作物杂志, 2019, 35(2): 136-141 doi: 10.16035/j.issn.1001-7283.2019.02.021

生理生化·植物营养·栽培耕作

叶面喷施5-氨基乙酰丙酸对不同密度春玉米生长特性和产量的影响

唐会会, 许艳丽, 王庆燕, 马正波, 李光彦, 董会, 董志强

中国农业科学院作物科学研究所,100081,北京

Effects of Foliar Spraying 5-Aminolevulinic Acid on Spring Maize Growth and Yield under Different Planting Densities

Tang Huihui, Xu Yanli, Wang Qingyan, Ma Zhengbo, Li Guangyan, Dong Hui, Dong Zhiqiang

Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China

通讯作者: 董志强,研究员,研究方向为作物栽培生理与化学调控

第一联系人:

唐会会,硕士研究生,研究方向为作物栽培生理与化学调控

收稿日期: 2018-10-21   修回日期: 2019-02-26   网络出版日期: 2019-04-15

基金资助: 国家重点研发计划-新型高效植物生长调节剂和生物除草剂研发项目.  2017YFD0201301
国家自然科学基金面上项目.  31470087

Received: 2018-10-21   Revised: 2019-02-26   Online: 2019-04-15

摘要

为探讨5-氨基乙酰丙酸(5-aminolevulinic acid,ALA)对不同密度春玉米生长发育和产量的影响,以中单909为材料,在玉米9展叶时期叶面喷施100mg/L ALA,以喷施等量清水为对照,在大田条件下研究ALA对不同密度春玉米群体(45 000、75 000、105 000株/hm 2)的作用效果。结果表明:ALA处理下,玉米45 000和75 000株/hm 2密度群体产量分别比对照增加9.7%和4.9%,75 000株/hm 2密度群体千粒重和收获期单株干物质积累量分别增加3.7%和4.3%,花后叶面积指数平均增加6.8%,SPAD值比对照分别平均增加2.5%,净光合速率平均比对照高15.7%。综上所述,ALA处理改善了大田玉米的群体结构,延缓了叶片衰老进程,提高了地上部分干物质积累量,进而提高了玉米单产水平,因此可作为东北春玉米高产稳产的技术措施。

关键词: 5-氨基乙酰丙酸 ; 春玉米 ; 密度 ; 群体结构 ; 产量

Abstract

To explore the effects of 5-aminolevulinic acid (ALA) on spring maize growth and yield and its physiological mechanisms, a split-spot experiment was conducted in Gongzhuling experiment station, Jilin Province. Maize hybrid Zhongdan 909 was used with low (45 000 plant/hm 2), middle (75 000 plant/hm 2) and high (105 000 plant/hm 2) planting population. The results showed that the yield increased by 9.7% and 4.9% in the densities of 45 000 and 75 000 plant/hm 2 compared with the control ALA treatments, respectively. Under the density of 75 000 plant/hm 2, ALA treatment increased maize thousand kernel weight, dry matter accumulation at harvest, leaf area index after anthesis, SPAD value and net photosynthetic rate by 3.7%, 4.3%, 6.8%, 2.5% and 15.7%, respectively, compared with that of control treatment under the same plant density. In conclusion, ALA improved canopy structure of maize plant in field conditions, delayed leaf senescence process, increased aboveground dry matter accumulation, which in turn, increased maize yield. Therefore, ALA could be used as a technical measure for high and stable yield of spring maize in Northeast China.

Keywords: ALA ; Spring maize ; Density ; Population structure ; Yield

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本文引用格式

唐会会, 许艳丽, 王庆燕, 马正波, 李光彦, 董会, 董志强. 叶面喷施5-氨基乙酰丙酸对不同密度春玉米生长特性和产量的影响[J]. 作物杂志, 2019, 35(2): 136-141 doi:10.16035/j.issn.1001-7283.2019.02.021

Tang Huihui, Xu Yanli, Wang Qingyan, Ma Zhengbo, Li Guangyan, Dong Hui, Dong Zhiqiang. Effects of Foliar Spraying 5-Aminolevulinic Acid on Spring Maize Growth and Yield under Different Planting Densities[J]. Crops, 2019, 35(2): 136-141 doi:10.16035/j.issn.1001-7283.2019.02.021

