夏玉米籽粒胚乳细胞增殖及产量对不同光照的响应

引用本文

高佳, 史建国, 董树亭, 刘鹏, 赵斌, 张吉旺. . 夏玉米籽粒胚乳细胞增殖及产量对不同光照的响应. 作物学报, 2017, 43(10): 1548-1558
GAO Jia, SHI Jian-Guo, DONG Shu-Ting, LIU Peng, ZHAO Bin, ZHANG Ji-Wang. . Response of Endosperm Cell Proliferation and Grain Yield of Summer Maize to Different Light Conditions. ACTA AGRONOMICA SINICA, 2017, 43(10): 1548-1558 复制到剪切板

Permissions

作物学报编辑部

夏玉米籽粒胚乳细胞增殖及产量对不同光照的响应

高佳, 史建国, 董树亭, 刘鹏, 赵斌, 张吉旺^*

作物生物学国家重点实验室 / 山东农业大学农学院, 山东泰安 271018

^* 通讯作者(Corresponding author): 张吉旺, E-mail: jwzhang@sdau.edu.cn, Tel: 0538-8241485

第一作者联系方式: E-mail: gaoj0803@163.com, Tel: 15505385417

收稿日期:2016-12-06 接受日期:2017-05-10 网络出版日期:2017-05-22

基金:本研究由国家自然科学基金项目(31671629), 国家现代农业产业技术体系建设专项(CARS-02-20), 山东省农业重大应用技术创新项目和山东省玉米育种与栽培技术企业重点实验室开放课题资助

摘要

为了探讨不同光照条件对夏玉米籽粒胚乳细胞增殖及产量的影响, 2012—2014年于山东农业大学试验农场进行了大田试验, 选玉米品种郑单958和登海605为试验材料, 设置大田自然光照(CK)、开花至收获期遮阴(S1)、拔节至开花期遮阴(S2)、出苗至收获期遮阴(S3)和开花至收获期增光(L) 5个处理, 研究其对夏玉米籽粒胚乳细胞增殖、籽粒灌浆、淀粉含量、干物质积累及产量的影响。结果表明, 遮阴后籽粒胚乳细胞增殖速率、细胞数目和淀粉积累量降低, 细胞充实度下降, 粒重显著降低, 且S3对其影响最大, S1次之, S2影响相对较小。郑单958和登海605两个品种S1、S2和S3成熟期的胚乳细胞数目较CK分别降低33%、6%、29%和41%、5%、29%, DH605 L处理显著提高了胚乳细胞增殖速率、胚乳细胞数目及淀粉含量, 细胞充实度和粒重显著上升。遮阴导致籽粒灌浆速率减缓, 灌浆高峰期推迟, 2个品种S1、S2和S3的最大灌浆速率较CK分别降低34%、13%、58%和38%、13%、64%, 登海605 L处理提高了籽粒灌浆速率, 籽粒灌浆高峰期提前出现。2个品种S1、S2和S3较CK分别减产58%、26%、81%和67%、27%、81%, 登海605 L处理增产9%。即遮阴降低了籽粒胚乳细胞数目、淀粉含量和灌浆速率, 进而影响产量, 花粒期充足的光照则有助于籽粒胚乳细胞增殖和产量提高。

关键词: 夏玉米; 光照; 胚乳细胞增殖; 籽粒灌浆; 产量

Response of Endosperm Cell Proliferation and Grain Yield of Summer Maize to Different Light Conditions

GAO Jia, SHI Jian-Guo, DONG Shu-Ting, LIU Peng, ZHAO Bin, ZHANG Ji-Wang^*

State Key Laboratory of Crop Biology / Agronomy College of Shandong Agricultural University, Tai’an 271018, China

Fund:This study was supported by National Natural Science Funds (31671629), the China Agriculture Research System (CARS-02-20), Shandong Key Agricultural Technology Innovation Project and Shandong Provincial Key Laboratory of Corn Breeding and Cultivation Technology.

Abstract

During the period from 2012 to 2014, Zhengdan 958 (ZD958) and Denghai 605 (DH605) were used with five treatments in a randomized complete block design with three replicates, including increasing light (L) from tassel stage (VT) to maturity stage (R6) and three shading treatments (40% of ambient light intensity) from VT to R6 (S1), from the sixth leaf stage (V6) to VT (S2), and from emergence stage (VE) to R6 (S3), and natural lighting in the field (CK). The illumination intensity of L on cloudy day was 1600-1800 μmol m^-2 s^-1. This experiment was conducted to explore the effects of light on endosperm cell proliferation, grain filling, starch content, dry matter accumulation and yield of summer maize. The dry matter accumulation of embryo and endosperm was significantly decreased under shading, as well as the content of starch and the endosperm cell plumpness decreased also. Effect of S3 on endosperm cell proliferation was the most, while that of S2 was the smallest. At R6, the endosperm cells number of S3, S2, S1 decreased by 33%, 6%, 29% in ZD958 and 41%, 5%, 29% in DH605. The dry matter accumulation of embryo and endosperm as well as the content of starch and the endosperm cell plumpness were increased after increasing light. The grain filling rate decreased and the days to maximum grain filling rate postponed under shading, while those increased after increasing light. The maximum grain filling rate of S1, S2, S3 decreased by 34%, 13%, 58% in ZD958 and 38%, 13%, 64% in DH605. The grain yield of S1, S2, S3 decreased by 58%, 26%, 81% in ZD958 and 67%, 27%, 81% in DH605. Results indicate that the number of endosperm cells, starch content and grain filling rate are decreased under shading, resulting in grain yield reducing, while L treatment can contribute to increasing the number of endosperm cell and grain yield.

Keyword: Summer maize; Illumination; Endosperm cell proliferation; Grain filling; Grain yield

Show Figures

玉米胚乳质量占籽粒质量的80%左右, 胚乳细胞的数量和充实度对粒重有决定作用^{[1, 2]}, 同时也代表着“ 库” 的高产潜力。胚乳细胞增殖速率快, 数量多, 有助于光合同化产物在籽粒中积累和淀粉粒形成, 提高籽粒充实度, 进而获得高产^{[3, 4, 5]}。玉米籽粒积累的淀粉数量决定其产量和品质^{[6, 7]}。淀粉作为光合作用的主要产物, 通过酶催化转运至胚乳细胞贮藏, 其积累速率和持续活跃时间决定粒重^{[8, 9]}, 即胚乳细胞越多, 淀粉积累量越多, 充实度越好, 粒重越高^[10]。

遮阴导致玉米籽粒胚乳细胞数和充实度降低^[10]。贾士芳等^[1]研究表明, 遮阴显著降低玉米籽粒中胚乳细胞数、淀粉及籽粒充实度, 最终降低粒重。杜成凤等^[11]和史建国等^[12]研究认为, 充足的光照有助于促进玉米光合作用, 也是玉米高产的基础; 曾希柏等^[13]研究指出, 在一定范围内植物生物量随光强的增加而提高, 玉米产量也会在一定范围随光照强度的上升而增加。

光是玉米光合作用的能量来源, 光照变化对植株光合作用、同化产物形成有极为显著的影响^[14]。开花后的光照时数、光照强度和叶面积决定了80%的玉米产量^[15]。遮阴导致夏玉米生育时期推迟, 株高、茎粗、叶面积显著减少^{[16, 17, 18, 19, 20]}, 倒伏率升高^[21], 叶片光合特性和生殖器官发育受到影响^{[22, 23, 24]}, 籽粒败育^[25], 产量下降^{[18, 19]}。1951年以来, 气候变化加剧, 日照时数减少, 太阳辐射减弱, 年平均气温不断升高, 极端天气频繁发生^{[26, 27]}, 以黄淮海区为例, 在夏玉米生长期内(6月至10月), 尤其是花粒期, 阴雨寡照天气频繁发生, 导致玉米总产量减少3%~ 6%^[15]。因此, 研究弱光胁迫对夏玉米胚乳细胞特性的影响具有重要意义。前人关于光照对夏玉米生产影响的研究多集中于营养体建成, 而关于不同时期光照对夏玉米籽粒生长发育影响的研究鲜见报道。本文通过设计不同时期遮阴和花粒期增光处理, 探讨不同光照对夏玉米胚乳细胞增殖及籽粒产量形成的影响, 为应对光照不足气候变化条件下的夏玉米高产高效生产提供理论和技术支持。

1 材料与方法

1.1 试验地点

山东农业大学试验农场(36.09° N, 117.09° E)具温带大陆性季风气候, 夏玉米从开花到成熟期的平均积温为1636℃ d。土壤类型为棕壤土, 播种前0~20 cm土壤养分状况为有机质9.3 g kg^-1, 全氮0.8 g kg^-1, 全磷0.9 g kg^-1, 速效磷37.2 mg kg^-1, 速效钾84.2 mg kg^-1, 碱解氮80.6 mg kg^-1。

1.2 试验设计

2012— 2014年, 选用郑单958和登海605, 于6月15日播种, 种植密度67 500株 hm^-2。设计5个处理, 即对照CK: 大田自然光照; S1: 开花至收获期遮阴; S2: 拔节至开花期遮阴; S3: 出苗至收获期遮阴; L: 开花至收获期增光(L处理仅选用登海605品种)。遮光度为60%, 增光强度为1600~1800 μ mol m^-2 s^-1。利用脚手架和遮光率为60%的黑色遮阴网搭建可升降式遮阴棚, 遮阴网与玉米冠层始终保持2 m距离, 从而保证遮阴棚内小气候与大田自然光照基本一致(表1)。利用金卤灯在阴天时增光, 光照时长及强度与晴朗天气相当, 正常光照下夏玉米花粒期的日照时数平均为359.1 h, 而增光处理在该基础上大约增加光照时数206 h。小区面积为3 m× 9 m, 重复3次, 随机排列。按12 000 kg hm^-2的产量水平施用纯N 210 kg hm^-2、P₂O₅75 kg hm^-2和K₂O 150 kg hm^-2, 氮肥为尿素(含N 46%), 磷肥为过磷酸钙(含P₂O₅ 17%), 钾肥为氯化钾(含K₂O 60%)。氮肥分拔节期40%和大喇叭口期60%两次施入, 磷肥和钾肥在拔节期一次性施入, 按高产田水平进行田间管理。

1.3 测定项目与方法

1.3.1 田间小气候参照崔海岩等^[28]方法测定光照强度、气温、CO₂浓度、相对湿度、风速和地温(5 cm)。开花后连续测定7 d, 均在11:00开始测定, 计算平均值(表1)。

表1 不同光照条件对田间小气候的影响 Table 1 Effects of different light conditions on the microclimate in the experimental field

1.3.2 胚乳细胞选择长势一致的植株, 于花后5、10、15、20、25、30、40、50 d各取每处理3个果穗, 剥取穗中部正常籽粒10粒, FAA固定液固定后保存于冰箱中。参照张祖建等^[29]方法测定玉米籽粒胚乳细胞数, 用Logistic方程拟合籽粒胚乳细胞增殖动态, 导出增殖特征参数。

1.3.3 籽粒灌浆特性于吐丝期选择生长一致的植株标记, 授粉后每隔10 d取各处理3个果穗, 取中部籽粒100粒, 直至成熟, 测定籽粒体积和干重, 3次重复。用Logistic方程拟合籽粒灌浆增殖动态, 导出灌浆特征参数。

1.3.4 籽粒淀粉和粗脂肪含量采用双波长法测定淀粉含量及组分^[30]; 采用残余油重法测定粗脂肪含量^[30]。

1.3.5 干物质积累与分配于成熟期取样, 每个处理各5株, 每株分为茎、叶、穗轴和籽粒四部分, 烘干并称重, 用于计算干物质的积累和分配。

1.3.6 叶面积指数(LAI)与光合势(LAD) 从每个小区连续选择10株长势一致的植株, 分别在各生育时期测定每片叶的长度和最大宽度。

单叶叶面积(cm²)=叶长(cm)× 叶宽(cm)× 0.75

LAI=(单株叶面积× 每个小区的植株数)/小区面积

LAD(m²d m^-2)=(某阶段起始叶面积+该阶段结束叶面积)/2 × 间隔天数

1.3.7 叶绿体色素含量(SPAD) 于各生育时期使用SPAD-502便携式叶绿素仪(Soil-plant Analysis Development Section, Minolta Camera Co., Osaka, Japan)在每小区连续选取10株生长一致的植株测定SPAD值(开花前测定最新展开叶, VT及其以后各时期测穗位叶, 测定时避开叶脉), 取平均值。

1.3.8 测产与考种于每小区中间3行, 随机选取30株玉米的果穗, 自然风干用于室内考种、测产。产量(kg hm^-2) = 收获穗数(ears hm^-2)× 穗粒数× 千粒重(g)× 10^-6 × (1-含水量%)/(1-14%), 收获穗数为田间调查有效穗数。

1.4 数据处理

采用Microsoft Excel 2007软件处理数据, SPSS 16.0 (LSD, Duncan)软件统计分析, SigmaPlot 10.0作图。

表2 不同光照条件对夏玉米产量及其构成因素的影响 Table 2 Effect of different light conditions on yield and yield components of summer maize

