that lignin was the key index to improve the structure of the transport tissue, coordinate the lodging index and the population yield. The higher lignin content and the population lignin content of the stem could simultaneously obtain stronger lodging resistance of the stem and a higher population yield. (3) When the density increased from 15×104 hm-2 to 60×104 hm-2, the activities of peroxidase (POD), cinnamyl alcohol dehydrogenase (CAD), phenylalanine ammonia lyase (PAL) and 4-coumaryl: CoA ligase (4CL) under T1 increased, and those decreased significantly with sowing date delayed from T1 to T2. 【Conclusion】Optimizing planting density under different sowing dates could significantly increase the population yield, and the decreased yield at the delayed sowing date could be compensated by increasing planting density. The ability of lignin synthesis enhanced, and the lignin content increased under delaying sowing dates with higher plant densities, and coordinated the contradiction between high yield and lodging resistance in rapeseed.
Keywords：rapeseed;sowing date;planting density;lodging resistance;yield


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本文引用格式
袁圆, 汪波, 周广生, 刘芳, 黄俊生, 蒯婕. 播期和种植密度对油菜产量和茎秆抗倒性的影响[J]. 中国农业科学, 2021, 54(8): 1613-1626 doi:10.3864/j.issn.0578-1752.2021.08.004
YUAN Yuan, WANG Bo, ZHOU GuangSheng, LIU Fang, HUANG JunSheng, KUAI Jie. Effects of Different Sowing Dates and Planting Densities on the Yield and Stem Lodging Resistance of Rapeseed[J]. Scientia Acricultura Sinica, 2021, 54(8): 1613-1626 doi:10.3864/j.issn.0578-1752.2021.08.004


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0 引言

【研究意义】目前,我国食用植物油自给率已降至30%左右,对外依存度进一步加大。油菜是我国第五大作物,也是我国最主要油料作物,常年种植面积约760万hm2;菜籽油约占我国国产食用植物油的55%,是当前我国居民食用油的重要来源[1]。提高油菜产量与种植效益、扩大生产规模是缓解我国食用植物油供给矛盾的重要举措。但实际生产中,各产区油菜普遍发生的茎秆倒伏现象,不仅限制了油菜产量的进一步提高,而且制约了机械化收获技术的应用,导致油菜产量、效益及种植面积长期徘徊不前[2,3]。因此,加强油菜抗倒机理研究,形成配套技术,可提高油菜生产效益,推动油菜生产的发展。【前人研究进展】作物倒伏包括根倒和茎倒2种类型[4]。长江流域是我国油菜主产区,该区油菜以稻-油轮作为主,且主要采用直播栽培技术。直播油菜主要以茎倒为主,一般在终花后20 d左右发生。受前茬水稻收获期、油菜播种期气候条件、土壤墒情、整地技术等的影响,即使在相同产区,油菜播种出苗差异大,导致种植密度存在较大差异;一般而言,油菜肥料施用量及施用方式较固定。因此,除品种及气候因素外,播期与密度是影响我国长江流域主产区直播油菜倒伏的关键因素。油菜播期推迟,根颈粗、株高及茎秆抗折力均有所降低,且木质素和纤维素含量下降,茎秆倒伏指数增大[5,6];也有研究表明,播期推迟,作物生育进程缩短,且显著降低了株高、重心高度和基部节间长度,倒伏风险降低[7,8]。适期播种,水稻茎秆中K、Ca、Si及木质素等含量较高,倒伏指数下降[9]。播期推迟,作物干物质累积能力下降,地上部干重下降,单株产量显著降低[10]。密度是影响油菜产量及抗倒性另一个重要因素。密度适宜,个体、群体及高产、抗倒矛盾协调较好,油菜生产效益显著提高。其机理是油菜倒伏主要发生在茎秆中上部,适当增大密度,茎秆中上部倒伏指数降低,抗倒能力增强,可实现产量和抗倒性的协同提高[11,12]。纤维素和木质素对茎秆硬度和弹性起着重要作用,适当增加种植密度,油菜茎秆木质素、纤维素含量增加[13,14,15],抗倒性增强,且增产效应明显。PAL、4CL、CAD、POD是木质素合成的关键酶[16],适当增加密度,木质素合成关键基因表达量及关键酶活性均增加[11, 17-18]。因此,适当增加油菜种植密度,可实现“以密增产、以密补迟、以密省肥、以密控草、以密适机”的五大效应[10]。【本研究切入点】虽然前人在油菜农艺性状[19]、产量构成[20,21]、倒伏特性[22]等方面已有较多研究,但油菜抗倒机理相关研究仍不够系统和深入。【拟解决的关键问题】本研究设置不同播期和种植密度处理,拟从农艺性状、力学特性、理化性质及显微结构等层面较为系统地探究油菜茎秆抗倒性变化规律及内在机理,以期为油菜高产抗倒栽培提供理论及技术支撑。

1 材料与方法

1.1 试验设计

2017—2019连续2年在华中农业大学试验基地（30.52°N,114.31°E）进行。试验地前茬为水稻,2017年土壤碱解氮、速效磷及速效钾含量分别为86.34、16.25、154.51 mg·kg-1;2018年上述指标分别为95.7、10.2、165.0 mg·kg-1。试验裂区设计,以播期（9月25日、10月25日）为主区;品种（华油杂62和沣油520）为裂区,其中华油杂 62为甘蓝型半冬性细胞质雄性不育三系杂交种,抗倒性强;沣油 520为甘蓝型细胞质雄性不育三系杂交种,易倒伏;密度（15×104、30×104、45×104和60×104株/hm2）为裂裂区;3次重复,计48个小区。小区面积为20 m2（2 m×10 m）,基肥为复合肥（N﹕P2O5﹕K2O为15%﹕15%﹕15%）600 kg·hm-2和硼肥15 kg·hm-2,苗肥为氮含量为46%的尿素150 kg·hm-2。采用25 cm行距直播,油菜出苗后间去窝堆苗,3叶期至5叶期定苗,其他管理同常规大田。

1.2 测定指标与方法

1.2.1 产量及产量构成 成熟期各小区连续取样10株,考察单株有效角果数、每角粒数及千粒重;小区单独收获,测定实际产量。

1.2.2 倒伏相关指标 终花后20 d,各小区连续取样10株,将主茎有效分枝部位以下均分为2段,基部、上部分别标记为1段、2段,用YYD-1茎秆强度测定仪（浙江托普仪器有限公司）测定茎秆2段中间部位抗折力。倒伏指数（cm·g·g-1）=高度（cm）×鲜重（g）/抗折力（g）。此处高度与鲜重为测定茎段至植株顶部对应高度与鲜重。