合理密植是玉米(Zea mays L.)高产栽培最重要的技术措施之一,而密植使玉米群体冠层通透性变差、叶片光合效率降低,进而容易导致干物质生产量下降、群体空秆增多、雌穗秃尖增长、茎秆细软和根系分布浅、倒伏等,这些因素均严重制约玉米的增产潜力[1,2]。因此,提高密植条件下的冠层叶片光合效率,改善干物质生产能力,对挖掘玉米高产潜力具有重要意义。

前人研究[3,4,5,6,7,8,9,10]表明,叶面喷施5-氨基乙酰丙酸(5-aminolevulinic acid,ALA)可有效提高作物光合速率。ALA又称5-氨基酮戊酸,是卟啉类化合物生物合成的关键前体,在绿色植物中,它是植物叶绿素合成的前体[11]。ALA具有调节植物生长发育的功能,是一种新的植物生长调节物质[12]。高浓度的外源ALA具有光敏除草剂的作用;低浓度的ALA可调节植物生长发育,促进作物生长,增强抗逆性,提高作物产量。因而在农业生产中ALA被认为有重要的潜在应用价值[13]。叶面喷施ALA能够提高光合速率、促进光合作用[3,4,5,6,7,8,9,10]、增强抗逆性[3-4,14-15],促进植物叶绿素合成与稳定,提高叶绿素含量[4-6,16]。低浓度ALA能够促进玉米幼苗[7]、西瓜[3]、西葫芦[5]、油菜[14]、辣椒[15,17]和水稻[18]植株的生长及干物质积累,增加水稻[18,19]、番茄[20]、西葫芦[21]、番茄和辣椒[22]的产量。截至目前,ALA在农业上的研究大多集中在蔬菜、瓜果等经济作物上,对大田作物,如玉米,只有少量盆栽[3]和浸种[23]的研究,将ALA运用于大田试验的研究尚未见报道。因此,本研究通过大田试验,研究ALA对不同密度群体玉米生长发育特性和产量的影响,探讨大田条件下ALA在玉米上的应用效果,为建立密植群体高产稳产栽培技术提供理论与技术支撑。

1 材料与方法

1.1 试验地概况

本试验于2016、2017年在中国农业科学院吉林省公主岭试验站(43°29′55″N,124 °48′43″E)进行。试验地土壤为黑土,土壤耕层(0~20cm)有机质含量26.7g/kg、全氮含量1.4g/kg、速效氮含量155.3mg/kg,速效磷含量34.4mg/kg、速效钾含量184.2mg/kg,pH 5.8。

1.2 试验材料与设计

试验品种为中单909(中国农业科学院作物科学研究所选育)。采用裂区设计,ALA喷施为主区,密度为副区。主区在玉米9展叶时期叶面喷施100mg/L ALA,对照(CK)喷施等量清水;副区设3个密度处理,即45 000、75 000、105 000株/hm2,不同密度处理和对照分别用TR4.5、TR7.5、TR10.5和CK4.5、CK7.5、CK10.5表示。小区为8行区,60cm等行距播种。小区长6m,宽4.8m,小区面积28.8m2,每个处理4次重复。全生育期施肥量N∶P2O5∶K2O=247.5∶97.5∶120kg/hm2;田间管理同当地大田生产。2016年4月30日播种,5月14日出苗,9月29日收获;2017年4月27日播种,5月21日出苗,9月27日收获。

1.3 测定项目与方法

1.3.1 产量及产量构成因素 玉米成熟后,在小区中部选10m2称重测产,调查穗部性状,测定出子率和含水量,并折算产量(按14%含水量计)。

1.3.2 地上部分干物质积累量及叶面积指数 分别在拔节期(V6)、大口期(V12)、吐丝期(VT)、花后15d(VT+15)、花后30d(VT+30)、花后45d(VT+45)和收获期(R6)选取有代表性的植株3株,按部位(叶片、叶鞘、茎秆、苞叶、子粒、穗轴)分开,于105℃杀青30min后于85℃烘干至恒重,称量并计算地上部分干物质积累量。同时测量每片叶片的长宽。采用长宽系数法计算叶面积,叶面积指数(LAI)=该土地面积上的总叶面积/土地面积。