年份 Year	品种 Hybrid	处理 Treatment	产量 Yield (kg hm^-2)	千粒重1000-kernel weight (g)	穗粒数 No. of kernels per ear	公顷穗数 Harvest ear number (ears hm^-2)	收获指数Harvest index (%)	空秆率Barrenness (%)
2012	郑单958 Zhengdan 958	S1	3518 c	299 b	296 c	39684 c	43 c	40 a
		S2	6878 b	330 b	407 b	51233 b	50 b	8 b
		S3	1313 d	205 c	188 d	34129 d	38 d	46 a
		CK	10070 a	353 a	502 a	56919 a	56 a	8 b
	登海605 Denghai 605	S1	2336 d	231 c	266 c	38096 c	39 d	42 a
		S2	7649 c	357 b	344 b	59530 b	45 c	11 b
		S3	1168 e	226 c	150 d	35981 d	33 e	38 a
		CK	11322 b	384 a	478 a	61679 b	51 b	6 bc
		L	12317 a	389 a	515 a	64934 a	57 a	1 c
2013	郑单958 Zhengdan 958	S1	5539 c	279 c	368 c	54077 c	37 d	16 a
		S2	8101 b	310 b	459 b	57040 b	50 b	9 b
		S3	2324 d	278 c	171 d	49633 d	44 c	19 a
		CK	11541 a	351 a	530 a	62117 a	58 a	4 c
	登海605 Denghai 605	S1	4485 d	315 b	253 c	56300 c	40 d	16 a
		S2	9754 c	347 a	478 b	58892 b	46 c	8 b
		S3	2809 e	229 b	156 d	54829 c	33 e	16 a
		CK	12554 b	362 a	545 a	63593 a	51 b	4 c
		L	12923 a	369 a	527 a	66485 a	55 a	1 d
2014	郑单958 Zhengdan 958	S1	4797 c	250 c	355 c	54077 c	45 c	14 b
		S2	8094 b	277 b	478 b	61114 b	50 b	7 c
		S3	1518 d	203 d	165 d	45187 d	33 d	19 a
		CK	11819 a	334 a	555 a	63670 a	53 a	5 c
	登海605 Denghai 605	S1	4361 d	290 d	282 d	53336 d	34 d	12 ab
		S2	9064 c	304 c	493 c	60373 c	41 c	8 b
		S3	1527 e	216 e	162 e	43706 e	27 e	17 a
		CK	10886 b	320 b	520 b	65337 b	57 b	4 c
		L	13605 a	363 a	578 a	70004 a	59 a	1 d
ANOVA
年份Year (Y)			* *	NS	NS	*	NS	* *
品种Hybrid (H)			* *	* *	NS	NS	NS	NS
处理Treatment (T)			* *	* *	* *	* *	* *	* *
Y× H			NS	NS	*	NS	NS	NS
Y× T			NS	*	NS	NS	NS	NS
H× T			NS	NS	NS	NS	NS	NS
Y× H× T			NS	NS	NS	NS	NS	NS

Values followed by different letters in the same column are significantly different at the 0.05 probability level. NS: not signiﬁ cant. * : signiﬁ cant at the 0.05 probability level; * * : signiﬁ cant at the 0.01 probability level. S1: shading from tassel stage (VT) to maturity stage (R6; S2: shading from six-leaf stage (V6) to VT; S3: shading from emergence stage to R6; CK: natural lighting in the field; L: increasing light from VT to R6.

同列标以不同小写字母的值差异达5%显著水平。NS: 差异不显著; * 表示在0.05水平下差异显著; * * 表示在0.01水平下差异显著。S1: 开花至收获期遮阴; S2: 拔节至开花期遮阴; S3: 出苗至收获期遮阴; CK: 大田自然光照; L: 开花至收获期增光

表2 不同光照条件对夏玉米产量及其构成因素的影响 Table 2 Effect of different light conditions on yield and yield components of summer maize

2 结果与分析

2.1 产量及其构成因素

不同光照条件对玉米产量影响显著。郑单958品种S1、S2和S3较CK分别减产58%、26%和81%; 登海605品种S1、S2和S3较CK分别减产67%、27%和81%, L较CK增产9%。产量构成因素也发生了改变, 遮阴导致千粒重、穗粒数和公顷穗数降低, 空秆率增加, 增光后千粒重和穗粒数增加, 公顷穗数无显著变化。

表3 不同光照条件对夏玉米成熟期干物质积累与分配的影响 Table 3 Effects of different light conditions on the dry matter accumulation and distribution at maturity of summer maize

年份 Year	品种 Hybrid	处理 Treatment	茎秆 Stalk		叶片 Leaf		籽粒 Grain		总重 Total dry matter weight (g plant^-1)
年份 Year	品种 Hybrid	处理 Treatment	干重 Dry matter weight (g plant^-1)	比例 Proportion (%)	干重 Dry matter weight (g plant^-1)	比例 Proportion (%)	干重 Dry matter weight (g plant^-1)	比例 Proportion (%)	总重 Total dry matter weight (g plant^-1)
2012	郑单958 Zhengdan 958	S1	43.2 c	30.0	27.6 c	19.2	67.5 c	46.8	144.0 c
		S2	58.0 b	24.6	35.8 b	15.1	123.0 b	52.1	236.1 b
		S3	30.3 d	43.6	18.1 d	26.1	15.8 d	22.8	69.4 d
		CK	82.1 a	23.9	39.3 a	11.4	200.9 a	58.5	343.7 a
	登海605 Denghai 605	S1	41.0 c	40.6	27.4 c	27.2	25.7 d	25.4	100.9 c
		S2	65.2 b	27.9	37.3 a	16.0	107.6 c	46.0	233.8 b
		S3	35.9 d	43.4	25.6 c	31.0	14.5 d	17.6	82.7 d
		CK	76.2 a	26.4	32.7 b	11.3	156.6 b	54.2	288.8 a
		L	76.0 a	24.3	30.4 b	9.7	177.4 a	56.7	312.9 a
2013	郑单958 Zhengdan 958	S1	48.7 c	28.7	35.7 ab	21.1	75.0 c	44.2	169.5 c
		S2	60.0 b	24.6	39.2 a	16.1	121.6 b	49.8	244.0 b
		S3	43.5 d	32.8	31.9 b	24.0	49.3 d	37.1	132.7 d
		CK	80.0 a	23.4	39.8 a	11.6	198.8 a	58.1	342.0 a
	登海605 Denghai 605	S1	51.4 c	37.3	32.3 c	23.5	45.1 d	32.7	137.8 c
		S2	79.0 b	27.7	48.4 a	17.0	130.4 c	45.7	285.3 b
		S3	40.1 d	34.7	25.3 d	21.9	36.5 d	32.7	115.5 d
		CK	88.9 a	29.5	39.3 b	13.1	152.4 b	50.6	301.1 b
		L	88.7 a	26.6	38.1 b	11.4	183.6 a	55.1	332.9 a
2014	郑单958 Zhengdan 958	S1	43.9 b	31.0	29.5 b	20.8	60.8 c	42.9	141.8 c
		S2	78.1 a	27.5	47.2 a	16.6	140.9 b	49.7	283.6 b
		S3	34.4 c	32.6	24.0 c	22.7	37.3 d	35.3	105.6 d
		CK	80.8 a	25.2	45.8 a	14.3	171.5 a	53.4	321.0 a
	登海605 Denghai 605	S1	53.3 c	34.2	32.4 b	20.8	51.3 d	32.9	156.0 c
		S2	65.8 b	31.3	36.7 a	17.5	86.7 c	41.3	209.9 b
		S3	34.3 d	39.5	21.8 c	25.1	24.6 e	28.4	86.8 d
		CK	78.7 a	26.4	37.6 a	12.6	168.6 b	56.6	297.7 a
		L	79.4 a	24.7	38.5 a	12.0	189.8 a	59.0	321.5 a

Values followed by a different small letter within a column are significantly different at the 0.05 probability level. S1: shading from tassel stage (VT) to maturity stage (R6); S2: shading from six-leaf stage (V6) to VT; S3: shading from emergence stage to R6; CK: natural lighting in the field; L: increasing light from VT to R6.

同列不同小写字母表示达到0.05显著差异。S1: 开花至收获期遮阴; S2: 拔节至开花期遮阴; S3: 出苗至收获期遮阴; CK: 大田自然光照; L: 开花至收获期增光。

2.2 干物质积累与分配

遮阴后玉米成熟期干物质积累量显著下降, 郑单958品种S1、S2和S3总干物质积累量较CK分别降低55%、24%和69%; 登海605品种S1、S2和S3较CK分别降低52%、18%和65%, L总干物质积累量较CK增加9%。由各个器官的干物质积累量可知, 遮阴条件下籽粒中的积累量大幅度下降, 且均小于CK。郑单958品种S1、S2和S3籽粒干重比例分别为45%、50%和33%; 登海605品种S1、S2、S3和L所占比例分别为34%、45%、27%和57%。

表4 不同光照条件对夏玉米籽粒胚乳细胞增殖特征参数的影响 Table 4 Effects of different light conditions on the characteristic parameters in endosperm cell proliferation of summer maize

2.3 胚乳细胞增殖特性

由图1可以看出, 各处理籽粒胚乳细胞增殖均呈“ S” 型曲线。各时期的遮阴处理胚乳细胞数目始终低于CK, 登海605 L处理籽粒胚乳细胞数目高于CK。各方程的决定系数(R²)均在0.99以上, 配合度高(表4)。遮阴导致ET_max升高, EW_max和EG_max均降低。郑单958品种S1、S2和S3的最大增殖速率较CK分别降低30%、7%和54%; 登海605品种S1、S2和S3较CK分别降低32%、23%和52%, L较CK提高15%。

Figure Option
View Download New Window

图1 不同光照条件对夏玉米籽粒胚乳细胞增殖的影响(标准差) ZD958: 郑单958; DH605: 登海605。S1: 开花至收获期遮阴; S2: 拔节至开花期遮阴; S3: 出苗至收获期遮阴; CK: 大田自然光照; L: 开花至收获期增光。Fig. 1 Effects of different light conditions on endosperm cell proliferation of summer maize (standard deviation) S1: shading from tassel stage (VT) to maturity stage (R6); S2: shading from six-leaf stage (V6) to VT; S3: shading from emergence stage to R6; CK: natural lighting in the field; L: increasing light from VT to R6.

2.4 胚和胚乳的干重及其比例

从表5可知, 遮阴显著降低了籽粒胚和胚乳的干重, 登海605 L处理干重显著增加, 且籽粒胚和胚乳干重表现为L> CK> S2> S1> S3。遮阴后胚/胚乳比值增高, 说明遮阴对胚乳干重的影响程度大于胚。

表5 不同光照条件对夏玉米籽粒胚、胚乳及胚/胚乳的影响 Table 5 Effects of different light conditions on embryo, endosperm and embryo/endosperm ratio of summer maize

2.5 籽粒灌浆特性

遮阴延缓了最大籽粒灌浆速率的出现, 其中S3影响最大, S1次之, S2最小。遮阴导致W_max和G_max下降, 登海605品种S3、S2和S1处理的W_max和G_max较CK分别下降50%、6%、8%和44%、5%、11%, 即遮阴降低了玉米籽粒最大灌浆速率及其生长量, 郑单958与登海605品种的变化趋势一致。登海605 L处理的最大灌浆速率提前出现, 提高了籽粒最大灌浆速率及其生长量。

表6 不同光照条件对夏玉米籽粒灌浆特征参数的影响 Table 6 Effects of different light conditions on the characteristic parameters of grain-filling of summer maize

2.6 籽粒直链淀粉、支链淀粉和粗脂肪含量

从表7可知, 遮阴降低了玉米籽粒总淀粉含量, 且直链淀粉降幅更大。郑单958和登海605两品种S1、S2和S3处理的总淀粉含量较CK分别降低11%、5%、16%和10%、5%、12%, 登海605 L处理籽粒淀粉含量较CK显著增加5.9%。遮阴增加了籽粒中粗脂肪含量, 且S3增幅最大。

表7 不同光照条件对夏玉米籽粒淀粉含量和粗脂肪含量的影响 Table 7 Effects of different light conditions on starch and gross fat content of summer maize

2.7 籽粒产量与各生理指标的相关性分析

由表8可知, W_max、G_max等籽粒灌浆参数与籽粒产量呈极显著正相关, 即W_max、G_max等越大, 籽粒产量越高, 而T_max与产量呈显著负相关。EW_max、EG_max等籽粒胚乳增殖参数与籽粒产量也呈极显著正相关, ET_max与产量呈显著负相关, 籽粒胚乳干重与产量呈显著正相关。由表9可知, LAI和LAD与籽粒产量呈显著正相关, SPAD值与籽粒产量呈极显著正相关, 即提高LAI、LAD和SPAD值有助于增加产量。

表8 籽粒产量、籽粒灌浆特性和胚乳细胞增殖特性的相关系数 Table 8 Correlation coefficients between grain yield and characters of grain filling and endosperm cell proliferation