1.2.3 茎秆化学成分 将终花后20 d主茎样品分装入牛皮纸袋中,105℃杀青30min后置85℃烘干至恒重,粉碎过筛,测定纤维素及木质素含量,方法参照文献[23,24]。

1.2.4 花期木质素及纤维素合成关键酶活性 花期在各小区中取主茎相同部位样品,液氮固定后置-80℃超低温冰箱保存,采用酶联免疫（ELISA）方法（江苏晶美实业有限公司）,测定苯丙氨酸解氨酶（PAL）、4-香豆酰-CoA连接酶（4CL）、肉桂醇脱氢酶（CAD）、过氧化物酶（POD）等木质素合成关键酶。具体步骤为：将样品用磷酸缓冲液10%组织匀浆（0.1 g鲜样加入0.9 mL磷酸缓冲液）,37℃反应30 min,洗板5次,加入酶试剂,37℃反应30 min,洗板5次,加入显色液,37℃反应10 min,加入终止液,450 nm读取OD值。用标准物的浓度与OD值计算出标准曲线的直线回归方程,将样品的OD值代入方程,计算出样品浓度,再乘以稀释倍数,即为样品的实际浓度。

1.2.5 花期茎秆显微结构测定 于初花期选取适播期（T1）油菜单株,切取易倒伏部位,即第一有效分枝以下5 cm处的主茎段,置 FAA 固定液中,用番红-固绿染色后制作切片,用Pannoramic 250、Pannoramic MIDI切片扫描仪（3D Histech, Hungary）扫描成像,采用CaseViewer软件分析维管束数量,观察细胞形态、排列方式。

1.2.6 气象数据 气象数据来自华中农业大学试验基地气象站。

1.3 数据处理

采用SPSS10.0软件进行数据统计分析;采用Origin2018软件绘图。

2 结果

2.1 不同播期和密度下油菜生育期及气象因子差异

2017—2018和2018—2019年油菜生长季有效积温和降水量存在差异（表1）。与2017—2018年相比,2018—2019年总降雨量增加,有效积温下降。播期推迟,油菜全生育期缩短30 d左右。相同播期,增加种植密度,生育期缩短3—5 d,2个品种规律一致;相同处理下,华油杂62生育期比沣油520长7 d左右。华油杂62所需有效积温高于沣油520;9月25日（T1）播期的有效积温高于10月25日（T2）播期。

Table 1
表1
表12017—2019年油菜生育期及气象数据
Table 1Rapeseed growth process and meteorological factor during 2017-2019
播期
Sowing date		品种
Variety		密度
Density		2017-20182018-2019
生育期
Growth days (d)		有效积温
Effective accumulated temperature (℃)		降雨量
Rainfall
(mm)		生育期
Growth days (d)		有效积温
Effective accumulated temperature (℃)		降雨量
Rainfall
(mm)
T1华油杂62
HZ 62		D12182772.2207.32182669.9492.8
D22162727.5207.32162623.5492.8
D32142678.6206.32142581.4492.8
D42132653.4206.32132564.0492.8
沣油520
FY 520		D12102586.1206.12102519.2449.5
D22082548.0206.12082478.0449.5
D32062516.1199.32062433.3449.5
D42052494.2181.82052415.4448.4
T2华油杂62
HZ 62		D11912318.3183.01932010.4475.9
D21892275.3180.01921974.8453.1
D31882254.8180.01911949.8453.1
D41872234.6180.01901927.4453.1
沣油520
FY 520		D11852195.1144.21871995.8474.7
D21842172.7144.21861955.9474.7
D31832150.4144.21851933.3474.7
D41822126.3144.21841909.2474.7
D1、D2、D3 和 D4 分别表示密度为15×104、30×104、45×104和 60×104株/hm2;T1和T2分别表示9月25日和10月25日播种。下同
D1, D2, D3, and D4 indicate the planting densities of 15×104, 30×104, 45×104 , and 60×104 plants/hm2, respectively. T1 and T2 represent respectively sowing time on September 25 and October 25, respectively. The same as below

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2.2 不同播期和密度对油菜机械收获关键性状的影响

2.2.1 产量及产量构成 同一密度,播期推迟,2个品种的单株角果数、每角粒数及千粒重均下降,单株产量下降27.63%;同一播期,增加密度,2个品种的单株角果数和每角粒数下降,千粒重受密度影响无显著差异,密度增至60×104 hm-2,单株产量显著下降64.36%,2年结果一致（表2）。实际产量的变化在播期及品种间有所不同。9月25日播种（T1处理）,实际产量随密度增加呈先增后减的趋势,2个品种、2年度变化规律相近;10月25日播种（T2处理）,随密度增加,华油杂62实际产量呈增加趋势,沣油520呈先增后减趋势;2年度规律相近。方差分析表明,播期、密度和品种对单株角果数、每角粒数、单株产量和实际产量的影响达显著水平,对单株角果数和实际产量的三因素互作效应达到显著水平。