1.3.3 株高及穗位高 在吐丝期(VT)、花后15d(VT+15)、花后30d(VT+30)、花后45d(VT+45)和收获期(R6)选取有代表性的植株3株,测株高及穗位高。

1.3.4 叶绿素相对含量(SPAD值) 在大口期(V12)、吐丝期(VT)、花后20d(VT+20)和花后40d(VT+40)用手持式SPAD-502型叶绿素计测定穗位叶叶绿素相对含量。

1.3.5 单粒重测定 玉米吐丝后,选长势、穗型一致的植株进行挂牌标记。从吐丝第10天起开始取样,每隔10d在小区标记植株中随机取3个果穗,每穗取中部子粒100粒,于105℃烘箱中杀青30min,85℃烘至恒重后称重。利用Curve Expert 1.3进行Logistic模拟子粒单粒重变化。

1.3.6 光合速率测定 在花后第15天,用Li-6400便携式光合测定仪测定穗位叶净光合速率(Pn)。

1.4 试验数据处理

采用Microsoft Excel 2007进行数据整理计算及作图,采用SPSS 18.0进行统计分析。除子粒灌浆特征曲线和净光合速率选用2016年数据外,其余均采用2016和2017年的数据。

2 结果与分析

2.1 玉米产量及其构成因素

表1可知,化学调控处理比对照处理提高了春玉米产量,2016和2017年ALA处理春玉米产量相比对照平均分别增加6.0%(731.7kg/hm2)和1.7%(202.3kg/hm2)。2016年,TR4.5、TR7.5和TR10.5处理相比CK4.5、CK7.5和CK10.5处理春玉米产量分别增加15.0%(1 788.0kg/hm2)、8.4%(1 082.4kg/hm2)和-5.8%(-675.1kg/hm2);2017年分别增加4.3%(528.6kg/hm2)、1.4%(173.3kg/hm2)和-0.8%(-95.1kg/hm2)。综上,ALA处理下TR4.5和TR7.5的春玉米产量分别比CK4.5和CK7.5平均增加9.7%和4.9%。

表1   2016-2017年ALA对不同密度春玉米产量及其构成因素的影响

Table 1  Effects of ALA on maize yield components of three planting populations in 2016 and 2017

年份
Year
处理
Treatment
穗长(cm)
Ear length
秃尖长(cm)
Bare tip length
穗粗(mm)
Ear diameter
穗粒数
Kernel number per ear
千粒重(g)
Thousand kernel weight
产量(kg/hm2)
Yield
2016CK4.519.8±0.60.5±0.654.5±0.7623.9±53.2371.9±5.911 888.4±838.5
TR4.520.1±0.10.3±0.554.4±0.8627.8±65.4348.2±9.013 676.3±244.5**
CK7.517.5±0.80.9±0.351.6±0.6582.0±58.2309.0±10.212 848.3±603.0
TR7.517.6±0.60.7±0.452.2±0.5578.3±73.8318.2±9.013 930.8±116.0
CK10.516.2±0.60.6±0.448.5±0.7503.3±42.5277.8±13.011 691.3±1 380.0
TR10.515.6±1.10.9±0.649.7±0.4*475.6±51.8300.6±6.6**11 016.2±684.9
2017CK4.522.0±0.80.5±0.751.6±3.2689.9±75.8303.0±20.912 395.5±554.4
TR4.521.8±0.90.5±0.652.1±1.4702.4±49.8330.0±6.9*12 923.7±594.0*
CK7.519.9±0.80.9±0.648.9±1.6638.6±74.9266.3±8.112 589.3±710.1
TR7.518.7±1.51.2±0.748.5±1.7555.3±80.6277.9±16.912 762.5±648.9
CK10.518.2±0.81.6±0.846.4±2.3539.8±61.4250.2±13.211 618.7±461.6
TR10.518.0±1.01.6±0.545.9±1.6530.4±69.4248.7±27.611 523.6±1 113.2

Note: "*" and "**" mean significant difference between ALA and control treatment at P=0.05 and P=0.01 level under the same planting density, respectively