表9 籽粒产量和LAI、LAD、SPAD值的相关性系数 Table 9 Correlation coefficients of grain yield, LAI, LAD, and SPAD

3 讨论

3.1 不同光照对夏玉米产量和干物质积累量的影响

光合作用决定了玉米90%以上的干物质, 且干物质的积累、分配与产量显著正相关^{[31, 32, 33]}, 尤其是生育后期的干物质积累分配特性决定了夏玉米的最终产量^[34]。前人研究表明, 光照不足, 尤其是生育后期光照不足严重制约玉米生产, 表现为叶片光合性能下降, 籽粒灌浆强度降低, 干物质向籽粒中的转运量下降, 直接影响夏玉米产量^{[35, 36]}。Cui等^[31]研究指出, 雌穗小花分化对光照非常敏感, 光照不足降低小花分化总数, 降低穗粒数, 导致产量降低。本试验结果进一步表明, 花前遮阴降低了光合产物积累量, 花后遮阴降低了后期干物质积累量和籽粒灌浆速率, 导致产量降低, 即花前遮阴削弱了“ 源” 的生产能力, 花后遮阴影响了“ 库” 的充实。各处理玉米产量和干物质积累量降低幅度为S3> S1> S2, 说明夏玉米花粒期对光照更为敏感, 花粒期充足的光照更有利于夏玉米高产^{[37, 38, 39]}。

3.2 不同光照对夏玉米胚乳细胞增殖的影响

胚乳细胞数目代表了籽粒“ 库” 的潜力, 也代表该潜力的实现程度, 而单个胚乳细胞的充实状态也对产量的提高具有重要作用^[7]。胚乳细胞越多, 积累的淀粉粒越多, 充实度越高, 粒重越高^[2]。本研究表明, 在授粉后30 d左右, 籽粒胚乳细胞数基本达到最高值(图1), 胚乳细胞停止或减少分裂, 胚乳细胞构建基本完成, 以淀粉积累为主, 籽粒生长进入淀粉粒的充实阶段, 淀粉含量和粒重快速增加。遮阴后玉米籽粒胚乳细胞增殖的最大速率下降, 导致胚乳细胞数目降低(降低幅度为S3> S1> S2), 最终表现为籽粒发育进程延缓, 淀粉积累量下降, 籽粒库容量变小, 充实度下降, 粒重降低, 相反, 增光条件下胚乳细胞增殖速度快、数目增多(表8), 籽粒积累同化产物的速度加快, 容纳较多的淀粉粒^[8], 提高产量。

3.3 不同光照对夏玉米籽粒灌浆的影响

胚乳细胞形成后, 籽粒生长进入有效灌浆期, 即籽粒淀粉粒的充实扩大期, 完成淀粉的积累和单粒质量的增加。胚乳细胞数目代表着“ 库” 的潜力, 增库则需要提高灌浆速率, 提前最大灌浆速率出现的日期, 保证光合产物高效地运转到籽粒^[40]。本研究表明, 遮阴降低了籽粒灌浆速率, 推迟了灌浆速率最大值的出现日期, 导致灌浆活跃期缩短, 严重影响籽粒的正常灌浆, 相反, 增光则有利于提高籽粒的灌浆效率。

增光处理一方面有利于玉米积累更多的光合产物, 提高干物质量^[39], 达到增“ 源” 的效果, 茎、叶等临时性贮藏器官向籽粒输送的光合产物增加, 籽粒灌浆速率和生长量增加, 籽粒干重所占比例升高; 另一方面增殖速率和最大生长量显著升高, 胚乳细胞能够较快地达到最大增殖速率, 胚乳细胞数目增多, 淀粉含量显著提高, 即扩大了籽粒的“ 库” 容量, 胚乳细胞有较高的充实度, 籽粒灌浆速率提高, 粒重增加。相反, 光照不足导致玉米光合速率下降, 光合产物向籽粒中的转运量降低^[37]。遮阴下胚乳细胞发育异常, 表现为增殖速率降低、最大增殖速率延迟出现以及最大生长量降低, 籽粒中的胚乳细胞数目严重下降, 淀粉含量、籽粒充实度、籽粒灌浆速率降低, 且开花后遮阴处理影响最大, 其粒重和产量降低幅度分别达20%和60%左右。

4 结论

花粒期光照显著影响夏玉米籽粒的生长发育。遮阴降低玉米籽粒胚乳细胞数目、淀粉含量和细胞充实度, 导致减产, 增光促进粒重增加, 提高产量。因此, 在实际生产中通过适当调整播期, 避开阴雨寡照天气, 合理布局种植密度, 改善田间光照条件有助于提高胚乳细胞的增殖速率及灌浆速率, 保证光合产物高效向籽粒运转, 实现高产。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