Table 2
表2
表2不同播期和密度对油菜产量及产量构成的影响
Table 2Effects of different sowing dates and planting densities on rapeseed yield and yield composition
年份
Year		播期
Sowing date		品种
Variety		密度
Density		单株角果数
Effective pods per plant		每角粒数
Seeds
per pod		千粒重
1000-seed
weight (g)		单株产量
Yield
per plant (g)		成株率
Survival
rate (%)		实际产量
Yield
(kg·hm-2)
2017-2018T1华油杂62
HZ 62		D1315.0b17.41a3.68a20.51a93.60a2742.3c
D2185.0ef17.00ab3.76a11.82d87.93bc2901.0b
D3156.8gh16.46bcd3.60a9.24e84.22cd3290.9a
D4103.7k16.14cde3.69a6.11h82.59de2649.1cd
沣油520
FY 520		D1372.4a16.88b3.18bc19.99a93.53a2555.7de
D2222.8d16.58bcd3.08bcd11.39d87.06bc2939.5b
D3172.6fg15.45fgh3.13bcd8.36f81.57de2731.0c
D4128.4ij15.03h3.04cde7.03g77.30fg2625.9cde
T2华油杂62
HZ 62		D1262.9c16.63bc3.26b14.25b94.51a1866.0i
D2171.8fg16.17cde3.23bc8.96ef88.57b2261.0gh
D3141.7hi15.88ef3.19bc7.19g83.99cd2467.1ef
D4106.9k15.01h3.19bc5.12i79.91ef2484.9def
沣油520
FY 520		D1274.3c16.07de3.02cde13.33c96.94a1857.7i
D2190.8e15.66efg2.92de8.73ef87.36bc2203.5gh
D3154.9h15.38fgh2.94de6.99g79.94ef2362.2fg
D4113.5jk15.29gh2.87e4.99i75.55g2126.8h
2018-2019T1华油杂62
HZ 62		D1284.0b16.26a3.58c17.19a97.60a2311.5c
D2173.6e16.00ab3.77ab10.45b93.10ab2704.5ab
D3139.6g15.77bcd3.79a8.28d84.50c2854.7a
D496.1ij15.49de3.70b5.52f83.30c2655.0b
沣油520
FY 520		D1312.6a16.16a3.30de16.60a95.20a2113.5d
D2182.6de15.80bcd3.26ef9.67bc93.30ab2660.3b
D3152.7fg15.50de3.29ef7.75de83.80c2534.6b
D4102.3ij15.34ef3.21 f5.06 fg80.70 cde2339.1 c
T2华油杂62
HZ 62		D1194.1 cd16.04ab3.21f10.00bc94.80a1384.5h
D2141.4fg15.95abc3.24ef7.31e84.40c1669.5fg
D3108.7hi15.66cd3.37d5.74f78.30def1792.0ef
D486.0j15.06f3.28ef4.26gh76.90ef1932.7e
沣油520
FY 520		D1206.0c15.73bcd2.94g9.39c89.70b1198.2i
D2157.4f15.32ef2.92g7.05e82.20cd1514.6gh
D3123.4h15.13f2.98g5.58f77.30ef1836.3ef
D489.8j14.74g2.95g3.90h75.60f1570.7g
方差分析Variance analyses
2017-2018T********NS**
V***********
D****NS******
T×V*****NSNSNS
T×D**NSNS**NS**
V×D**NSNS****
T×V×D***NSNSNS**
2018-2019T************
V***********
D************
T×VNS***NSNSNS
T×D**NSNS*****
V×D**NS**NSNS*
T×V×D**NSNSNSNS*
*, **表示在 0.05 和 0.01 水平显著差异,NS 表示差异不显著;T：播期;V：品种;D：种植密度;T×V, T×D, V×D, T×V×D分别表示因素间互作。下同
* and **, significant difference at 0.05 and 0.01 probability levels, respectively; NS: not significant; T: Sowing date; V: Variety; D: Density; T×V,T×D,V×D, T×V×D mean the interaction between factors. The same as below

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2.2.2 倒伏相关性状 同一密度,播期推迟,2个品种茎秆抗折力和地上部鲜重均显著下降,倒伏指数下降;同一播期,密度增大,2个品种茎秆抗折力和地上部鲜重均显著下降,T1、T2处理,2个品种倒伏指数分别在密度为45×104 hm-2、60×104 hm-2达最小值;相同条件下,华油杂62的茎秆抗折力和地上部鲜重均高于沣油520,倒伏指数低于沣油520;2年规律一致（图1）。同一植株,茎秆基部抗折力及地上部鲜重高于上部,基部倒伏指数低于上部;同一播期,密度增加,茎秆基部倒伏指数增加,茎秆上部倒伏指数降低,且上部倒伏指数变幅大于基部;同一密度,播期推迟,茎秆基部和上部倒伏指数均降低,且上部倒伏指数降幅大于基部;品种间和年际间趋势一致。方差分析表明,播期、密度和品种对茎秆抗折力、地上部鲜重和倒伏指数的影响均达显著水平;播期、密度及品种对茎秆抗折力的三因素互作效应达显著水平,对地上部鲜重及倒伏指数的互作效应未达显著水平。

图1

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图1不同播期和密度对油菜茎秆机械强度、地上鲜重和倒伏指数的影响

D1、D2、D3 和 D4 分别表示种植密度为 15×104、30×104、45×104和 60×104株/hm2;T1和T2分别表示9月25日和10月25日播种时间; HZ62和FY520分别表示品种为华油杂62和沣油520。下同
Fig. 1Effects of different sowing dates and planting densities on mechanical strength, above-ground fresh weight and lodging index of rapeseed stem

D1, D2, D3, and D4 indicate the planting densities of 15×104, 30×104, 45×104 and 60×104 plants/hm2, respectively; T1 and T2 represent respectively sowing time on September 25 and October 25. HZ62 and FY520 represent respectively varieties of huayouza 62 and fengyou 520. The same as below


2.3 播期和密度对茎秆发育的影响

2.3.1 关键生育时期茎秆高度、干重变化 播期和密度影响着2个品种各生育期的茎秆高度及干重,导致成熟期株高及茎秆干重存在差异（图2）。同一密度、播期推迟,及同一播期、密度增加,2个品种成熟期株高及茎秆干重均显著降低,2年规律一致。同一密度,播期推迟,2个品种苗期至薹期株高增幅显著提高,茎秆干重增幅降低;同一播期,密度增加,2个品种苗期至薹期茎秆高度及干重增加幅度均显著降低。薹期—花期是油菜茎秆高度及干重变化最大的时期。同一密度、播期推迟,及同一播期、密度增加,2个品种薹期至花期茎秆高度及干重增幅均显著降低,2年规律一致。同一密度,播期推迟,2个品种花期至成熟期茎秆高度及干重增幅显著降低;同一播期,密度增加,2个品种花期至成熟期茎秆高度及干重增幅均显著降低。

图2

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图2不同播期和密度对株高和茎秆干重的影响

SS、BS、FS和MS分别表示苗期、蕾薹期、花期、角果期
Fig. 2Effects of different sowing dates and planting densities on plant height and stem dry weight

SS, BS, FS and MS represent seedling stage, bolting stage, flowering stage, and maturity stage, respectively


2.3.2 密度对茎秆结构的影响 密度增加,2个品种茎秆维管束数目和面积均逐渐减小,维管束长度/髓腔外组织宽度和维管束面积/茎横截面积均逐渐增大。相同密度条件下,华油杂62的维管束数目和面积、维管束长度/髓腔外组织宽度和维管束面积/茎横截面积均大于沣油520。方差分析表明,密度和品种对维管束面积、维管束长度/髓腔外组织宽度和维管束面积/茎横截面积的影响均达显著水平;密度及品种对维管束数目和面积、维管束长度/髓腔外组织宽度和维管束面积/茎横截面积的互作效应未达显著水平（图3）。

图3

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图3不同密度下油菜花期茎秆显微结构特点

Fig. 3Microstructural characteristics of rapeseed stem at different planting densities