注:“*”和“**”表示在同一播种密度ALA处理与对照在P=0.05和P=0.01水平上差异显著

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表1所示,两年试验春玉米雌穗穗长、穗粗、穗粒数和千粒重随播种密度的增加均呈下降趋势。ALA处理下,2016年TR4.5和TR7.5相比CK4.5和CK7.5,秃尖长分别缩短40.0%和22.2%,而TR4.5穗粒数增加0.6%,TR7.5和TR10.5千粒重分别增加3.0%和8.2%;2017年TR4.5和TR7.5的千粒重相比CK4.5和CK7.5分别增加8.9%和4.4%。综上,ALA处理下TR7.5的千粒重比CK7.5平均增加3.7%。

2.2 地上部分干物质积累动态

图1所示,2016年ALA处理提高了各密度花后干物质积累,其中,TR4.5在VT+30和VT+45时干物质积累量均分别较CK4.5高12.3%和7.7%;TR7.5在R6时高于CK7.5,增幅为4.3%;TR10.5在VT+30、VT+45和R6分别比相应对照高11.7%、5.6%和16.3%。2017年TR4.5在R6时期干物质积累比CK4.5高7.3%,TR7.5在VT+15、VT+45和R6时期分别比CK7.5高5.7%、1.7%和1.2%。

图1

图1   ALA对不同密度玉米干物质积累的影响(2016-2017年)

Fig.1   Effects of ALA on maize dry matter accumulation under different planting densities in 2016 and 2017


2.3 叶面积指数变化动态

随密度增加叶面积指数呈增加趋势,而不同密度群体叶面积指数对ALA处理响应不一致(图2)。2016年,播种密度为45 000株/hm2时,V6至VT+15期间,CK4.5和TR4.5叶面积指数无明显差异,而在VT+15至VT+45,TR4.5比CK4.5叶面积指数增加7.4%;播种密度为75 000株/hm2时,V6至VT期间,TR7.5叶面积指数比CK7.5降低4.1%,VT至VT+30无明显差异,VT+30至VT+60 TR7.5叶面积指数比CK7.5平均高27.2%;播种密度为105 000株/hm2时,ALA处理叶面积指数呈增加趋势,在VT+15、VT+30和VT+60 TR10.5叶面积指数分别比CK10.5增加1.0%、10.7%和37.0%。ALA处理叶面积指数在VT+15、VT+30、VT+45和VT+60分别平均比对照增加0.7%、5.3%、3.1%和18.1%,花后叶面积指数平均增加6.8%。2017年,TR4.5叶面积指数在VT+45和R6时期较CK4.5分别高4.4%和27.6%;TR7.5在VT+30、VT+45和R6时期较CK4.5分别高5.0%、7.3%和31.0%;TR10.5在VT+30、VT+45和R6时期较CK4.5分别高9.5%、9.5%和78.4%。

图2

图2   ALA对不同密度玉米LAI的影响(2016-2017年)

Fig.2   Effects of ALA on maize LAI under different planting densities in 2016 and 2017


2.4 玉米穗位叶SPAD值变化

表2所示,2016年玉米穗位叶SPAD值随播种密度增加呈下降趋势,从V12至VT+40呈现先增加后下降的趋势。2016年,ALA处理增加各密度梯度SPAD值,V12、VT和VT+40时各密度SPAD值比对照分别平均增加3.3%、2.9%和1.4%,平均增加2.5%。2017年,TR10.5在V12、VT、VT+20和VT+40时期SPAD值分别比CK10.5高1.6%、7.7%、2.5%和5.7%。

表2   ALA对不同密度春玉米SPAD值的影响(2016-2017年)

Table 2  Effects of ALA on maize ear leaf SPAD value of three planting densities in 2016 and 2017

年份处理
Treatment
生育时期Growth stage
V12VTVT+20VT+40
2016CK4.559.4±1.262.4±2.563.5±2.261.8±2.4
TR4.560.2±0.662.5±0.464.0±2.563.3±1.3
CK7.558.4±0.661.3±2.263.0±2.358.9±3.7
TR7.559.8±0.861.7±0.860.3±1.258.4±1.9
CK10.557.0±0.954.9±1.361.3±0.654.4±3.0
TR10.560.5±0.559.6±1.159.4±1.555.8±3.7
2017CK4.558.2±4.263.5±2.467.4±3.460.9±1.8
TR4.562.5±1.659.9±3.365.0±3.759.8±2.5
CK7.554.4±3.660.4±1.561.6±4.557.1±5.9
TR7.554.6±5.056.4±5.160.6±1.760.7±2.7
CK10.551.2±4.553.0±3.355.7±3.849.2±3.7
TR10.552.0±5.757.1±3.556.9±3.852.0±2.5