参考文献

View Option

[1]	贾士芳, 李从锋, 董树亭, 张吉旺. 花后不同时期遮光对玉米粒重及品质影响的细胞学研究. 中国农业科学, 2010, 43: 911-921 Jia S F, Li C F, Dong S T, Zhang J W. Effects of shading at different stages after anthesis on maize grain weight and quality at cytology level. Sci Agric Sin, 2010, 43: 911-921 (in Chinese with English abstract) [本文引用:2]
[2]	秦大鹏, 刘鹏, 王空军, 董树亭, 张吉旺, 刘月娥. 高淀粉玉米籽粒胚乳细胞增殖与淀粉积累的关系. 山东农业科学, 2008, (8): 35-39 Qin D P, Liu P, Wang K J, Dong S T, Zhang J W, Liu Y E. Relationship between endosperm cell proliferation and starch accumulation in high-starch maize grain. Shand ong J Agric Sin, 2008, (8): 35-39 (in Chinese with English abstract) [本文引用:2]
[3]	徐云姬, 顾道健, 张博博, 张耗, 王志琴, 杨建昌. 玉米果穗不同部位籽粒激素含量及其与胚乳发育和籽粒灌浆的关系. 作物学报, 2013, 39: 1452-1461 Xu Y J, Gu D J, Zhang B B, Zhang H, Wang Z Q, Yang J C. Hormone contents in kernels at different positions on an ear and their relationship with endosperm development and kernel filling in maize. Acta Agron Sin, 2013, 39: 1452-1461 (in Chinese with English abstract) [本文引用:1]
[4]	孙庆泉, 吴元奇, 胡昌浩, 董树亭, 荣廷昭, 张颖. 不同产量潜力玉米籽粒胚乳细胞增殖与籽粒充实期的生理活性. 作物学报, 2005, 31: 612-618 Sun Q Q, Wu Y Q, Hu C H, Dong S T, Rong T Z, Zhang Y. Physiological activities and multiplication of endosperm cell at filling stage of kernels with different yield in maize. Acta Agron Sin, 2005, 31: 612-618 (in Chinese with English abstract) [本文引用:1]
[5]	Zhao L, Pan T, Cai C, Wang J, Wei C. Application of whole sections of mature cereal seeds to visualize the morphology of endosperm cell and starch and the distribution of storage protein. J Cereal Sci, 2016, 71: 19-27 [本文引用:1]
[6]	张智猛, 戴良香, 胡昌浩, 董树亭, 王空军, 宁堂原. 玉米灌浆期水分差异供应对籽粒淀粉积累及其酶活性的影响. 植物生态学报, 2005, 29: 636-643 Zhang Z M, Dai L X, Hu C H, Dong S T, Wang K J, Ning T Y. Effects of different water treatments on starch accumulation and related enzyme activity in grain filling period. Chin J Plant Ecol, 2005, 29: 636-643 (in Chinese with English abstract) [本文引用:1]
[7]	谷岩, 王振民, 何文安, 梁烜赫, 吴春胜. 不同类型玉米品种籽粒胚乳细胞增殖与籽粒建成的关系. 西北农林科技大学学报, 2010, 38(1): 90-96 Gu Y, Wang Z M, He W A, Liang X H, Wu C S. Relationship between endosperm cell proliferation and seed formation in maize of different types. J Northwest A&F Univ, 2010, 38(1): 90-96 (in Chinese with English abstract) [本文引用:2]
[8]	李栋梁, 李小刚, 顾蕴洁, 王忠. 不同类型水稻品种胚乳发育的研究. 中国农业科学, 2014, 47: 3757-3768 Li D L, Li X G, Gu Y J, Wang Z. Investigation of endosperm cell development of different rice varieties. Sci Agric Sin, 2014, 47: 3757-3768 (in Chinese with English abstract) [本文引用:2]
[9]	王贺正, 马均, 李旭毅, 张荣萍. 水分胁迫对水稻籽粒灌浆及淀粉合成有关酶活性的影响. 中国农业科学, 2009, 42: 1550-1558 Wang H Z, Ma J, Li X Y, Zhang R P. Effects of water stress on grain filling and activities of enzymes involved in starch synthesis in rice. Sci Agric Sin, 2009, 42: 1550-1558 (in Chinese with English abstract) [本文引用:1]
[10]	李伯航, 崔彦宏. 夏玉米胚乳细胞建成与粒重关系. 河北农业大学学报, 1989, 12(4): 39-45 Li B H, Cui Y H. Studies on the relationship between endosperm cell development and kernel weight of summer maize. J Agric Univ Hebei, 1989, 12(4): 39-45 (in Chinese with English abstract) [本文引用:2]
[11]	杜成凤, 李潮海, 刘天学, 赵亚丽. 遮阴对两个基因型玉米叶片解剖结构及光合特性的影响. 生态学报, 2011, 31: 6633-6640 Du C F, Li C H, Liu T X, Zhao Y L. Response of anatomical structure and photosynthetic characteristics to low light stress in leaves of different maize genotypes. Acta Ecol Sin, 2011, 31: 6633-6640 (in Chinese with English abstract) [本文引用:1]
[12]	史建国, 朱昆仑, 曹慧英, 董树亭, 刘鹏, 赵斌, 张吉旺. 花粒期光照对夏玉米干物质积累和养分吸收的影响. 应用生态学报, 2015, 26: 46-52 Shi J G, Zhu K L, Cui H Y, Zhao B, Dong S T, Liu P, Zhang J W. Effect of light from flowering to maturity stage on dry matter accumulation and nutrient absorption of summer maize. Chin J Appl Ecol, 2015, 26: 46-52 (in Chinese with English abstract) [本文引用:1]
[13]	曾希柏. 作物生长中光照和氮肥施用量的相互关系研究. 土壤学报, 2008, 37: 380-386 Zeng X B. Research on the correlation of light and nitrogen application in crop growth. Acta Pedol Sin, 2008, 37: 380-386 (in Chinese with English abstract) [本文引用:1]
[14]	Ren B Z, Cui H Y, James J C, Dong S T, Liu P, Zhang J W. Effects of shading on the photosynthetic characteristics and mesophyll cell ultrastructure of summer maize. Sci Nat, 2016, 103: 67 [本文引用:1]
[15]	崔海岩, 靳立斌, 李波, 赵斌, 董树亭, 刘鹏, 张吉旺. 大田遮阴对夏玉米光合特性和叶黄素循环的影响. 作物学报, 2013, 39: 478-485 Cui H Y, Jin L B, Li B, Zhao B, Dong S T, Liu P, Zhang J W. Effects of shading on photosynthetic characteristics and xanthophyll cycle of summer maize in the field. Acta Agron Sin, 2013, 39: 478-485 (in Chinese with English abstract) [本文引用:2]
[16]	张吉旺, 董树亭, 王空军, 胡昌浩, 刘鹏. 遮阴对夏玉米生长发育和产量的影响. 应用生态学报, 2006, 17: 657-662 Zhang J W, Dong S T, Wang K J, Hu C H, Liu P. Effects of shading on the growth, development and grain yield of summer maize. Chin J Appl Ecol, 2006, 17: 657-662 (in Chinese with English abstract) [本文引用:1]
[17]	王洋, 齐晓宁, 邵金锋, 刘兆永, 李泽兴. 光照强度对不同玉米品种生长发育和产量构成的影响. 吉林农业大学学报, 2008, 30: 769-773 Wang Y, Qi X N, Shao J F, Liu Z Y, Li Z X. Effects of light intensity at full growing stage on the growth and yield of different maize varieties. J Jilin Agric Univ, 2008, 30: 769-773 (in Chinese with English abstract) [本文引用:1]
[18]	付景, 李潮海, 赵久然, 马丽, 刘天学. 玉米品种耐阴性指标的筛选与评价. 应用生态学报, 2009, 20: 2705-2709 Fu J, Li C H, Zhao J R, Ma L, Liu T X. Shade-tolerance indices of maize: Selection and evaluation. Chin J Appl Ecol, 2009, 20: 2705-2709 (in Chinese with English abstract) [本文引用:2]
[19]	王秀萍, 刘天学, 李潮海, 李大鹏. 遮光对不同株型玉米品种农艺性状和果穗发育的影响. 江西农业学报, 2010, 22(1): 5-7 Wang X P, Liu T X, Li C H, Li D P. Effects of shading on agronomic traits and ear development of maize cultivars ( Zea mays L. ) with different plant types. Acta Agric Jiangxi, 2010, 22(1): 5-7 (in Chinese with English abstract) [本文引用:2]
[20]	Ma B L, Morrison M J, Dwyer L M. Canopy light reflectance and field greenness to assess nitrogen fertilization and yield of maize. Agron J, 1996, 88: 915-920 [本文引用:1]
[21]	刘仲发, 勾玲, 赵明, 张保军. 遮阴对玉米茎秆形态特征、穿刺强度及抗倒伏能力的影响. 华北农学报, 2011, 26(4): 91-96 Liu Z F, Gou L, Zhao M, Zhang B J. Effects of shading on stalk morphological characteristics, rind penetration strength and lodging-resistance of maize. Acta Agric Boreali-Sin, 2011, 26(4): 91-96 (in Chinese with English abstract) [本文引用:1]
[22]	贾士芳, 董树亭, 王空军, 张吉旺, 刘鹏. 弱光胁迫对玉米产量及光合特性的影响. 应用生态学报, 2007, 18: 2456-2461 Jia S F, Dong S T, Wang K J, Zhang J W, Liu P. Effects of weak light stress on grain yield and photosynthetic traits of maize. Chin J Appl Ecol, 2007, 18: 2456-2461 (in Chinese with English abstract) [本文引用:1]
[23]	Hashemi-Dezfouli A, Herbert S J. Intensifying plant density response of corn with artificial shade. Agron J, 1992, 84: 547-551 [本文引用:1]
[24]	张亚勤, 杨华, 祁志云, 袁亮, 王楠, 金川, 邱正高. 遮光胁迫对玉米植株性状的影响研究. 中国农学通报, 2011, 27(33): 40-43 Zhang Y Q, Yang H, Qi Z Y, Yuan L, Wang N, Jin C, Qiu Z G. Effect of light stress on the plants characters of maize inbred lines. Chin Agric Sci Bull, 2011, 27(33): 40-43 (in Chinese with English abstract) [本文引用:1]
[25]	贾士芳, 董树亭, 王空军, 张吉旺, 刘鹏. 玉米花粒期不同阶段遮光对籽粒品质的影响. 作物学报, 2007, 33: 1960-1967 Jia S F, Dong S T, Wang K J, Zhang J W, Liu P. Effect of shading on grain quality at different stages from flowering to maturity in maize. Acta Agron Sin, 2007, 33: 1960-1967 (in Chinese with English abstract) [本文引用:1]
[26]	任国玉, 郭军, 徐铭志, 初子莹, 张莉, 邹旭恺, 李庆祥, 刘小宁. 近50年中国地面气候变化基本特征. 气象学报, 2005, 63: 942-956 Ren G Y, Guo J, Xu M Z, Chu Z Y, Zhang L, Zou X K, Li Q Y, Liu X N. Climate changes of Chinese mainland over the past half century. Acta Meteorol Sin, 2005, 63: 942-956 (in Chinese with English abstract) [本文引用:1]
[27]	杨彦武, 于强, 王靖. 近40年华北及华东局部主要气候资源要素的时空变异性. 资源科学, 2004, 26(4): 45-50 Yang Y W, Yu Q, Wang J. Spatio-temporal variations of principal climatic factors in north china and part of east China within past 40 years. Resour Sci, 2004, 26(4): 45-50 (in Chinese with English abstract) [本文引用:1]
[28]	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: 3497-3505 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. Sci Agric Sin, 2012, 45: 3497-3505 (in Chinese with English abstract) [本文引用:1]
[29]	张祖建, 朱庆森, 王志琴. 水稻胚乳细胞计数方法研究. 江苏农学院学报, 1996, 17(6): 7-11 Zhang Z J, Zhu Q S, Wang Z Q. A method of counting endosperm cells in rice grain. Jiangsu Agric Res, 1996, 17(6): 7-11 (in Chinese with English abstract) [本文引用:1]
[30]	何照范. 粮油籽粒品质及其分析技术. 北京: 农业出版社, 1985. pp 329-331 He Z F. Oil Seeds Quality and Analysis Techniques. Beijing: China Agriculture Press, 1985. pp 329-331(in Chinese with English abstract) [本文引用:2]
[31]	Cui H Y, James J C, Jin L B, Zhang J W. Effects of shading on spike differentiation and grain yield formation of summer maize in the field. Int J Biometeorol, 2015, 59: 1189-1200 [本文引用:2]
[32]	宋凤斌, 童淑媛. 不同株型玉米的干物质积累、分配及转运特征. 江苏农业学报, 2010, 26: 700-705 Song F B, Tong S Y. Characteristics of dry matter accumulation, distribution and translocation in maize with different plant types. Jiangsu J Agric Sci, 2010, 26: 700-705 (in Chinese with English abstract) [本文引用:1]
[33]	戴明宏, 陶洪斌, 王利纳, 王璞. 不同氮肥管理对春玉米干物质生产、分配及转运的影响. 华北农学报, 2008, 23(1): 154-157 Dai M H, Tao H B, Wang L N, Wang P. Effects of different nitrogen managements on dry matter accumulation, partition and transportation of spring maize ( Zea mays L. ). Acta Agric Boreali-Sin, 2008, 23(1): 154-157 (in Chinese with English abstract) [本文引用:1]
[34]	刘伟, 张吉旺, 吕鹏, 杨今胜, 刘鹏, 董树亭, 李登海, 孙庆泉. 种植密度对高产夏玉米登海661产量及干物质积累与分配的影响. 作物学报, 2011, 37: 1301-1307 Liu W, Zhang J W, Lyu P, Yang J S, Liu P, Dong S T, Li D H, Sun Q Q. Effect of plant density on grain yield dry matter accumulation and partitioning in summer maize cultivar Denghai 661. Acta Agron Sin, 2011, 37: 1301-1307 (in Chinese with English abstract) [本文引用:1]
[35]	孙小玲, 许岳飞, 马鲁沂, 周禾. 植株叶片的光合色素构成对遮阴的响应. 植物生态学报, 2010, 34: 989-999 Sun X L, Xu Y F, Ma L Y, Zhou H. A review of acclimation of photosynthetic pigment composition in plant leaves to shade environment. Chin J Plant Ecol, 2010, 34: 989-999 (in Chinese with English abstract) [本文引用:1]
[36]	李潮海, 栾丽敏, 尹飞, 王群, 赵亚丽. 弱光胁迫对不同基因型玉米生长发育和产量的影响. 生态学报, 2005, 25: 824-830 Li C H, Luan L M, Yin F, Wang Q, Zhao Y L. Effects of seedling shading and light intensity transfer on photosynthetic efficiency of different maize hybrids. Acta Ecol Sin, 2005, 31: 381-385 (in Chinese with English abstract) [本文引用:1]
[37]	崔海岩, 靳立斌, 李波, 董树亭, 刘鹏, 赵斌, 张吉旺. 遮阴对夏玉米干物质积累及养分吸收的影响. 应用生态学报, 2013, 24: 3099-3105 Cui H Y, Jin L B, Li B, Dong S T, Liu P, Zhao B, Zhang J W. Effects of shading on dry matter accumulation and nutrient absorption of summer maize. Chin J Appl Ecol, 2013, 24: 3099-3105 (in Chinese with English abstract) [本文引用:2]
[38]	贾士芳, 董树亭, 王空军, 张吉旺, 刘鹏. 弱光胁迫影响夏玉米光合效率的生理机制初探. 植物生态学报, 2010, 34: 1439-1447 Jia S F, Dong S T, Wang K J, Zhang J W, Liu P. Physiological mechanism of shading stress on photosynthetic efficiency in summer maize ( Zea mays L. ). Chin J Plant Ecol, 2010, 34: 1439-1447 (in Chinese with English abstract) [本文引用:1]
[39]	史建国, 崔海岩, 赵斌, 董树亭, 刘鹏, 张吉旺 . 花粒期光照对夏玉米产量和籽粒灌浆特性的影响. 中国农业科学 , 2013, 46: 4427-4434 Shi J G, Cui H Y, Zhao B, Dong S T, Liu P, Zhang J W. Effect of light on yield and characteristics of grain-filling of summer maize from flowering to maturity. Sci Agric Sin, 2013, 46: 4427-4434 (in Chinese with English abstract) [本文引用:2]
[40]	Ren B Z, Zhang J W, Dong S T, Liu P, Zhao B. Effects of duration of water logging at different growth stages on grain growth of summer maize ( Zea mays L. ) under field conditions. J Agron Crop Sci, 2016, 202: 564-575 [本文引用:1]

2010

0.0

贾士芳, 李从锋, 董树亭, 张吉旺. 花后不同时期遮光对玉米粒重及品质影响的细胞学研究. 中国农业科学, 2010, 43: 911-921

Jia S

, Li C

, Dong S

, Zhang J

. Effects of shading at different stages after anthesis on maize grain weight and quality at cytology level. Sci Agric Sin, 2010, 43: 911-921 (in Chinese with English abstract)

... 玉米胚乳质量占籽粒质量的80%左右, 胚乳细胞的数量和充实度对粒重有决定作用^[1,2], 同时也代表着#cod#x0201c ...

... 贾士芳等^[1]研究表明, 遮阴显著降低玉米籽粒中胚乳细胞数、淀粉及籽粒充实度, 最终降低粒重 ...

2008

0.0

秦大鹏, 刘鹏, 王空军, 董树亭, 张吉旺, 刘月娥. 高淀粉玉米籽粒胚乳细胞增殖与淀粉积累的关系. 山东农业科学, 2008, (8): 35-39

Qin D

, Liu

, Wang K

, Dong S

, Zhang J

, Liu Y

. Relationship between endosperm cell proliferation and starch accumulation in high-starch maize grain. Shand ong J Agric Sin, 2008, (8): 35-39 (in Chinese with English abstract)

... 玉米胚乳质量占籽粒质量的80%左右, 胚乳细胞的数量和充实度对粒重有决定作用^[1,2], 同时也代表着#cod#x0201c ...

... 胚乳细胞越多, 积累的淀粉粒越多, 充实度越高, 粒重越高^[2] ...

2013

0.0

徐云姬, 顾道健, 张博博, 张耗, 王志琴, 杨建昌. 玉米果穗不同部位籽粒激素含量及其与胚乳发育和籽粒灌浆的关系. 作物学报, 2013, 39: 1452-1461

Xu Y

, Gu D

, Zhang B

, Zhang

, Wang Z

, Yang J

. Hormone contents in kernels at different positions on an ear and their relationship with endosperm development and kernel filling in maize. Acta Agron Sin, 2013, 39: 1452-1461 (in Chinese with English abstract)

Kernels at the upper position of a maize ear usually show poorer filling and lower weight in contrast to those at the basal position. The mechanism has not been understood . The experiments were conducted using a maize cultivar Denghai 11, grown in both field and greenhouse conditions, to determine number of endosperm cells, kernel filling rate, and contents of zeatin (Z) + zeatin riboside (ZR), indole-3-acetic acid (IAA), abscisic acid (ABA), and gibberellin-3 (GA 3 ) in the kernels at different positions on an ear from silking to maturity. The results indicated that the maximum number of endosperm cells, maximum division rate of endosperm cell s, mean endosperm cell division rate, maximum kernel filling rate, mean kernel filling rate and 100-kernel weight for kernels showed an order of basal position > middle position> upper position. During the active endosperm development and the active kernel filling period, contents of Z+ZR, IAA and ABA in kernels were the greatest at the basal position, the mediate at the middle position, and the least at the upper position on an ear. The two experiments exhibited similar results. The endosperm cell division rate and kernel filling rate were very significantly and positively correlated with contents of Z+ZR, IAA, and ABA in kernels, whereas significantly and negatively correlated with content of GA 3 . The results suggested that a lower kernel weight at the upper position on a maize ear is mainly due to a smaller division rate of endosperm cells, leading to less number of endosperm cells, which is closely associated with lower contents of Z+ZR, IAA, and ABA, as well as higher content of GA 3 in the kernels during kernel filling.

以玉米品种登海 11 为材料 , 分别进行大田和温室试验 , 观察灌浆期果穗不同部位籽粒玉米素 (Z)+ 玉米素核苷 (ZR) 、吲哚 -3- 乙酸 (IAA) 、脱落酸 (ABA) 和赤霉素 (GA 3 ) 含量变化及其与胚乳发育和籽粒灌浆的关系。结果显示 , 籽粒最大胚乳细胞数目、最大胚乳细胞增殖速率及平均速率、最大灌浆速率、平均灌浆速率和百粒重表现为果穗下部籽粒 > 中部籽粒 > 上部籽粒。在胚乳细胞活跃增殖期或活跃灌浆期 , 籽粒 Z+ZR 、 IAA 和 ABA 含量以果穗下部籽粒最高 , 中部籽粒其次 , 上部籽粒最低。 GA 3 含量则为果穗上部籽粒 > 中部籽粒 > 下部籽粒。两个试验的结果趋势一致。胚乳细胞增殖速率和籽粒灌浆速率与籽粒 Z+ZR 、 IAA 和 ABA 含量呈极显著正相关 , 与籽粒 GA 3 含量呈显著负相关。说明玉米果穗上部籽粒轻主要是由于这些籽粒的胚乳细胞增殖速率小 , 导致其胚乳细胞数少 , 这与其灌浆期较低的 Z+ZR 、 IAA 和 ABA 含量及较高的 GA 3 含量有密切关系。

... 胚乳细胞增殖速率快, 数量多, 有助于光合同化产物在籽粒中积累和淀粉粒形成, 提高籽粒充实度, 进而获得高产^[3,4,5] ...