2.3.3 播期和密度对茎秆主要成分的影响 同一密度,播期推迟,2个品种茎秆酸不溶木质素、酸溶木质素、总木质素、半纤维素、纤维素和可溶性糖含量均显著下降;同一播期,密度增大,2个品种酸不溶木质素、总木质素、半纤维素和纤维素含量均呈先增后降的趋势,2个品种酸溶木质素和可溶性糖含量呈先降后增的趋势（表3）。T1及T2处理,2个品种茎秆酸不溶木质素、总木质素、半纤维素和纤维素分别在密度为45×104 hm-2、60×104 hm-2达最大值,2个品种酸溶木质素和可溶性糖含量分别在45×104 hm-2、60×104 hm-2的密度达最小值。相同处理条件下,华油杂62茎秆酸不溶木质素、总木质素、半纤维素、纤维素及可溶性糖含量均高于沣油520;酸溶木质素含量低于沣油520;2年规律一致。方差分析表明,播期、密度和品种对酸不溶木质素、总木质素、可溶性糖、半纤维素、纤维素含量影响均达显著水平,播期、密度和品种对酸不溶木质素、酸溶木质素、总木质素、半纤维素、纤维素及可溶性糖的三因素互作效应未达到显著水平。

Table 3
表3
表3不同播期和密度对油菜茎秆主要成分的影响
Table 3Effects of different sowing dates and planting densities on main composition of rapeseed stem (%)
年份
Year
播期
Sowing date		品种
Variety
密度
Density		酸不溶木质素
Acid insoluble lignin		酸溶木质素
Soluble
lignin		总木质素
Total
lignin		半纤维素
Hemicel
lulose		纤维素
Cellulose		可溶性糖
Soluble
sugar
2017-2018T1华油杂62
HZ 62		D124.00bc1.73abc25.73abcd19.03cde34.74bc3.44a
D224.23abc1.70abc25.93abc20.11b36.00ab3.24b
D324.96a1.57c26.53a21.13a37.25a3.05c
D423.38cde1.73abc25.11cde18.75def35.98ab3.40a
沣油520
FY 520		D122.43fg1.83ab24.26fg18.66def32.78cd3.29b
D223.74bcd1.80ab25.54bcd18.61def34.84bc3.11c
D324.53ab1.74abc26.26ab19.84bc35.61ab3.06c
D423.78bcd1.77ab25.55bcd19.33bcd34.78bc3.27b
T2华油杂62
HZ 62		D121.84g1.73abc23.57g16.96hi29.14e3.08c
D222.72ef1.72abc24.43efg17.56gh30.89de2.86e
D324.03bc1.68abc25.71abcd17.98fg31.51d2.71f
D424.48ab1.63bc26.11ab18.09efg32.61d2.68f
沣油520
FY 520		D121.73g1.87a23.59g16.91hi22.32h2.96d
D222.32fg1.82ab24.14fg17.22ghi23.79gh2.72f
D323.08def1.82ab24.90def16.301i25.46fg2.70f
D423.36cde1.81ab25.17cde17.95fg27.09f2.63f
2018-2019T1华油杂62
HZ 62		D124.43b2.03bcde26.46b18.32abc27.97cd5.50a
D225.43a1.99cdef27.42a18.90a28.85bc4.87c
D325.66a1.94efg27.60a19.25a32.45a4.06h
D422.58cd2.11ab24.68cd17.10defg27.88cd4.41f
沣油520
FY 520		D120.96fgh2.06abcd23.02efg17.38cdef26.70de4.96b
D222.44cd1.98def24.42cd18.10abcd29.06bc4.49e
D323.20c1.85gh25.05c18.36abc30.59b3.21l
D421.22efg2.01bcde23.23ef17.53bcde28.96bc4.33f
T2华油杂62
HZ 62		D120.23hi1.91fgh22.14gh17.44cde26.36de5.04b
D221.17efgh1.88gh23.05efg17.62bcde26.75de4.57d
D322.04de1.86gh23.90de18.21abcd28.14cd4.18g
D423.40c1.76i25.16c18.63ab28.90bc3.76j
沣油520
FY 520		D119.73i2.13a21.87h15.13h23.71g4.80c
D220.55ghi2.08abc22.64fgh16.15g24.68fg3.95i
D321.84def1.87gh23.71de16.30fg26.01ef3.43k
D423.06c1.84hi24.90c16.64efg27.01de3.36k
方差分析Variance analyses
2017-2018T**NS********
V************
D**NS********
T×VNSNSNSNS**NS
T×D**NS*******
V×DNSNSNS**NS*
T×V×D**NS**NSNSNS
2018-2019T************
V************
D************
T×V**********NS
T×D************
V×DNS**NSNSNS**
T×V×DNSNSNSNSNS**

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2.3.4 茎秆主要成分与显微结构、倒伏指数及小区产量的关系 将茎秆主要组成成分与茎秆显微结构进行相关分析（图4）,结果表明,酸不溶木质素、总木质素、半纤维素及纤维素含量均与维管束长度/髓腔外组织宽度呈显著正相关。

图4

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图4茎秆显微结构与主要成分的相关关系

Fig. 4Correlation between microstructure of stem and main components



逐步回归所建立的最优回归方程包含了对因变量有显著影响的自变量,不包含对因变量没有显著影响的自变量[25],利用所建立的回归方程,筛选出一些对油菜抗倒性和产量有显著影响的指标。本试验中,将茎秆6个成分指标与倒伏指数进行逐步回归分析（表4）,2个播期的倒伏指数回归方程：Y1=1.975+0.412X2- 0.042X3,R2=0.314（T1）,Y1=2.614+0.285X2-0.078X3,R2=0.668（T2）。式中,Y1：倒伏指数;X2：酸溶木质素;X3：总木质素。T1和T2回归结果表明,酸溶木质素和总木质素对倒伏指数的影响远大于其他指标,在茎秆抗倒性中起着重要的作用。

Table 4
表4
表4茎秆主要成分与倒伏指数和小区产量的回归系数及显著性检验
Table 4Regression coefficients and significance tests between lodging index and yield with stem main components
播期
Sowing date		因变量
Dependent variable		自变量
Independent variable		回归系数
Regression coefficients		校准误
Calibration
error		标准系数
Standard coefficient		显著性Saliency
TP
T1倒伏指数
Lodging index		酸溶木质素soluble lignin0.4120.1240.4143.3340.002
总木质素Total lignin-0.0420.016-0.328-2.6430.011
小区产量
Yield		群体可溶性糖Group soluble sugar-2.3990.382-0.654-6.2830.000
群体总木质素Group total lignin0.6700.0850.8217.8890.000
T2倒伏指数
Lodging index		酸溶木质素soluble lignin-0.0780.013-0.636-6.2260.000
总木质素Total lignin0.2850.1050.2782.7210.009
小区产量
Yield		群体总木质素Group total lignin1.1440.0711.00116.0740.000
群体可溶性糖Group soluble sugar-3.3920.643-0.329-5.2740.000