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2.5 玉米子粒灌浆特征

单粒重与灌浆天数模拟结果(图3)显示,播种密度为45 000株/hm2时,灌浆10~30d,TR4.5单粒重高于CK4.5,灌浆50~60d相反;播种密度为75 000株/hm2时灌浆50d和60d子粒单粒重较CK7.5分别低3.6%和4.4%,播种密度为105 000株/hm2时灌浆50d和60d子粒单粒重较CK7.5分别高6.6%和11.5%。

图3

图3   ALA对不同密度玉米子粒灌浆特征的影响(2016年)

Fig.3   Effects of ALA on maize grain-filling of three plant densities in 2016


2.6 穗位叶Pn

2016年,通过测定玉米抽雄后15d穗位叶Pn,发现ALA处理显著增加了玉米穗位叶Pn,其中随密度增加Pn呈降低趋势,ALA处理显著增加各密度群体春玉米的Pn,其中TR4.5、TR7.5和TR10.5处理相比CK4.5、CK7.5和CK10.5处理春玉米Pn分别增加17.7%、7.0%和22.9%,平均比对照高15.7%(图4)。

图4

图4   ALA对不同密度玉米Pn的影响(2016年)

“*”表示在同一播种密度ALA处理与对照在P=0.05水平上差异显著

Fig.4   Effects of ALA on maize ear leaf net photosynthetic rate of three planting densities in 2016

"*" means significant difference between ALA and control treatment at P=0.05 level under the same planting density


3 讨论

3.1 ALA对不同密度群体春玉米干物质及产量的调控效应

ALA具有类似激素的生理作用,具有调节作物生长发育的功能。Hotta等[24]研究了低浓度ALA对多种作物生长及产量的调控效应,发现ALA能促进萝卜、大麦、马铃薯、大蒜、蚕豆等作物的干物质积累,增加产量。在研究ALA对粮食作物[7,18-19]和瓜果类作物[3,5,17,20-22]的研究中均证明ALA能促进植株生长、干物质积累和产量增加。本试验中,ALA处理不同密度群体春玉米,75 000、105 000株/hm2群体植株花后干物质积累增加,45 000和75 000株/hm2群体分别比对照增产9.7%和4.9%,表明ALA对中低密度玉米群体具有增产效应,这与前人在其他作物上的研究结果一致。

3.2 ALA对不同密度群体春玉米SPAD值的调控效应

研究发现,外源ALA处理能够提高作物叶片的叶绿素含量[25]。汪良驹等[12]研究表明,外源ALA处理明显提高甜瓜叶片叶绿素含量。在禾本科作物上的研究也表明添加了ALA的叶面肥能提高水稻功能叶[18]和玉米幼苗叶片[7]的SPAD值。本试验中,随密度增加春玉米功能叶SPAD值表现出降低趋势,其中,45 000和75 000株/hm2密度群体功能叶SPAD值显著大于105 000株/hm2,ALA处理增加了各密度群体SPAD值,该结果与要娟娟等[26]研究结果一致。但叶绿素含量与外源ALA施用量之间不存在线性关系,而是出现了类似于植物激素作用的效应。因此,有学者认为,ALA不是直接作为叶绿素合成的前体起作用,而是作为一种植物生长调节物质参与叶绿素合成,调节植物生长[23]

3.3 ALA对不同密度群体春玉米光合特性的调控效应

不同密度群体冠层中,光、热、气的通透性不同,直接影响着穗位功能叶片的光合作用效率。本试验结果表明,随密度增加,春玉米穗位功能叶Pn呈下降趋势;而ALA处理显著增加了各密度群体春玉米穗位叶的Pn。Hotta等[25]研究发现,光照条件下,ALA提高了不同作物叶片CO2的固定速率,抑制黑暗条件下叶片CO2的释放;汪良驹等[6]研究发现,ALA可促进萝卜中下部叶片的光合作用,并对叶片光系统Ⅱ光化学效率有显著的促进作用。由此可见,ALA通过诱导光抑制保护机制提高了叶片的光合效率。Tanaka等[27]以黄瓜白化苗子叶为材料,发现ALA处理诱导叶绿素a的合成,当叶绿素a转化成叶绿素b,并与LHCⅡ脱辅基蛋白结合后,形成了结构稳定的蛋白,该蛋白可以不受蛋白水解酶的影响,因而,叶片光合能力显著提高。Sasaki等[28]以螺旋藻为材料,证实ALA可以促进螺旋藻合成藻蓝蛋白,提高光系统Ⅱ活性,提高了光合效率。