2005

0.0

孙庆泉, 吴元奇, 胡昌浩, 董树亭, 荣廷昭, 张颖. 不同产量潜力玉米籽粒胚乳细胞增殖与籽粒充实期的生理活性. 作物学报, 2005, 31: 612-618

Sun Q

, Wu Y

, Hu C

, Dong S

, Rong T

, Zhang

. Physiological activities and multiplication of endosperm cell at filling stage of kernels with different yield in maize. Acta Agron Sin, 2005, 31: 612-618 (in Chinese with English abstract)

... 胚乳细胞增殖速率快, 数量多, 有助于光合同化产物在籽粒中积累和淀粉粒形成, 提高籽粒充实度, 进而获得高产^[3,4,5] ...

2016

0.0

... 胚乳细胞增殖速率快, 数量多, 有助于光合同化产物在籽粒中积累和淀粉粒形成, 提高籽粒充实度, 进而获得高产^[3,4,5] ...

2005

0.0

张智猛, 戴良香, 胡昌浩, 董树亭, 王空军, 宁堂原. 玉米灌浆期水分差异供应对籽粒淀粉积累及其酶活性的影响. 植物生态学报, 2005, 29: 636-643

Zhang Z

, Dai L

, Hu C

, Dong S

, Wang K

, Ning T

. Effects of different water treatments on starch accumulation and related enzyme activity in grain filling period. Chin J Plant Ecol, 2005, 29: 636-643 (in Chinese with English abstract)

... 玉米籽粒积累的淀粉数量决定其产量和品质^[6,7] ...

2010

0.0

谷岩, 王振民, 何文安, 梁烜赫, 吴春胜. 不同类型玉米品种籽粒胚乳细胞增殖与籽粒建成的关系. 西北农林科技大学学报, 2010, 38(1): 90-96

, Wang Z

, He W

, Liang X

, Wu C

. Relationship between endosperm cell proliferation and seed formation in maize of different types. J Northwest A&F Univ, 2010, 38(1): 90-96 (in Chinese with English abstract)

【Objective】 The proliferation process of endosperm cell and its relationship with the correlative properties of grain in filling stage were studied.【Method】 The upright maize varieties Xianyu 335 and Zhengdan 958 and the common maize varieties Changcheng 799 and Nongda 364 were chosen as the experiment material.Central kernels were collected after the pollination stage,endosperm cell number,kernel weight,filling rate,starch content,starch phosphorylase and invertase were determined,and the relationship was analysed.【Result】 The endosperm cell,grain weight and starch content kept increasing in the entire grain-filling period and the increase appeared like an “S” shape.The endosperm cell number of corn grain was determined on the 20 th day after pollination.The endosperm cell number had significantly positive correlation with grain weight,starch content and the filling rate of kernels.The filling rate of kernels was evidently affected by physiological activity of kernels.The activities of invertase and starch phosphorylase appeared a single peak during the filling period,and the peak in upright maize occurred later than the common maize varieties. The endosperm cell had significantly positive correlation with the activities of invertase and starch phosphorylase.【Conclusion】 The endosperm cell number had significantly positive correlation with the kernel weight,filling rate,starch content,starch phosphorylase and invertase.In order to improve high-super yield,we must enhance the rate of endosperm cell proliferation and the filling before the filling period,and ensure the transportation of photosynthetic production to kernels quickly and repletely.

【目的】研究不同类型玉米品种籽粒胚乳细胞增殖与籽粒充实期相关性状的关系。【方法】以紧凑型玉米品种(先玉335和郑单958)和平展型玉米品种(长城799和农大364)为试验材料，于授粉后3，5，7，10，15，20，25,30,35,40,45,50,55，60 d取果穗中部籽粒，观测胚乳细胞数目、单粒质量、灌浆速率、淀粉含量、籽粒淀粉磷酸化酶和蔗糖转化酶活性等指标的变化动态，并分析不同玉米品种籽粒胚乳细胞增殖动态与籽粒灌浆期相关生理指标的关系。【结果】随着灌浆期的延长，不同玉米品种籽粒胚乳细胞数、单粒质量、淀粉含量均呈“S”型曲线增长，籽粒胚乳细胞数在授粉后20 d基本稳定。胚乳细胞数与单粒质量、淀粉含量和灌浆速率均呈极显著正相关；籽粒充实期的生理状态影响着籽粒灌浆，淀粉磷酸化酶和蔗糖转化酶活性在灌浆期呈现单峰曲线，但峰值出现时间因品种而异，紧凑型玉米酶活性最高点出现时间要晚于平展型玉米；籽粒胚乳细胞数与这两种酶活性均呈显著相关。【结论】玉米籽粒中胚乳细胞数与胚乳细胞的充实状态、籽粒单粒质量、灌浆速率、淀粉含量、淀粉磷酸化酶蔗糖转化酶活性均呈显著正相关；要提高玉米产量，应在灌浆前期提高胚乳细胞增殖速率和灌浆速率，保证光合产物快速充分地运送到籽粒中去，使籽粒淀粉积累达到最佳状态，进而达到高产。

... 玉米籽粒积累的淀粉数量决定其产量和品质^[6,7] ...

... 的潜力, 也代表该潜力的实现程度, 而单个胚乳细胞的充实状态也对产量的提高具有重要作用^[7] ...

2014

0.0

李栋梁, 李小刚, 顾蕴洁, 王忠. 不同类型水稻品种胚乳发育的研究. 中国农业科学, 2014, 47: 3757-3768

Li D

, Li X

, Gu Y

, Wang

. Investigation of endosperm cell development of different rice varieties. Sci Agric Sin, 2014, 47: 3757-3768 (in Chinese with English abstract)

... 淀粉作为光合作用的主要产物, 通过酶催化转运至胚乳细胞贮藏, 其积累速率和持续活跃时间决定粒重^[8,9], 即胚乳细胞越多, 淀粉积累量越多, 充实度越好, 粒重越高^[10] ...

... S2), 最终表现为籽粒发育进程延缓, 淀粉积累量下降, 籽粒库容量变小, 充实度下降, 粒重降低, 相反, 增光条件下胚乳细胞增殖速度快、数目增多(表8), 籽粒积累同化产物的速度加快, 容纳较多的淀粉粒^[8], 提高产量 ...

2009

0.0

王贺正, 马均, 李旭毅, 张荣萍. 水分胁迫对水稻籽粒灌浆及淀粉合成有关酶活性的影响. 中国农业科学, 2009, 42: 1550-1558

Wang H

, Ma

, Li X

, Zhang R

. Effects of water stress on grain filling and activities of enzymes involved in starch synthesis in rice. Sci Agric Sin, 2009, 42: 1550-1558 (in Chinese with English abstract)

1989

0.0

李伯航, 崔彦宏. 夏玉米胚乳细胞建成与粒重关系. 河北农业大学学报, 1989, 12(4): 39-45

Li B

, Cui Y

. Studies on the relationship between endosperm cell development and kernel weight of summer maize. J Agric Univ Hebei, 1989, 12(4): 39-45 (in Chinese with English abstract)

... 遮阴导致玉米籽粒胚乳细胞数和充实度降低^[10] ...

2011

0.0

杜成凤, 李潮海, 刘天学, 赵亚丽. 遮阴对两个基因型玉米叶片解剖结构及光合特性的影响. 生态学报, 2011, 31: 6633-6640

Du C

, Li C

, Liu T

, Zhao Y

. Response of anatomical structure and photosynthetic characteristics to low light stress in leaves of different maize genotypes. Acta Ecol Sin, 2011, 31: 6633-6640 (in Chinese with English abstract)

Prolonged rainy weather during the maize growing season can lead to insufficient light, and thus dramatically lower yields. Maize, Zea mays L., is one of the most important food crops in China, and around the world, and a high and stable yield is necessary to guarantee food safety. Previous studies into the effects of low light stress on maize have predominantly focused on aboveground biomass, leaf area, chlorophyll content, net photosynthetic rate, root activity, soluble protein content, grain yield, nutrient levels, and the activity of some enzymes, including superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), sucrose-phosphate synthase (SPS), sucrose synthase (SS), soluble starch synthase (SSS), granule-bound-starch synthase (GBSS), adenosine diphosphate glucose pyrophorylase (ADPGPPase), and uridine diphosphate glucose pyrophorylase (UDPGPPase). However, the potentially damaging effects of low light stress on the anatomical structure of maize photosynthetic systems have seldom been mentioned in the literature, and the relationships between anatomical structure and photosynthetic characteristics remain unclear. Our objective was to determine the effects of low light stress on the anatomical structure of maize photosynthetic systems, and the relationship between photosynthetic characteristics and anatomical structure, in different maize cultivars at the seedling stage. We experimented on the seedling leaves of two maize cultivars: the shade-tolerant cultivar Zhengdan 958 (ZD958) and the shade-sensitive cultivar Yuyu 22(YY22). These were grown in experimental pots in the field. 50% shade treatment was used, with non-shaded seedlings as controls. Under low light stress, plant growth was inhibited; leaves became thinner, numbers of chloroplasts per unit of leaf area decreased, number of grana increased, grana thickness increased, and the thickness of the grana lamellae increased. In the shade-sensitive cultivar YY22, most of the chloroplasts became dysplastic, the stroma lamellae swelled and the grana lamellae became loose. In contrast, in the shade-tolerant cultivar ZD958 the chloroplasts were almost normal with some healthy parts. The net photosynthetic rate ( Pn ), photochemical efficiency ( Fv/Fm ), and actual photochemical efficiency ( Φ PS Ⅱ ) all decreased under shade. Chlorophyll content, intercellular CO 2 concentration ( Ci ) and non-photochemical quenching ( NPQ ) increased; however these were all reversible. The values of all variables investigated were significantly higher or lower in the shade-sensitive cultivar YY22 than in the corresponding control. Shading had no significant influence on the shade-tolerant cultivar ZD958 except for Pn and Ci . We speculate, therefore, that low light stress has different significant effects on the anatomy and photosynthesis of maize with different genotypes. The shade-tolerant cultivar ZD958 maintains more normal anatomical structure and higher photosynthetic function than the shade-sensitive YY22 under low light stress; these are the major anatomical and physiological traits necessary for shade tolerance.

以耐荫性不同的玉米品种郑单958(ZD958,耐荫性较强)和豫玉22(YY22,耐荫性较弱)为材料,研究了苗期50%遮荫对玉米叶片形态结构和光合特性的影响。结果表明:形态结构上,苗期遮荫处理后,玉米叶片变薄,单位面积叶绿体数目减少,基粒数、基粒厚度和片层数增加,但是YY22的叶绿体大部分发育不良,肿胀呈球形,基粒片层和基质片层出现不同程度的松散;而ZD958大部分叶绿体结构良好,各部分发育基本正常。光合特性上,弱光胁迫使玉米叶片叶绿素含量升高,净光合速率( Pn )、PSⅡ最大光化学量子产量( Fv/Fm )和实际光化学效率( Φ PS Ⅱ )降低,胞间CO 2 浓度( Ci )和非光化学猝灭( NPQ )增加,但是YY22的 Pn、Fv/Fm 和 Φ PS Ⅱ 显著下降, Ci 和 NPQ 显著升高;而ZD958的 Fv/Fm 和 Φ PS Ⅱ 下降幅度较小,且 NPQ 增加亦不显著。研究结果提示,弱光胁迫对玉米叶片形态结构和光合特性影响较大,且存在基因型差异,耐荫性较强的品种对弱光环境的适应性较强。

... 杜成凤等^[11]和史建国等^[12]研究认为, 充足的光照有助于促进玉米光合作用, 也是玉米高产的基础 ...

2015

0.0

史建国, 朱昆仑, 曹慧英, 董树亭, 刘鹏, 赵斌, 张吉旺. 花粒期光照对夏玉米干物质积累和养分吸收的影响. 应用生态学报, 2015, 26: 46-52

Shi J

, Zhu K

, Cui H

, Zhao

, Dong S

, Liu

, Zhang J

. Effect of light from flowering to maturity stage on dry matter accumulation and nutrient absorption of summer maize. Chin J Appl Ecol, 2015, 26: 46-52 (in Chinese with English abstract)

Using Denghai 605 (DH605) as the experimental material, shading (S) and increasing light (L) treatments from flowering to maturity stage were designed in a field experiment to explore effects of light on dry matter accumulation and nutrient absorption of summer maize. Results showed that grain yield, dry matter accumulation and nutrient absorption decreased significantly after shading but increased after increasing light. Yields in S treatment from 2011 to 2013 were reduced by 59.4%, 79.0% and 60.6% compared to those in CK, while that in L treatment were increased by 16.3%, 12.9% and 6.8%, respectively. The relative N and P absorption increased to a certain extent because of the greater effect of shading on dry matter accumulation than that of N and P absorption. After shading, K absorption of whole plant corn decreased significantly to a greater extent than that of dry matter accumulation decrease. The proportion of nutrient allocated to grains decreased significantly after shading. Dry matter accumulation and N and P absorption rates increased significantly after increasing light, and effects of increasing light on N and P absorption were greater than that of dry matter accumulation. Nutrient accumulation and the proportion allocated to grain increased significantly after increasing light.