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将茎秆6个成分群体值与小区产量进行逐步回归分析（表4）,2个播期的小区产量回归方程：Y2=136.532-2.399X6+0.670X3,R2=0.596（T1）;Y2=68.662-3.392X6+1.144X3,R2=0.846（T2）。式中,Y2：小区产量;X3：群体总木质素,X6：群体可溶性糖。T1和T2回归结果表明,群体总木质素和群体可溶性糖对小区产量的影响远大于其他指标。

2个播期条件下,茎秆倒伏指数及小区产量分别与茎秆主要成分的逐步回归分析结果表明,茎秆木质素指标均在2个方程中出现,且与倒伏指数为负效应,与小区产量为正效应。这说明,茎秆的各主要成分中,木质素是协调倒伏指数及小区产量的最重要的指标,茎秆木质素含量及群体木质素总量高,则可同时有较强的茎秆抗倒性及较高的小区产量。

2.3.5 播期和密度对木质素合成关键酶活性的影响 同一密度,播期推迟,2个品种的茎秆POD、CAD、PAL、4CL酶活性均降低;同一播期,密度增大,2个品种的茎秆POD、CAD、PAL、4CL酶活性均增加;相同处理条件下,华油杂62茎秆的POD、CAD、PAL、4CL酶活性均高于沣油520;且2年规律一致（图5）。方差分析表明,播期、密度和品种对茎秆POD、CAD、PAL、4CL酶活性影响达到显著水平;播期、密度和品种对茎秆CAD、PAL、4CL酶活性的互作效应未达到显著水平。

图5

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图5不同播期和密度对木质素合成有关的酶活性的影响

HZD1和HZD3分别表示华油杂62的种植密度为15×104和45×104株/hm2;FYD1和FYD4分别表示沣油520的种植密度为15×104和60×104株/hm2
Fig. 5Effects of different sowing dates and planting densities on enzyme activities related to lignin synthesis

HZD1 and HZD3 indicate huayouza 62 with the planting densities of 15×104 and 45×104 plants/hm2, respectively; FYD1 and FYD4 indicate fengyou 520 with the planting densities of 15×104 and 60×104 plants/hm2, respectively


3 讨论

本研究选用抗倒性强、但产量稍低的华油杂62和抗倒性较弱、但产量潜力较大的沣油520进行不同的密度及播期试验,探究密度和播期协调直播油菜产量和抗倒性的生理机制。这2个油菜品种代表了当前生产中众多主推品种的主要性状,品种的高产与抗倒性依然存在不小的矛盾,限制了油菜产量的进一步提高。因此,选用这2个品种进行相关试验,其结果具有较强的代表性。

3.1 不同播期和密度对油菜产量的影响

一定范围内的合理密植,虽然油菜的单株产量降低,但群体产量增加;适宜的种植密度亦与播期密切相关。这在较多的作物密度试验中均得到证实[26,27]。本试验2年的结果均表明,2个品种均随着种植密度的增加,油菜单株角果数、每角粒数下降,从而导致单株产量亦随之下降;而适期播种条件下（T1处理）,油菜群体产量随密度的增加呈先增后降的变化规律,推迟播种（T2处理）,群体产量随密度的增加逐渐增加。本试验中,常规肥料运筹模式下,T1和T2处理分别在45×104 hm-2、60×104 hm-2密度条件下,群体产量达最大值,说明在不同的播期条件下,优化种植密度,可以显著提高油菜的群体产量;且播期推迟,可通过进一步增大种植密度的方法来有效弥补单株产量的不足。

3.2 不同播期和密度对油菜茎秆抗倒性的影响

直播油菜根冠比较大,倒伏以茎秆倒伏为主,本研究亦关注各处理对茎秆抗倒性的变化。不同种植密度条件下,油菜植株的形态结构和茎秆成分等的改变最终显著影响着茎秆抗倒性。倒伏指数是衡量作物茎秆抗倒伏能力的一个重要的指标[11,12]。株高、地上部鲜重及茎秆抗折力显著影响作物的抗倒性。本研究表明,适期播种时（T1）,增大密度,油菜株高、地上部鲜重及茎秆抗折力均降低,茎秆抗倒伏能力呈先增后降的趋势。适宜密度条件下,株高及地上部鲜重乘积的降幅高于茎秆抗折力降幅,导致其倒伏指数下降,抗倒性增强。播期推迟（T2）,增大密度,油菜株高、地上部鲜重及茎秆抗折力均降低,但株高及地上部鲜重乘积的降幅高于茎秆抗折力降幅,抗倒性增强。

本研究结果表明,不同种植密度亦影响油菜茎秆易倒伏部位。从油菜茎秆不同部位的倒伏指数值可以看出,适期播种（T1）条件下,从D1至D2密度,油菜茎秆上下部位的倒伏指数值差异较大,易倒伏部位主要为茎秆上部（倒伏指数最大）;在D3密度条件下,茎秆上部下部倒伏指数接近;在D4密度条件下,基部倒伏指数增加,易倒伏部位主要在茎秆基部。这是因为油菜茎秆干重和茎秆高度发育进程前慢后快,低密度种植,苗期至薹期茎秆干物质较多,茎秆基部粗壮,薹期—花期是油菜茎秆高度及干重变化最大的时期,茎秆上部细弱,导致油菜易倒伏部位往往偏高;高密种植,油菜生育期缩短,茎秆发育进程加快,基部细弱,倒伏易发生部位降低;适宜种植密度,油菜茎秆发育比较均匀适度,茎秆不同部位粗细接近,抗倒性能接近。