4 结论

ALA处理通过延缓春玉米在灌浆中、后期的衰老速率(叶面积指数和穗位叶SPAD值增加)和穗位叶Pn的增加,促进了春玉米在灌浆中、后期地上部分干物质的积累和灌浆速率的提高,并最终增加了玉米的千粒重和产量。因此,ALA可作为东北地区春玉米高产稳产的技术措施之一加以推广。

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

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以两种不同耐密性的4个玉米品种为材料,探讨在不同密度压力下茎秆性状的变化,重点研究了地面上第三节茎秆形态及内部中央大维管束生理结构的变化,并分析了各性状与倒伏的关系。结果表明,不同类型间和同类型不同品种间对密度的反应存在明显差异。郑单958和辽单526对密度变化更加敏感。4个品种的田间倒伏率与种植密度、茎秆第三节的长粗比、茎秆中央大维管束单位视野个数呈显著正相关;与第三节茎粗、中央大维管束的平均面积呈负相关。

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以秋冬季塑料大棚中盆栽西瓜幼苗为材料,研究了50~200mg/L 5-氨基乙酰丙酸(ALA)处理对低温弱光条件下西瓜植株光合特性、碳水化合物积累、抗氧化酶活性和丙二醛(MDA)含量的影响。结果表明,外源ALA处理显著提高了西瓜幼苗叶片净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(Tr)。ALA处理还显著提高了叶片SOD和POD等抗氧化酶活性,但对CAT及APX活性影响不大,暗示ALA处理可能主要通过提高植株SOD和POD活性来促进西瓜幼苗活性氧代谢,从而保护细胞光合性能。处理后30d分析表明,ALA处理植株干物质重量比对照高出42%~54%,淀粉含量增加62.2%~207.0%,可溶性糖含量也增加32.0%~87.1%。以上结果说明,ALA处理可以增强低温弱光条件下西瓜幼苗抗氧化系统活性,促进光合作用与光合产物积累,并促进植株生长。

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The role of 5-aminolevulinic acid (5-ALA) as a precursor of chlorophyll and a herbicide is well documented. 5-ALA as a Plant Growth substance is also proven in recent times. In the present report, to elucidate the physiological effects of 5-ALA, the compound was used in in vitro studies using MS medium supplemented with 5-ALA at 2, 5 and 10 mg l -1 . Leaf and cotyledonary node were used as the explants. In vitro studies confirmed the hormonal role of 5-ALA by striking proliferation of callus paripassu induction of rooting and shooting with a profound effect of the former than the latter. Thus, 5-ALA has the dual properties of auxin and cytokinin in the induction of callusing and rhizogenesis, and shooting respectively.

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In order to utility ALA in summer squash, we need a better understanding of the mechanism of ALA to photosynthetic and physiological characteristic of summer squash. Using ‘Zaoqingyidai’ as experiment material, an experiment was conducted to examine the effects of 5-aminolevulinic acid (ALA) on photosynthetic characteristics of leaves of summer squash seedlings under chilling stress. The results were as follows: Under chilling stress,the chlorophyll contents decreased, net photosynthesis rate (Pn), leaf area and dry matter of per seedling decreased greatly. ALA of 60 mg/L could relieve the effects of chilling on summer squash seedlings by increasing chlorophyll contents, photosynthesis rate, the leaf area and dry matter with 29.96%, 52.96%, 39.80% and 43.75% respectively. Because of increasing the chlorophyll contents and decreasing nonstomatal limitation effects, ALA of 60 mg/L could relieve some of the effects of chilling stress on summer squash seedlings.