选用登海605(DH605)为试验材料,在大田条件下通过花粒期遮阴(S)和增光(L)研究不同光照对花后夏玉米产量、干物质积累和氮、磷、钾养分吸收与运转特性的影响.结果表明: 遮阴后夏玉米产量、干物质积累量及氮、磷、钾养分吸收量均显著降低;而增光有利于夏玉米干物质积累量及氮、磷、钾养分吸收量和产量的提高.2011—2013年,遮阴处理的产量较对照(自然光下)分别降低了59.4%、79.0%、60.6%;增光后产量分别增加16.3%、12.9%、6.8%.遮阴对干物质积累的影响大于其对植株氮、磷吸收量的影响,植株体内氮、磷等养分含量有所上升,而钾吸收量的下降幅度大于干物质积累,钾素相对含量降低,氮、磷、钾向籽粒的分配比例降低.增光后干物质积累和氮、磷吸收量显著上升,但对氮、磷吸收的影响更大;增光条件下,氮、磷、钾等养分向籽粒的分配比例增加.

... 杜成凤等^[11]和史建国等^[12]研究认为, 充足的光照有助于促进玉米光合作用, 也是玉米高产的基础 ...

2008

0.0

曾希柏. 作物生长中光照和氮肥施用量的相互关系研究. 土壤学报, 2008, 37: 380-386

Zeng X

. Research on the correlation of light and nitrogen application in crop growth. Acta Pedol Sin, 2008, 37: 380-386 (in Chinese with English abstract)

By using the method of pot culture experiment in the case of artificial climate the result indicated that the effects of light intensity(X 1 )and N application(X 2 ) on lettuce yield(y)were mathematically represented by the following equation in this study: y=-0.375+0.0230X 1 +9.421X 2 +0.0251X 1 X 2 -0.131×10 -4 X 1 2 -17.794X 2 2 ;The efficiency of N fertilizer was influenced by light intensity. A quadratic multinomial equation y=b 0 +b 1 X + b 2 X 2 , could be used to describe the correlation between N application and lettuce yield. Also, quadratic or cubic multinomial equation could be used to describe the correlation between light and lettuce yield. Accordingly, the effects of light intensity(X 1 )on N application (X 2 ) were mathematically represented by the following equation: X 2 =0.446-0.477×10 -2 X 1 +0.269×10 -4 X 1 2 -0.394×10 -7 X 1 3 . Moreover, nitrogen nutrition of lettuce was bigger influenced by light and N application.

应用盆栽试验,在人工气候箱内进行的研究结果表明;光照强度(X 1 ,μmol/m 2 ·s)和氮肥施用量(X 2 ,g/pot)均对莴笋生长及生物量产生影响,且二者的变化与莴笋生物量的关系可表示为y=-0.375+0.0230X 1 +9.42lX 2 +0.025lX 1 X 2 -0.131×10 -4 X 1 2 -17.794X 2 2 形式;不同光照强度下氮肥施用量与莴笋生物量的关系均可表示为y=b 0 +b 1 X+b 2 X 2 形式;不同氮肥施用量下,光照强度与莴笋生物量的关系亦可用二次或三次多项式来表示。根据研究结果,还求得了氮肥施用量(X 2 )与光照强度(X 1 )的关系为;X 2 =0.446-0.477×10 -2 X 1 +0.269×10 -4 X 1 2 -0.394×10 -7 X 1 3 .此外,本研究还探讨了光照强度、氮肥施用量与莴笋的氮素营养状况的关系。

... 曾希柏等^[13]研究指出, 在一定范围内植物生物量随光强的增加而提高, 玉米产量也会在一定范围随光照强度的上升而增加 ...

2016

0.0

... 光是玉米光合作用的能量来源, 光照变化对植株光合作用、同化产物形成有极为显著的影响^[14] ...

2013

0.0

崔海岩, 靳立斌, 李波, 赵斌, 董树亭, 刘鹏, 张吉旺. 大田遮阴对夏玉米光合特性和叶黄素循环的影响. 作物学报, 2013, 39: 478-485

Cui H

, Jin L

, Li

, Zhao

, Dong S

, Liu

, Zhang J

. Effects of shading on photosynthetic characteristics and xanthophyll cycle of summer maize in the field. Acta Agron Sin, 2013, 39: 478-485 (in Chinese with English abstract)

... 开花后的光照时数、光照强度和叶面积决定了80%的玉米产量^[15] ...

... 1951年以来, 气候变化加剧, 日照时数减少, 太阳辐射减弱, 年平均气温不断升高, 极端天气频繁发生^[26,27], 以黄淮海区为例, 在夏玉米生长期内(6月至10月), 尤其是花粒期, 阴雨寡照天气频繁发生, 导致玉米总产量减少3%~ 6%^[15] ...

2006

0.0

张吉旺, 董树亭, 王空军, 胡昌浩, 刘鹏. 遮阴对夏玉米生长发育和产量的影响. 应用生态学报, 2006, 17: 657-662

Zhang J

, Dong S

, Wang K

, Hu C

, Liu

. Effects of shading on the growth, development and grain yield of summer maize. Chin J Appl Ecol, 2006, 17: 657-662 (in Chinese with English abstract)

Under field condition,this paper studied the effects of shading on the growth, development,and grain yield of summer maize varieties ND108 and YD13.The resul ts showed that shading decreased maize yield significantly,and the effect was d iffered with different shading period and intensity.With a shading intensity of 50% and 90%,the grain yield of ND108 and YD13 was decreased by 67.5% and 79.4% ,and 82.9% and 86.7% when shading at flowering-maturing stage,and by 34.1% and 55.3%,and 47.2%,65.7% when shading at joining-flowering stage,respectively. Shading at seedling-joining stage had a relatively smaller effect,with the grai n yield decreased by 16.9% and 24.5%,and 18.9% and 24.3%,respectively.Shading had a larger effect on YD13 than on ND108,and the effect of shading period was larger than that of shading intensity.Under shading,the growth and developmen t of maize was retarded,and the effect was increased with increasing shading in tensity.Shading at joining-flowering stage affected spike differentiation signi ficantly,reflecting in the marked decrease of the numbers of silks and tassels, and the effect was also larger on YD13 than on ND108.The leaf and plant growth was restrained significantly when shading at seedling stage and at flowering-m aturing stage.

Key Laboratory of Crop Biology of Shandong Province,College of Agronomy,Shand ong Agricultural University,Tai’an 271018,China

在大田条件下研究了不同时期和不同程度遮荫对夏玉米产量及生长发育的影响.结果表明，遮荫显著降低玉米产量.不同时期遮荫对其影响不同，花粒期（从开花到成熟期）遮荫的影响最显著，农大108（ND108）和掖单13号（YD13）遮荫50%、90%处理分别减产67. 5%、79.4%和82.9%和86.7%，其次是穗期（从拔节到开花期）遮荫，ND108和YD13分别减产34.1%、55.3%和47.2%、65.7%，而苗期（从出苗到拔节期）遮荫对其影响相对较小，ND108和YD13分别减产16.9%、24.5%和18.9%、24.3%.遮荫对YD13产量的影响大于ND10 8.遮荫时期对玉米产量的影响显著地大于遮荫程度.遮荫后两个玉米品种的生育进程都延迟，并随着遮荫程度的增加，对其影响加剧.穗期遮荫显著影响玉米的穗分化，花丝数和雄穗分枝数显著降低，对YD13的影响大于ND108.苗期和穗期遮荫显著抑制玉米叶面积、株高和茎节的生长.

... 遮阴导致夏玉米生育时期推迟, 株高、茎粗、叶面积显著减少^{[16,17,18,19,20]}, 倒伏率升高^[21], 叶片光合特性和生殖器官发育受到影响^[22,23,24], 籽粒败育^[25], 产量下降^[18,19] ...

2008

0.0

王洋, 齐晓宁, 邵金锋, 刘兆永, 李泽兴. 光照强度对不同玉米品种生长发育和产量构成的影响. 吉林农业大学学报, 2008, 30: 769-773

Wang

, Qi X

, Shao J

, Liu Z

, Li Z

. Effects of light intensity at full growing stage on the growth and yield of different maize varieties. J Jilin Agric Univ, 2008, 30: 769-773 (in Chinese with English abstract)

2009

0.0

付景, 李潮海, 赵久然, 马丽, 刘天学. 玉米品种耐阴性指标的筛选与评价. 应用生态学报, 2009, 20: 2705-2709

, Li C

, Zhao J

, Ma

, Liu T

. Shade-tolerance indices of maize: Selection and evaluation. Chin J Appl Ecol, 2009, 20: 2705-2709 (in Chinese with English abstract)

2010

0.0

王秀萍, 刘天学, 李潮海, 李大鹏. 遮光对不同株型玉米品种农艺性状和果穗发育的影响. 江西农业学报, 2010, 22(1): 5-7

Wang X

, Liu T

, Li C

, Li D

. Effects of shading on agronomic traits and ear development of maize cultivars ( Zea mays L. ) with different plant types. Acta Agric Jiangxi, 2010, 22(1): 5-7 (in Chinese with English abstract)

1996

0.0

2011

0.0

刘仲发, 勾玲, 赵明, 张保军. 遮阴对玉米茎秆形态特征、穿刺强度及抗倒伏能力的影响. 华北农学报, 2011, 26(4): 91-96

Liu Z

, Gou

, Zhao

, Zhang B

. Effects of shading on stalk morphological characteristics, rind penetration strength and lodging-resistance of maize. Acta Agric Boreali-Sin, 2011, 26(4): 91-96 (in Chinese with English abstract)

Under field condition,effects of shading on stalk mechanical and rind penetration strength were investigated using three maize cultivars differing in lodging resistant ability,i.e.JK519,CS1 and ZD958.The results showed that JK519 with a large spike under thinned planting density was sensitive to shade stress,its plant height and ear height were decreased significantly,and LAI(leaf area index) was also declined.With lower internodes diameter and shorter internodes length at basal stem,the ratio of dry weight to length of internodes was reduced in shading condition.The rind penetration strength of JK519 was decreased by 36.4% and 66.0% under a shading intensity of 30% and 60%,respectively;and field lodging seriously.However,with a shading intensity of 30%,CS1(a compact and tolerant to high density type) was declined slightly in the ear height,LAI and internodes diameter,and its rind penetration strength was only decreased by 5.9%.The rind penetration strength was decreased obviously by 60 % shading intensity,lodging heavily.Meanwhile,the finally maize yield,the number of harvest ear and grain per ear,and weight of 1 000 kernels of three maize cultivars were decreased significantly with increasing shading intensity.The 30% shading intensity should be definite as an appraise and selection condition of the mechanical strength of maize high yield and lodging resistance.

1. College of Agronomy, Northwest Agriculture and Forestry University, Yangling 712100, China; 2. Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China; 3. Key Laboratory of Oasis Ecology Agriculture of Xinjiang Construction Corps, Shihezi University, Shihezi 832003, China

大田条件下,研究遮荫处理对玉米不同类型品种的茎秆形态特征及抗倒力学强度的影响。结果表明:稀植大穗品种JK519对弱光胁迫反应较为敏感,遮荫处理后玉米株高和穗位高均显著下降,叶面积指数降低。基部节间缩短、直径变小,单位节间长干质量减少。茎秆穿刺强度在30%和60%遮荫条件下分别下降36.4%和66.0%,田间倒伏率严重。而紧凑耐密品种CS1在遮荫30%胁迫下穗位高、LAI下降不明显;节间缩短,但直径和单位节间长干质量变化不大;穿刺强度仅下降5.9%,田间倒伏较轻;在遮荫60%胁迫下,穿刺强度明显下降,倒伏加重。同时各个品种均随遮荫程度的增加,最终玉米产量、收获穗数、单穗粒数和千粒重明显降低。因此,30%遮荫处理(弱光胁迫)可推荐作为玉米高产抗倒品种茎秆力学强度鉴定和筛选条件之一。

2007

0.0

贾士芳, 董树亭, 王空军, 张吉旺, 刘鹏. 弱光胁迫对玉米产量及光合特性的影响. 应用生态学报, 2007, 18: 2456-2461

Jia S

, Dong S

, Wang K

, Zhang J

, Liu

. Effects of weak light stress on grain yield and photosynthetic traits of maize. Chin J Appl Ecol, 2007, 18: 2456-2461 (in Chinese with English abstract)

通过在玉米授粉前0～14d、授粉后1～14d、授粉后15～28d3个时期分别进行遮光处理,研究了弱光胁迫对两个玉米品种费玉3号和泰玉2号产量及光合特性的影响.结果表明:各时期弱光胁迫均使玉米产量降低,其中授粉前0～14d弱光处理的产量降幅最大,费玉3号对弱光胁迫的反应较泰玉2号敏感.弱光胁迫后籽粒灌浆高峰出现时间延迟、灌浆速率慢、积累量小;弱光胁迫开始的时间越早,籽粒达到最大灌浆速率的时间(Tmax)越晚.弱光胁迫期内,玉米Chl(a+b)、Chla/b、光合速率(Pn)、光系统Ⅱ最大光化学效率(Fv/Fm)和光系统Ⅱ实际光化学效率(ФPSⅡ)显著下降,Chlb相对含量提高,胞间二氧化碳浓度(Ci)和非光化学淬灭系数(NPQ)则显著上升;胁迫结束后,玉米Chl(a+b)、Chla/b、Pn、Fv/Fm、ФPSⅡ、Ci和NPQ逐渐恢复接近自然光照条件(CK)水平,而Chlb相对含量下降.表明非气孔因素是弱光胁迫下玉米光合速率降低的原因之一.