3.3 密度和播期协调油菜产量和抗倒性的生理机制

综合不同播期下油菜产量和倒伏表现,可知适宜播期下,产量在45×104 hm-2处理达峰值,此时倒伏指数最小,抗倒能力最强,产量及抗倒性协同提高,而晚播条件下,60×104 hm-2密度时,产量较高,抗倒性强。茎秆是影响油菜倒伏和产量的关键部位,茎秆的生化成分与抗倒性密切相关,茎秆结构承担营养物质向籽粒的转运。本研究中,增大密度,茎秆中维管束长度/髓腔外组织宽度增加,木质素和纤维素含量亦增加。纤维素是植物细胞壁的骨架,对茎秆韧性起重要作用[28],而木质素直接影响茎秆强度[29],高抗倒伏品种木质素含量和茎秆抗折力均大于中抗倒伏和易倒伏品种[30]。本研究结果表明,增大密度,茎秆中的木质素和纤维素含量亦会增加,此时产量和抗倒性均增大;通过逐步回归分析,木质素是对油菜抗倒性和产量有显著影响的指标。较高的木质素含量,改善了茎秆输导组织结构,茎秆维管束长度/髓腔外组织宽度增加,抗倒性提高而增产。汪灿等[17]研究表明,茎秆木质素含量高,其茎秆机械组织层数多、维管束数目多且面积大,其抗倒伏能力强。茎秆木质素含量增加,改善其抗倒伏性,同时产量提高[31]。本研究逐步回归分析表明,个体条件下,较高的茎秆总木质素含量,较低的茎秆酸溶木质素含量,则茎秆抗倒性强;群体条件下,较高的茎秆总木质素总量,较低的可溶性糖含量,有较高小区产量。这说明,在茎秆的各主要成分中,木质素是协调倒伏指数及小区产量的最重要的指标,茎秆木质素含量及群体木质素总量高,可同时获得较强的茎秆抗倒性及较高的小区产量。

适当增加种植密度时,与木质素合成的相关酶的活性（POD、CAD、4CL、PAL）明显提高,导致茎秆木质素含量增加。有研究表明,维管束长度/髓腔外组织宽度与油菜茎秆抗倒伏性正相关[32]。本研究中,适当增加种植密度,茎秆中维管束长度/髓腔外组织宽度和维管束面积/茎横截面积增加,油菜茎秆抗倒伏能力增强,但此时茎秆变细,维管束数目和面积显著减小,单株产量下降,但油菜群体优势显现。因此,采用合理种植密度,可协同提高油菜的产量与抗倒性。播期推迟,茎秆中的木质素和纤维素相对含量减小,油菜单株产量和倒伏指数均减小,但合理密植亦可提高茎秆中的木质素含量,提高抗倒性,虽单株产量减小,但是群体密度增大,可达到以密增产、以密补迟的效果。

4 结论

一定范围内的合理密植,虽然油菜的单株产量降低,但群体产量增加。在适播期条件下,按45×104 hm-2种植密度,可显著提高油菜的群体产量;且播期推迟,种植密度可进一步增大至60×104 hm-2,来有效弥补单株产量的不足。适宜增密条件下,株高及地上部鲜重乘积的降幅高于茎秆抗折力降幅,导致其倒伏指数下降,抗倒性增强。不同种植密度亦影响油菜茎秆易倒伏部位,随着种植密度的增加,油菜茎秆易倒伏部位逐渐下移;一般来说,当种植密度为45×104 hm-2或60×104 hm-2,茎秆上部下部倒伏指数接近,抗倒性最强。不同种植密度条件下,油菜植株形态结构和茎秆成分的改变最终显著影响着茎秆抗倒性。在茎秆的各主要成分中,木质素是协调倒伏指数及小区产量的最重要的指标,茎秆木质素含量及群体木质素总量高,可同时获得较强的茎秆抗倒性及较高的小区产量。

参考文献 原文顺序
文献年度倒序
文中引用次数倒序
被引期刊影响因子

刘成, 冯中朝, 肖唐华, 马晓敏, 周广生, 黄凤洪, 李加纳, 王汉中. 我国油菜产业发展现状、潜力及对策
中国油料作物学报, 2019,41(4):485-489.
[本文引用: 1]

LIU C, FENG Z C, XIAO T H, MA X M, ZHOU G S, HUANG F H, LI J N, WANG H Z. Development, potential and adaptation of Chinese rapeseed industry
Chinese Journal of Oil Crop Sciences, 2019,41(4):485-489. (in Chinese)
[本文引用: 1]

蒯婕, 孙盈盈, 左青松, 廖庆喜, 冷锁虎, 程雨贵, 曹石, 吴江生, 周广生. 机械收获模式下直播冬油菜密度与行距的优化
作物学报, 2016,42(6):898-908.
[本文引用: 1]

KUAI J, SUN Y Y, ZUO Q S, LIAO Q X, LENG S H, CHENG Y G, CAO S, WU J S, ZHOU G S. Optimization of plant density and row spacing for mechanical harvest inwinter rapeseed (Brassica napus L.)
Acta Agronomica Sinica, 2016,42(6):898-908. (in Chinese)
[本文引用: 1]

蒯婕, 王积军, 左青松, 陈红琳, 高建芹, 汪波, 周广生, 傅廷栋. 长江流域直播油菜密植效应及其机理研究进展
中国农业科学, 2018,51(24):4625-4632.
[本文引用: 1]

KUAI J, WANG J J, ZUO Q S, CHEN H L, GAO J Q, WANG B, ZHOU G S, FU T D. Effects and mechanism of higher plant density on directly-sown rapeseed in the Yangtze River Basin of China
Scientia Agricultura Sinica, 2018,51(24):4625-4632. (in Chinese)
[本文引用: 1]

BERRY P M, SPINK J H, GAY A P, CRAIGON J. A comparison of root and stem lodging risks among winter wheat cultivars
Journal of Agricultural Science, 2003,141:191-202.
[本文引用: 1]

雷小龙, 刘利, 刘波, 黄光忠, 马荣朝, 任万军. 杂交籼稻F优498机械化种植的茎秆理化性状与抗倒伏性
中国水稻科学, 2014,28(6):612-620.
[本文引用: 1]

LEI X L, LIU L, LIU B, HUANG G Z, MA R C, REN W J. Physical and chemical characteristics and lodging resistance of culm of indica hybrid rice F you 498 under mechanical planting
Chinese Journal of Rice Science, 2014,28(6):612-620. (in Chinese)
[本文引用: 1]

吴莲蓉. 油菜茎秆生化成分和倒伏相关性研究
[D]. 武汉: 华中农业大学, 2015.
[本文引用: 1]

WU L R. Study on the stem biochemical components in rapeseed and their relationship with lodging characters
[D]. Wuhan: Huazhong Agricultural University, 2015. (in Chinese)
[本文引用: 1]

李华英. 种植密度和播期对冬小麦籽粒产量和抗倒性能的影响
[D]. 泰安: 山东农业大学, 2015.
[本文引用: 1]

LI H Y. Effects of plant density and sowing date on grain yieldand lodging resistance of winter wheat
[D]. Taian: Shandong Agricultural University, 2015. (in Chinese)
[本文引用: 1]

BHAGAT K P, SAIRAM R K, DESHMUKH P S, KUSHWAHA S R. Biochemical analysis of stem in lodging tolerant and susceptible wheat (Triticum aestivum L.) genotypes under normal and late sown conditions
Indian Journal of Plant Physiology, 2011,16:68-74.
[本文引用: 1]