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ALA对萝卜不同叶位叶片光合作用与叶绿素荧光特性的影响

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以盆栽萝卜为材料,研究了叶面喷布100~300mg/LALA对萝卜植株生长、光合作用以及叶绿素荧光特性的影响.结果表明,ALA促进萝卜植株生长与其促进植株中下部叶片的光合作用有关.ALA处理提高了叶片光合表观量子效率,并降低了光补偿点.叶绿素荧光动力学资料显示,ALA处理不仅降低了叶绿素初始荧光Fo、稳态荧光Fs以及两者的差值ΔFo,还降低了暗适应以及光照下的最大荧光(Fm和Fm′)和可变荧光(Fv和Fv′),但是不影响PSⅡ最大光化学效率(Fv/Fm),对于中下部叶片PSⅡ实际光化学效率(ΦPSⅡ)还有明显的促进作用.ALA处理植株叶片光合非循环电子传递速率ETR显著提高,300mg/LALA处理还降低了光合相对限制值L(PFD).ALA具有促进叶绿素荧光光化学猝灭和非光化学猝灭的双重特性,其中,前者提高了叶片光化学速率,后者增加热耗散量以及提高热耗散速率.另外,本试验结果显示,ALA处理提高了植株基部叶片的叶绿素含量以及中下部叶片类萝卜素含量.以上参数变化说明ALA通过诱导光抑制保护机制来提高叶片光合效率.

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外源5-氨基乙酰丙酸对低温胁迫下玉米幼苗生长及光合特性的影响

作物杂志, 2017(5):87-93.

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为探究外源5-氨基乙酰丙酸(ALA)对低温胁迫下玉米幼苗生长及光合特性的调控效应,以绥玉13(低温敏感型)和郑单958(耐低温型)为试验材料,于三叶一心期叶面喷施20mg/L ALA,在昼/夜温度为14℃/5℃的低温条件下培养48h,恢复常温48h后取样,分析外源ALA对玉米幼苗生长、叶绿素含量、光合作用参数及叶绿素荧光特性的影响。结果表明:低温胁迫显著抑制两个玉米品种幼苗的株高、叶面积、鲜重和干重。低温处理导致幼苗SPAD值下降,叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(G_s)、胞间CO_2浓度(C_i)、最大荧光(F_m)、最大光化学效率(F_v/F_m)、潜在光化学效率(F_v/F_o)及光化学猝灭系数(q_P)水平降低,初始荧光(F_o)和非光化学猝灭系数(NPQ)升高。外源ALA能够促进低温胁迫下玉米幼苗形态建成与物质积累,叶片SPAD值增加。外源ALA能够缓解由低温胁迫导致的叶片P_n、T_r、G_s和C_i下降,提高F_m、F_v/F_m、F_v/F_o及q_P水平,降低F_o和NPQ水平。由此可见,外源ALA能够促进叶片光合作用,提高PSⅡ反应活性,促进物质积累,增强玉米苗期的抗低温能力,且对低温敏感型玉米品种的缓解低温伤害效果更好。

高晶晶, 冯新新, 段春慧 , .

ALA提高苹果叶片光合性能与果实品质的效应

果树学报, 2013,30(6):944-951.

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【目的】为了探讨5-氨基乙酰丙酸(ALA)在苹果生产上应用的可能性,【方法】以‘红星’和‘富士’苹果为材料,研究了生长季节喷施低质量浓度(0.05~5 mg·L-1)外源ALA对苹果叶片叶绿素相对含量(SPAD)、气体交换参数、叶绿素荧光快速诱导曲线(OJIP)、820 nm光吸收曲线以及果实品质的影响。【结果】0.05~5 mg·L-1ALA均能不同程度地提高苹果叶片SPAD,增加叶片净光合速率(P n)、瞬时羧化效率(P n/Ci)。M-PEA测定结果表明,ALA处理叶片单位面积吸收(ABS/CS)、捕获(TR0/CS)和传递(ET0/CS)的能量以及光合性能指数(PI ABS和PI CS)均有不同程度升高,PSI反应中心的氧化和再还原速率上升,同时以热形式耗散的能量(DI o/CS和DI o/RC)下降,表明低浓度ALA可以显著促进苹果叶片光合电子传递和能量转换。另外,果实成熟时,ALA处理果实的硬度、花青素、可溶性固形物、可溶性糖、可溶性蛋白和维生素C含量显著高于对照,而可滴定酸含量则低于对照。【结论】生长季节喷施低浓度ALA可以提高苹果叶片光合性能,改善果实内外品质。

Castelfranco P A, Beale S I .