1992

0.0

2011

0.0

张亚勤, 杨华, 祁志云, 袁亮, 王楠, 金川, 邱正高. 遮光胁迫对玉米植株性状的影响研究. 中国农学通报, 2011, 27(33): 40-43

Zhang Y

, Yang

, Qi Z

, Yuan

, Wang

, Jin

, Qiu Z

. Effect of light stress on the plants characters of maize inbred lines. Chin Agric Sci Bull, 2011, 27(33): 40-43 (in Chinese with English abstract)

Eight maize inbred lines were used as tested materials, with shading treatment (90% shading control) from seeding stage to maturity stage. The experiment of two factors split-plot design was carried out to study the shade-tolerance and plant traits’ changes of the maize inbred lines. The order of the shade-tolerance level of the materials was: 06CIn967-11>06CIn892-2>06CIn967-4>06CIn892-11>06CIn891-4>06CIn891-15>06C984-2>06 C984-8, the difference of the inbred lines of same source wasn’t significant; there was certain difference among inbred lines of different source. The plant and leaf growth, tassel differentiation was restrained significantly; the shading reaction between different genotype lines was different. The shading reaction of different character was also different. The reaction of the tassel differentiation was the strongest of all. The shade-tolerance type inbred lines should be selected based on the traits changes of different maize inbred lines.

以8个玉米自交系为供试材料，采用2因素裂区设计，在玉米苗期到成熟期遮阴（遮光率为90%），研究玉米自交系的耐阴性及其在遮光胁迫条件下的植株性状的变化。结果表明：参试材料耐阴湿的程度从高到低为：06CIn967-11>06CIn892-2>06CIn967-4>06CIn892-11>06CIn891-4>06CIn891-15>06C984-2>06C9 84-8，其中，同一来源的自交系耐阴性差异不显著，不同来源的自交系耐阴性有一定差异；在弱光下，玉米材料的雄穗分化、植株和叶片的生长受到了极显著抑制，不同基因型材料的植株性状对遮光的反应也不同；遮光对不同性状的影响不同，对雄穗分化的影响最大。因此可以根据不同材料性状的变化来筛选耐阴材料。

2007

0.0

贾士芳, 董树亭, 王空军, 张吉旺, 刘鹏. 玉米花粒期不同阶段遮光对籽粒品质的影响. 作物学报, 2007, 33: 1960-1967

Jia S

, Dong S

, Wang K

, Zhang J

, Liu

. Effect of shading on grain quality at different stages from flowering to maturity in maize. Acta Agron Sin, 2007, 33: 1960-1967 (in Chinese with English abstract)

Light deficiency from pollination to maturity is a common problem in summer maize ( Zea mays L.) production in Huang-Huai-Hai-River region in China. Some researches on rice ( Oryza sativa L.), wheat ( Triticum aestivum L.), and soybean [ Glycine max (L.) Merr.] have reported that light deficiency limited carbohydrate accumulation, and result in protein content increase in grain. However, the conclusion on maize is uncertain due to rare studies. To investigate the effects of low light intensity at different stages after pollination on grain quality and the activities of key enzymes of starch and protein syntheses, we conducted a field experiment by shading treatment in 2005–2006 with two maize cultivars, Feiyu 3 (high starch content) and Taiyu 2 (low starch content). The black fabric net was used for making light defiency with 45% sunlight transmission. The shading periods were 1–14 (S1), 15–28 (S2), and 29–42 (S3) days after pollination (DAP), respectively. Only medium position grains of the ear were sampled at 7, 14, 21, 28, 35, and 42 DAP to determinate the enzyme activities. Grain yield and quality parameters were measured after maturity. Compared with the control (no shading treatment, S0), the S1, S2, and S3 treatments showed obvious difference in grain quality and activities of tested enzymes, and the two cultivars had similar results. The grain yield, test weight, and starch content in the three shading treatments were significantly lower than those in control, while the water content, protein content and oil content were relatively higher. The grain nutrition quality indexes such as four protein component contents, glutenin/gliadin ratio, amylopectin /amylose ratio, sub-oleic acid content, flax acid content, un-saturated fatty acid content and sub-oleic acid/Oleic acid ratio increased by shading, these suggesting shading improved the grain nutrition quality. However, the grain starch RVA eigenvalues such as peak viscosity (PV), hot viscosity (HV) and breakdown (BD) decreased by shading, resulting in the decline of cooking and eating quality. The activities of the tested enzymes were significantly reduced by shading treatment, in which the activities of adenosine diphosphate glucose pyrophosphorylase (ADPG), uridine diphosphate glucose pyrophosphorylase (UDPG), soluble starch synthases (SSS), and granule bounded starch synthases (GBSS) were significantly and positively correlated with grain starch content, test weight, peak viscosity, and breakdown. The activities of glutamine synthase (GS) and glutamate synthase (GOGAT) were significantly and positively correlated with grain protein content and Glu/Gli ratio. Among three shading treatments, the commodity quality (test weight and water content) of maize grain was affected mostly by S1, moderately by S2, and least by S3, respectively. However, the effect of S2 treatment on grain nutrition quality was more decisive as compared with those of S1 and S3. This result suggested that the early and middle terms from flowering to maturity are the key stage for the impact of shading on grain qualities, with little effect for the light condition at the later stage from flowering to maturity stage.

选用高淀粉玉米费玉3号和普通玉米泰玉2号，大田条件下分别于花粒前期（授粉后1~14 d），中期（15~28 d）和后期（29~42 d）遮光（透光率45%）处理，研究了对籽粒品质和籽粒中蛋白质、淀粉合成关键酶活性的影响，以及影响品质的关键时期。结果表明，花粒期遮光显著降低了籽粒产量、容重、淀粉直/支比、淀粉及其组分含量，遮光后淀粉RVA谱特征值峰值黏度（PV）、热浆黏度（HV）及崩解值（BD）下降，而籽粒含水率、粗蛋白及其各组分含量、谷/醇比、粗脂肪含量、亚油酸含量、亚麻酸含量、亚油酸/亚麻酸比值以及不饱和脂肪酸含量均升高；遮光显著降低了籽粒蛋白质和淀粉合成关键酶活性，腺苷二磷酸葡萄糖焦磷酸化酶（ADPGPPase）、尿苷二磷酸葡萄糖焦磷酸化酶（UDPGPPase）、淀粉合酶（SSS）、束缚态淀粉合酶（GBSS）活性与籽粒淀粉含量、容重、峰值黏度和崩解值呈显著正相关，谷氨酰胺合酶（GS）和谷氨酸合酶（GOGAT）活性与蛋白质含量和谷/醇比也呈显著正相关。不同时期遮光对两品种玉米品质性状的影响基本一致，前期遮光处理对籽粒商品品质(容重，含水率)影响大，而中期遮光对籽粒营养品质影响大。可见花粒中前期的光照强度对籽粒品质形成具有关键作用，后期光强对其影响较小。

2005

0.0

任国玉, 郭军, 徐铭志, 初子莹, 张莉, 邹旭恺, 李庆祥, 刘小宁. 近50年中国地面气候变化基本特征. 气象学报, 2005, 63: 942-956

Ren G

, Guo

, Xu M

, Chu Z

, Zhang

, Zou X

, Li Q

, Liu X

. Climate changes of Chinese mainland over the past half century. Acta Meteorol Sin, 2005, 63: 942-956 (in Chinese with English abstract)

A data set of mean temperature, precipitation and other climate parameters for 740 stations across China′s Mainland was used to analyze climate change in mainland China. The data set has been processed more carefully, and the main in homogeneities existing in the monthly mean temperature data have been checked and corrected. The widely accepted procedures for creating area-averaged climatic time series and for calculating linear trend have been used. Analyses have been made for annual and monthly mean temperature, precipitation, pan evaporation, sunshine duration, wind speed, and the maximum depth of snow cover.The analyses show that annual mean surface air temperature in China′s Mainland as a whole rose by about 1.1 ℃ for the last 50 years, with a warming rate of about 0.22 ℃/10 a. The warming in the later half 20th century is more rapid than the average values of the world and the Northern Hemisphere. The most evident warming occurred in winter and spring as expected. Northeast China, North China and Northwest China experienced more significant warming in terms of annual mean temperature, while a cooling trend reported in earlier studies is still continuing. Summer mean temperature in the middle and lower reaches of the Yangtze River also decreased in the last 50 years.No significant long term change in the country averaged annual precipitation was seen for both the past 50 years. However, an obvious tendency of drying in the Yellow River Basin and the North China Plain in terms of precipitation has been found, and the largest drop in precipitation occurred in Shandong Province. Meanwhile, an insignificant wetting trend in the Yangtze River Basin and most parts of western China could be detectable. For the Yangtze River Basin, the increased annual precipitation mainly resulted from the significant rising of summer rainfall, though winter precipitation also tended to increase.Since 1956, the country veraged pan evaporation has had a significant trend to decrease, with a changing rate of -34.5 mm/10a. The most significant decreaseoccurred in spring and summer in the North China Plain and the lower reach of the Yangtze River. The largest decrease in Pan evaporation in terms of absolute values is in northwestern China. It is worth to note that average sunshine duration and wind speed are also experiencing a tremendous drop in most regions of the country during the past 50 years, and they usually bear a similarity with pan evaporation in spatial pattern of the tendencies. In parts of the North China Plain,annual sunshine duration in the recent years is almost 500 hours fewer than that of 50 years ago. We have also tentatively analyzed the possible causes for the observed changes. However, it is be premature at this moment to give a definite conclusion.

采用国家基准气候站和基本气象站的地面资料，系统地分析了中国大陆地区1951年以来近地表主要气候要素演化的时间和空间特征。结果表明，中国近50 a来年平均地表气温变暖幅度约为1.1 ℃，增温速率接近0.22 ℃/（10 a），比全球或半球同期平均增温速率明显偏高。地表气温增暖主要发生在最近的20余年，其季节和空间特征与前人分析结论基本一致。降水量变化趋势对所取时间段和区域范围敏感。1951年以来全国平均降水量变化趋势不明显，但1956年以来略有增加。降水变化的空间特征明显而相对稳定，东北北部、包括长江中下游的东南部地区和西部广大地区降水增加，而华北地区以及东北东南部和西北东部地区降水明显减少。分析还发现，近50 a来全国平均的日照时数、平均风速、水面蒸发等气候要素均呈显著下降趋势，但积雪地带的最大积雪深度却有所增加。中国日照时间和水面蒸发量变化的空间特征很相似，减少最明显的地区均发生在华北和华东，新疆次之。影响中国年代以上尺度气候变化的因子错综复杂，人类活动引起的大气中温室气体浓度增高可能在一定程度上影响了中国近50 a来的气候，但考虑到尚存的不确定性，目前仍不能给出明确结论。中国东部大部分地区日照时间和水面蒸发量减少可能均起源于人为排放的气溶胶影响，平均风速减弱也有利于水面蒸发量下降，而在西部地区云量和降水量的变化可能更重要。

2004

0.0

杨彦武, 于强, 王靖. 近40年华北及华东局部主要气候资源要素的时空变异性. 资源科学, 2004, 26(4): 45-50

Yang Y

, Yu

, Wang

. Spatio-temporal variations of principal climatic factors in north china and part of east China within past 40 years. Resour Sci, 2004, 26(4): 45-50 (in Chinese with English abstract)

该文利用时空数据模型引入时间维,在GIS平台上分析了我国华北地区和华东部分地区1961年～2000年温度、降水和日照时数的时空变化规律,以及三者的相互联系.结果表明,华北地区北部温度变化不大,而南部和华东部分地区温度升高明显,10年平均气温大约升高0.5℃/40年;华北地区降水呈东南高西北低的特点,并有明显的年代际变化,华北南部和北部的降水变化存在一定的反相关关系,华北北部降水减少,而东南部降水却呈上升势头;华北地区和华东部分地区的日照时数都有明显的降低.对三者的关系进行了定性分析,华北中北部温度和降水的关系为暖干型,而南部和华东地区为暖湿型;华北中北部日照时数的降低主要是因为大气污染加剧而引起,而华北东南部和华东部分地区是由于降水的增加,即云量的增加和晴天日数的减少而导致日照时数的降低.对温度、降水和日照时数时空变化规律的分析有助于对未来气候变化做出合理的预测,并制定相应的对策.