TANAKA K, MURATA K, YAMAZAKI M, ONOSATO K, MIYAO A, HIROCHIKA H. Three distinct rice cellulose synthase catalytic subunit genes required for cellulose synthesis in the secondary wall
Plant Physiology, 2003,133:78-83.
[本文引用: 1]

LI X Y, ZUO Q S, CHANG H B, BAI G P, KUAI J, ZHOU G S. Higher density planting benefits mechanical harvesting of rapeseed in the Yangtze River basin of China
Field Crops Research, 2018,218:97-105.
DOI:10.1016/j.fcr.2018.01.013URL [本文引用: 2]

孙盈盈. 不同栽培措施对油菜产量及抗倒性的影响
[D]. 武汉: 华中农业大学, 2016.
[本文引用: 3]

SUN Y Y. Effect of different cultivation measures on rapeseed yieldand lodging resistance
[D]. Wuhan: Huazhong Agricultural University, 2016. (in Chinese)
[本文引用: 3]

李小勇, 周敏, 王涛, 张兰, 周广生, 蒯婕. 种植密度对油菜机械收获关键性状的影响
作物学报, 2018,44(2):278-287.
[本文引用: 2]

LI X Y, ZHOU M, WANG T, ZHANG L, ZHOU G S, KUAI J. Effects of planting density on the mechanical harvesting characteristics of semi-winter rapeseed
Acta Agronomica Sinica, 2018,44(2):278-287. (in Chinese)
[本文引用: 2]

CROOK M J, ENNOS A R. The effect of nitrogen and growth regulators on stem and root characteristics associated with lodging in two cultivars of winter wheat
Journal of Experimental Botany, 1995,46(289):931-938.
[本文引用: 1]

LI X J, LI S Y, LIN J X. Effect of GA3 spraying on lignin and auxin contents and the correlated enzyme activities in bayberry (Myrica rubra Bieb.) during flower-bud induction
Plant Science, 2003,164:549-556.
DOI:10.1016/S0168-9452(03)00004-9URL [本文引用: 1]

赵小红, 白羿雄, 王凯, 姚有华, 姚晓华, 吴昆仑. 种植密度对2个青稞品种抗倒伏及秸秆饲用特性的影响
作物学报, 2020,46(4):586-595.
[本文引用: 1]

ZHAO X H, BAI Y X, WANG K, YAO Y H, YAO X H, WU K L. Effects of planting density on lodging resistance and straw forage characteristics in two hulless barley varieties
Acta Agronomica Sinica, 2020,46(4):586-595. (in Chinese)
[本文引用: 1]

VANHOLME R, MORREEL K, RALPH J, BOERJAN W. Lignin engineering
Current Opinion in Plant Biology, 2008,11:278-285.
DOI:10.1016/j.pbi.2008.03.005URLPMID:18434238 [本文引用: 1]
Lignins are aromatic polymers that are present mainly in secondarily thickened plant cell walls. Several decades of research have elucidated the main biosynthetic routes toward the monolignols and demonstrated that lignin amounts can be engineered and that plants can cope with large shifts in p-hydroxyphenyl/guaiacyl/syringyl (H/G/S) lignin compositional ratios. It has also become clear that lignins incorporate many more units than the three monolignols described in biochemistry textbooks. Together with the theory that lignin polymerization is under chemical control, observations hint at opportunities to design lignin structure to the needs of agriculture. An increasing number of examples illustrates that lignin engineering can improve the processing efficiency of plant biomass for pulping, forage digestibility and biofuels. Systems approaches, in which the plant's response to engineering of a single gene in the pathway is studied at the organismal level, are beginning to shed light on the interaction of lignin biosynthesis with other metabolic pathways and processes.

汪灿, 阮仁武, 袁晓辉, 胡丹, 杨浩, 林婷婷, 何沛龙, 李燕, 易泽林. 荞麦茎秆解剖结构和木质素代谢及其与抗倒性的关系
作物学报, 2014,40(10):1846-1856.
[本文引用: 2]

WANG C, RUAN R W, YUAN X H, HU D, YANG H, LIN T T, HE P L, LI Y, YI Z L. Relationship of anatomical structure and lignin metabolism with lodging resistance of culm in buckwheat
Acta Agronomica Sinica, 2014,40(10):1846-1856. (in Chinese)
[本文引用: 2]

束红梅, 赵新华, 周治国, 郑密, 王友华. 不同棉花品种纤维比强度形成的温度敏感性差异机理研究
中国农业科学, 2009,42(7):2332-2341.
[本文引用: 1]

SHU H M, ZHAO X H, ZHOU Z G, ZHENG M, WANG Y H. Physiological mechanisms of variation in temperature-sensitivity of cotton fiber strength formation between two cotton cultivars
Scientia Agricultura Sinica, 2009,42(7):2332-2341. (in Chinese)
[本文引用: 1]

SONG X, LIU F L, ZHENG P Y, ZHANG X K, LU G Y, FU G P, CHENG Y. Correlation analysis between agronomic traits and yield of rapeseed (Brassica napusL.) for high-density planting
Scientia Agricultura Sinica, 2010,43(9):1800-1806.
[本文引用: 1]

OZER H. Sowing date and nitrogen rate effects on growth, yield and yield components of two summer rapeseed cultivars
European Journal of Agronomy, 2003,19(3):453-463.
[本文引用: 1]

MATINFAR M, MATINFAR M, MAHJOOR M, SHIRANI RAD A H, MAHMODI R. Effect of plant density on yield and yield seed components of rapeseed (Brassica napus) cultivars
Journal of Crop Ecophysiology, 2013,6(4):405-414.
[本文引用: 1]

LI H G, CHENG X, ZHANG L P, HU J H, ZHANG F G, CHEN B Y, XU K, GAO G Z, LI H, LI L X, HUANG Q, LI Z Y, YAN G X, WU X M. An integration of genome-wide association study and gene co-expression network analysis identifies candidate genes of stem lodging-related traits in Brassica napus
Frontiers in Plant Science, 2018,9:796.
DOI:10.3389/fpls.2018.00796URLPMID:29946333 [本文引用: 1]
Lodging is a persistent problem which severely reduce yield and impair seed quality in rapeseed (Brassica napus L.). Enhancing stem strength (SS) has proven to be an effective approach to decrease lodging risk. In the present study, four interrelated stem lodging-related traits, including stem breaking resistance (SBR), stem diameter (SD), SS, and lodging coefficient (LC), were investigated among 472 rapeseed accessions. A genome-wide association study (GWAS) using Brassica 60K SNP array for stem lodging-related traits identified 67 significantly associated quantitative trait loci (QTLs) and 71 candidate genes. In parallel, a gene co-expression network based on transcriptome sequencing was constructed. The module associated with cellulose biosynthesis was highlighted. By integrating GWAS and gene co-expression network analysis, some promising candidate genes, such as ESKIMO1 (ESK1, BnaC08g26920D), CELLULOSE SYNTHASE 6 (CESA6, BnaA09g06990D), and FRAGILE FIBER 8 (FRA8, BnaC04g39510D), were prioritized for further research. These findings revealed the genetic basis underlying stem lodging and provided worthwhile QTLs and genes information for genetic improvement of stem lodging resistance in B. napus.