2012

0.0

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: 3497-3505

Cui H

, Jin L

, Li

, Zhang J

, Zhao

, Dong S

, Liu

. Effects of shading on stalks morphology, structure and lodging of summer maize in field. Sci Agric Sin, 2012, 45: 3497-3505 (in Chinese with English abstract)

... 1 田间小气候参照崔海岩等^[28]方法测定光照强度、气温、CO₂浓度、相对湿度、风速和地温(5 cm) ...

1996

0.0

张祖建, 朱庆森, 王志琴. 水稻胚乳细胞计数方法研究. 江苏农学院学报, 1996, 17(6): 7-11

Zhang Z

, Zhu Q

, Wang Z

. A method of counting endosperm cells in rice grain. Jiangsu Agric Res, 1996, 17(6): 7-11 (in Chinese with English abstract)

摘　要：在改进Ｓｉｎｇｈ小麦胚乳细胞计数方法的基础上，研究提出水稻单个籽粒胚乳细胞的计数方法，进行了细胞计数过程中各环节的误差分析，并对计数方法鉴别样品差异的能力做出统计推断，证明是一种可用于较大样本量、简便易行的水稻胚乳细胞计数方法。

... 参照张祖建等^[29]方法测定玉米籽粒胚乳细胞数, 用Logistic方程拟合籽粒胚乳细胞增殖动态, 导出增殖特征参数 ...

1985

0.0

... 4 籽粒淀粉和粗脂肪含量采用双波长法测定淀粉含量及组分^[30] ...

... 采用残余油重法测定粗脂肪含量^[30] ...

2015

0.0

... 1 不同光照对夏玉米产量和干物质积累量的影响光合作用决定了玉米90%以上的干物质, 且干物质的积累、分配与产量显著正相关^[31,32,33], 尤其是生育后期的干物质积累分配特性决定了夏玉米的最终产量^[34] ...

... Cui等^[31]研究指出, 雌穗小花分化对光照非常敏感, 光照不足降低小花分化总数, 降低穗粒数, 导致产量降低 ...

2010

0.0

宋凤斌, 童淑媛. 不同株型玉米的干物质积累、分配及转运特征. 江苏农业学报, 2010, 26: 700-705

Song F

, Tong S

. Characteristics of dry matter accumulation, distribution and translocation in maize with different plant types. Jiangsu J Agric Sci, 2010, 26: 700-705 (in Chinese with English abstract)

干物质积累及转运是影响玉米产量的重要因素,明确不同部位营养器官干物质对籽粒产量的贡献,对品种改良具有重要意义.本研究以平展型玉米农大364和紧凑型玉米郑单958为材料,探讨玉米不同部位叶片、叶鞘和茎秆的干物质积累、转运及其对籽粒的贡献.结果表明:玉米茎秆干物质的转运对籽粒的贡献最大,随着节位的升高贡献率降低;农大364叶鞘的干物质转运率高于叶片,郑单958则相反.通过提高农大364穗位叶以上叶片、第9～第11叶鞘和第15节以上节间的干物质转运率,以及郑单958穗位叶以下叶片、第12叶以上叶鞘和第16节以下节间的干物质转运率,可能在一定程度上会提高产量.

2008

0.0

戴明宏, 陶洪斌, 王利纳, 王璞. 不同氮肥管理对春玉米干物质生产、分配及转运的影响. 华北农学报, 2008, 23(1): 154-157

Dai M

, Tao H

, Wang L

, Wang

. Effects of different nitrogen managements on dry matter accumulation, partition and transportation of spring maize ( Zea mays L. ). Acta Agric Boreali-Sin, 2008, 23(1): 154-157 (in Chinese with English abstract)

The objective of this study is to provide a technique for high2yielding spring maize cultivation by analyzingeffects of different nitrogen managements on dry matter accumulation,partition and transportation. There are three nitrogenmanagements:Zero nitrogen fertilization(ZNF),Recommended nitrogen fertilization(RNF),Traditional nitrogen fertiliza2tion(TNF). The results showed that neither RNF nor TNF could obviously improve dry matter accumulation,LAI,ear2leave photosynthetic rate and grain yield in the first year. However,in the second year,ZNF achieved yield reduction of1210 % and 1116 % comparing with RNF and TNF respectively. Furthermore,RNF could not only reduce nitrogen input,but also improve dry matter transportation from other organs to grain,the proportion of which reached 2211 %,and was611 % higher than TNF.

为探索华北地区春玉米高产可持续栽培技术,研究了不同氮素管理(不施氮、推荐施氮、经验施氮)对春玉米的干物质积累、分配及转运的影响。结果表明,在高肥力土壤条件下,第一年推荐和经验施氮同不施氮相比在干物质积累、叶面积指数、籽粒产量、穗位叶光合速率等方面都没有起到明显的促进作用,但在第二年不施氮处理产量比推荐施氮和经验施氮分别下降了12.0%和11.6%。推荐施氮的优势不仅体现在减少氮肥投入的前提下保持产量的稳定,同时也明显促进了生育后期植株营养体干物质向籽粒的转运,各器官干物质转运总量占籽粒总干质量的22.1%,比经验施氮高6.1%。

2011

0.0

刘伟, 张吉旺, 吕鹏, 杨今胜, 刘鹏, 董树亭, 李登海, 孙庆泉. 种植密度对高产夏玉米登海661产量及干物质积累与分配的影响. 作物学报, 2011, 37: 1301-1307

Liu

, Zhang J

, Lyu

, Yang J

, Liu

, Dong S

, Li D

, Sun Q

. Effect of plant density on grain yield dry matter accumulation and partitioning in summer maize cultivar Denghai 661. Acta Agron Sin, 2011, 37: 1301-1307 (in Chinese with English abstract)

2010

0.0

孙小玲, 许岳飞, 马鲁沂, 周禾. 植株叶片的光合色素构成对遮阴的响应. 植物生态学报, 2010, 34: 989-999

Sun X

, Xu Y

, Ma L

, Zhou

. A review of acclimation of photosynthetic pigment composition in plant leaves to shade environment. Chin J Plant Ecol, 2010, 34: 989-999 (in Chinese with English abstract)

Chlorophylls function in harvesting light energy, funneling the excitation to reaction center and converting sunlight into chemical energy, and carotenoids are responsible for light harvesting and photoprotection. Both are vital for photosynthesis. We summarized the distribution and function of the main photosynthetic pigments and variation of pigment composition and content in sun and shade plants. Sun plants possess larger xanthophyll cycle pool size (violaxanthin + antheraxanthin + zeaxanthin), but de-epoxidation level is lower than that of shade plants. The ratio of lutein to xanthophyll cycle pool size is positively correlated to plant shade tolerance. Light intensity and spectral quality vary between different shade sources. Generally for plant growth, building shade is better than vegetation shade, and deciduous shade exceeds coniferous shade. Variation in light intensity may activate two cycles in plants, xanthophyll cycle and lutein epoxide cycle, for light harvesting or energy dissipation. Some species may alter chlorophyll content and Chl a/b ratio to acclimate to different light intensity, but this character is not related to their shade tolerance. Temporary shade is not necessarily detrimental. Xanthophyll cycle pool size is not only determined by daily photon receipt, but also by the way photon flux is distributed over the daylight hours, because light and temperature are both essential for optimal photosynthetic metabolism. The best photosynthetic performances of plants were obtained with the reinforcement of blue, red and far red wavelengths and with a red: far red ratio closer to that observed in nature. We reviewed internal and external factors affecting photosynthetic pigment content and composition, and determined that during the acclimation to different light environments, plants altered pigment composition and content mainly through adjusting the ratio of reaction center to light harvesting complex and PSI/PSII. We also discussed current research problems and provided insight into future relevant research.

叶绿素在植株体内负责光能的吸收、传递和转化, 类胡萝卜素则行使光能捕获和光破坏防御两大功能, 它们在光合作用中起着非常重要的作用。该文综述了几大主要光合色素的分布和功能, 以及不同物种的色素含量和构成差异。阳生植物的叶黄素库较大, 但脱环氧化水平不及阴生植物。黄体素与叶黄素库的比值与植物的耐阴性呈正相关关系。由不同的遮阴源造成的遮阴环境, 光强和光质有很大的差异, 总体来说对植物生长的影响, 建筑物遮阴<阔叶林遮阴<针叶林遮阴。光强的改变可诱导类胡萝卜素的两大循环——叶黄素循环和黄体素循环。由光强诱导的叶绿素含量和叶绿素a/b比值的改变与该物种的耐阴性无关。短时间的遮阴不会对植物的生长造成危害, 叶黄素库的大小不仅与每天接受的光量子有关, 更与光量子在一天的分布有关, 因为光照和温度是协同作用的。光合作用或色素构成是蓝光、红光和远红光共同作用的结果, 不是某一种单色光所能替代的。我们总结了影响植物色素构成的内因和外因, 指出植物主要通过调整光反应中心和捕光天线色素蛋白复合体的比例, 以及两个光系统的比值来调整色素含量和构成以适应不同的光照条件, 提出了现存研究中存在的一些问题, 旨在为今后的相关研究提供建议。

... 前人研究表明, 光照不足, 尤其是生育后期光照不足严重制约玉米生产, 表现为叶片光合性能下降, 籽粒灌浆强度降低, 干物质向籽粒中的转运量下降, 直接影响夏玉米产量^[35,36] ...

2005

0.0

李潮海, 栾丽敏, 尹飞, 王群, 赵亚丽. 弱光胁迫对不同基因型玉米生长发育和产量的影响. 生态学报, 2005, 25: 824-830

Li C

, Luan L

, Yin

, Wang

, Zhao Y

. Effects of seedling shading and light intensity transfer on photosynthetic efficiency of different maize hybrids. Acta Ecol Sin, 2005, 31: 381-385 (in Chinese with English abstract)

Light is one of the most important factors affecting corn (Zea mays L.) production. The light saturation point of corn is much higher than that of most other crops, allowing corn to take advantage of the higher light intensity. During the growing season, however, corn frequently encounters low light intensities that restrict growth capacity, therefore limiting the grain yield. The level of photosynthetic active radiation (PAR) can markedly change the growth environment, which directly impacts photosynthesis...

以不同基因型玉米为材料,在玉米生长发育的3个主要阶段(苗期、穗期、粒期)进行分期遮光试验,研究不同时期弱光胁迫对不同基因型玉米生长发育和产量的影响。结果表明,遮光延缓了玉米叶片的出生速度,使叶片变薄;遮光可以延缓叶片的衰老,但遮光解除后则加速叶片的衰老;遮光造成植株高度增加,但恢复正常光照后,其株高却逐渐低于对照;遮光使干物质积累下降,抽雄吐丝日期推迟,尤其是吐丝日期推迟更多,并使产量降低,但不同基因型玉米不同遮光处理下降程度不同。试验的4个品种中,掖单2 2和豫玉2号受遮光影响较小,而掖单36 38和丹玉13受影响较大,即不同基因型玉米对弱光胁迫的敏感性不同。

2013

0.0

崔海岩, 靳立斌, 李波, 董树亭, 刘鹏, 赵斌, 张吉旺. 遮阴对夏玉米干物质积累及养分吸收的影响. 应用生态学报, 2013, 24: 3099-3105

Cui H

, Jin L

, Li

, Dong S

, Liu

, Zhao

, Zhang J

. Effects of shading on dry matter accumulation and nutrient absorption of summer maize. Chin J Appl Ecol, 2013, 24: 3099-3105 (in Chinese with English abstract)

... S2, 说明夏玉米花粒期对光照更为敏感, 花粒期充足的光照更有利于夏玉米高产^[37,38,39] ...

... 相反, 光照不足导致玉米光合速率下降, 光合产物向籽粒中的转运量降低^[37] ...

2010

0.0

贾士芳, 董树亭, 王空军, 张吉旺, 刘鹏. 弱光胁迫影响夏玉米光合效率的生理机制初探. 植物生态学报, 2010, 34: 1439-1447

Jia S

, Dong S

, Wang K

, Zhang J

, Liu

. Physiological mechanism of shading stress on photosynthetic efficiency in summer maize ( Zea mays L. ). Chin J Plant Ecol, 2010, 34: 1439-1447 (in Chinese with English abstract)

... S2, 说明夏玉米花粒期对光照更为敏感, 花粒期充足的光照更有利于夏玉米高产^[37,38,39] ...

2013

0.0

... S2, 说明夏玉米花粒期对光照更为敏感, 花粒期充足的光照更有利于夏玉米高产^[37,38,39] ...

... 增光处理一方面有利于玉米积累更多的光合产物, 提高干物质量^[39], 达到增#cod#x0201c ...

2016

0.0

... 的潜力, 增库则需要提高灌浆速率, 提前最大灌浆速率出现的日期, 保证光合产物高效地运转到籽粒^[40] ...