KATAHIRA R, SLUITER J B, SCHELL D J, DAVIS M F. Degradation of carbohydrates during dilute sulfuric acid pretreatment can interfere with lignin measurements in solid residues
Journal of Agricultural and Food Chemistry, 2013,61(13):3286-3292.
URLPMID:23428141 [本文引用: 1]

PEI Y J, LI Y Y, ZHANG Y B, YU C B, FU T D, ZOU J, TU Y Y, PENG L C, CHEN P. G-lignin and hemicellulosic monosaccharides distinctively affect biomass digestibility in rapeseed
Bioresour Technol, 2016,203:325-333.
DOI:10.1016/j.biortech.2015.12.072URLPMID:26748046 [本文引用: 1]
In this study, total 19 straw samples from four Brassica species were determined with a diverse cell wall composition and varied biomass enzymatic digestibility under sulfuric acid or lime pretreatment. Correlation analysis was then performed to detect effects of cell wall compositions and wall polymer features (cellulose crystallinity, hemicellulosic monosaccharides and lignin monomers) on rapeseeds biomass digestibility. As a result, coniferyl alcohol (G-lignin) showed a strongly negative effect on biomass saccharification, whereas hemicellulosic monosaccharides (fucose, galactose, arabinose and rhamnose) were positive factors on lignocellulose digestions. Notably, chemical analyses of four typical pairs of samples indicated that hemicellulosic monosaccharides and G-lignin may coordinately influence biomass digestibility in rapeseeds. In addition, Brassica napus with lower lignin content exhibited more efficiency on both biomass enzymatic saccharification and ethanol production, compared with Brassica junjea. Hence, this study has at first time provided a genetic strategy on cell wall modification towards bioenergy rapeseed breeding.

周广生, 梅方竹, 周竹青, 朱旭彤. 小麦不同品种耐湿性生理指标综合评价及其预测
中国农业科学, 2003,36(11):1378-1382.
[本文引用: 1]

ZHOU G S, MEI F Z, ZHOU Z Q, ZHU X T. Comprehensive evaluation and forecast on physiological indices of waterlogging resistance of different wheat varieties
Scientia Agricultura Sinica, 2003,36(11):1378-1382. (in Chinese)
[本文引用: 1]

吕丽华, 陶洪斌, 夏来坤, 张雅杰, 赵明, 赵久然, 王璞. 不同种植密度下的夏玉米冠层结构及光合特性
作物学报, 2008,34(3):447-455.
[本文引用: 1]

Lü L H, TAO H B, XIA L K, ZHANG Y J, ZHAO M, ZHAO J R, WANG P. Canopy structure and photosynthesis traits of summer maize under different planting densities
Acta Agronomica Sinica, 2008,34(3):447-455. (in Chinese)
[本文引用: 1]

胡焕焕, 刘丽平, 李瑞奇, 李慧玲, 李雁鸣. 播种期和密度对冬小麦品种河农822产量形成的影响
麦类作物学报, 2008,28(3):490-495.
[本文引用: 1]

HU H H, LIU L P, LI R Q, LI H L, LI Y M. Effect of sowing date and planting density on the yield formation of a winter wheat cultivar Henong 822
Journal of Triticeae Crops, 2008,28(3):490-495. (in Chinese)
[本文引用: 1]

LIU W G, DENG Y C, HUSSAIN S, ZOU J L, YUAN J, LUO L, YANG C Y, YUAN X Q, YANG W Y. Relationship between cellulose accumulation and lodging resistance in the stem of relay intercropped soybean [Glycine max (L.) Merr.]
Field Crops Research, 2016,196:261-267.
[本文引用: 1]

崔海岩, 靳立斌, 李波, 张吉旺, 赵斌, 董树亭, 刘鹏. 遮阴对夏玉米茎秆形态结构和倒伏的影响
中国农业科学, 2012,45(17):3497-3505.
[本文引用: 1]

CUI H Y, JIN L B, LI B, ZHANG J W, ZHAO B, DONG S T, LIU P. Effects of shading on stalks morphology, structure and lodging of summer maize in field
Scientia Agricultura Sinica, 2012,45(17):3497-3505. (in Chinese)
[本文引用: 1]

佘恒志, 聂姣, 李英双, 刘星贝, 胡丹, 马珊, 次仁卓嘎, 汪灿, 吴东倩, 阮仁武, 易泽林. 不同抗倒伏能力甜荞品种茎秆木质素及其单体合成特征
中国农业科学, 2017,50(7):1202-1209.
[本文引用: 1]

SHE H Z, NIE J, LI Y S, LIU X B, HU D, MA S, CIREN Z G, WANG C, WU D Q, RUAN R W, YI Z L. Lignin and lignin monomer synthetic characteristics of culm incommon buckwheat with different lodging resistance capabilities
Scientia Agricultura Sinica, 2017,50(7):1202-1209. (in Chinese)
[本文引用: 1]

卢昆丽, 尹燕枰, 王振林, 李勇, 彭佃亮, 杨卫兵, 崔正勇, 杨东清, 江文文. 施氮期对小麦茎秆木质素合成的影响及其抗倒伏生理机制
作物学报, 2014,40(9):1686-1694.
[本文引用: 1]

LU K L, YIN Y P, WANG Z L, LI Y, PENG D L, YANG W B, CUI Z Y, YANG D Q, JIANG W W. Effect of nitrogen fertilization timing on lignin synthesis of stem and physiological mechanism of lodging resistance in wheat
Acta Agronomica Sinica, 2014,40(9):1686-1694. (in Chinese)
[本文引用: 1]

师恭曜. 甘蓝型油菜茎秆抗倒伏性构成因素的鉴定与评价
[D]. 郑州: 郑州大学, 2010: 27-31.
[本文引用: 1]

SHI G Y. Characterization and evaluation of stem lodging resistance in rapeseed (Brassica Campestris L.)
[D]. Zhengzhou: Zhengzhou University, 2010: 27-31. (in Chinese)
[本文引用: 1]

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