Chlorophyll biosynthesis:recent advances and areas of current interest

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Abstract BACKGROUND: Children with acute lymphoblastic leukemia (ALL) are at risk for developing musculoskeletal complications during and after their medical treatment. The objective of this study was to examine the feasibility of an in-hospital physical therapy- and home exercise program during the first four phases of medical treatment, for children with newly diagnosed ALL. PROCEDURE: Nine patients, between the ages of 2-14 years old were enrolled within 2 weeks of diagnosis in the study. Each patient was evaluated at study entry, after each of the first four phases of therapy and each time patients were re-admitted to the hospital. Following the initial physical therapy evaluation an individualized home exercise program was developed, consisting of stretching, strengthening, and aerobic exercises. The following end points were measured at each evaluation: gross motor assessment as measured by gross motor function measure (GMFM), health-related quality of life as measured by the PedsQL and parent satisfaction questionnaire. RESULTS: This study was feasible with 98% of the evaluation sessions completed. The GMFM and PedsQL improved steadily throughout the study; however, the PedsQL slightly decreased from interim maintenance to delayed intensification. The parents reported being satisfied with the physical therapy program. CONCLUSION: We demonstrated that an in-hospital- and home exercise physical therapy program during the first four phases of medical treatments is feasible for children with ALL. Future randomized studies are needed to confirm whether an initial physical therapy program at diagnosis in children with ALL will limit functional impairments, improve overall fitness and increase health-related quality of life. Copyright 2011 Wiley-Liss, Inc.

徐刚, 刘涛, 高文瑞 , .

5-氨基乙酰丙酸对蔬菜生理作用的研究进展

金陵科技学院学报, 2010,26(4):52-57.

DOI:10.3969/j.issn.1672-755X.2010.04.012      URL     [本文引用: 2]

5-氨基乙酰丙酸(ALA)作为一种新型植物生长调节剂,同五大类植物激素一样,能对植物生长发育起重要调节作用。近10多年来,关于ALA的研究取得了很大的进展。与五大类植物激素相比,ALA具有一些新的功能及其生理作用,综述了其在蔬菜作物上的应用。

刘玉梅, 艾希珍, 于贤昌 .

5-氨基乙酰丙酸对亚适宜温光条件下黄瓜幼苗光合特性的影响

园艺学报, 2010,37(1):65-71.

URL     Magsci     [本文引用: 1]

为了弄清外源5 - 氨基乙酰丙酸(ALA) 提高亚适宜温光下黄瓜幼苗光合作用的效果, 以‘津优1号’黄瓜为试材, 用0.5 mg·L - 1 ALA喷洒幼苗, 利用光照培养箱模拟亚适宜温光条件(昼/夜温度18 ℃/12 ℃, 光量子通量密度300μmol·m <SUP>- 2</SUP> ·s<SUP>- 1</SUP> ) , 研究ALA对幼苗气体交换参数、叶绿素荧光参数及光合酶活性的影响。结果表明: 亚适宜温光下, 黄瓜幼苗叶片的净光合速率( <EM>P</EM><SUB>n</SUB> ) 和气孔导度(<EM>G</EM><SUB>s </SUB>) 、羧化效率(<EM>CE</EM>) 、表观量子效率(<EM>AQY</EM>) 、光下实际光化学效(<EM>Φ</EM><SUB>PⅡ</SUB> ) 、光化学猝灭系数( <EM>q</EM><SUB>P </SUB>) 、天线转化效率(<EM>F</EM><SUB>v</SUB> ′/<EM>F</EM><SUB>m </SUB>′) 及核酮糖- 1, 5 - 二磷酸羧化酶(RuBPCase) 、果糖- 1, 6 - 二磷酸酶( FDPase) 活性等均明显降低, 胞间CO2浓度(<EM>C</EM><SUB>i </SUB>) 升高, 光补偿点(<EM>LCP</EM>) 、CO2补偿点(<EM>CCP</EM>) 显著下降。与水处理相比, ALA处理的<EM>P</EM><SUB>n</SUB>、<EM>G</EM><SUB>s</SUB>、CE、<EM>AQY</EM>、<EM>Φ</EM><SUB>PSⅡ</SUB>、<EM>q</EM><SUB>P</SUB>、<EM>F</EM><SUB>v</SUB> ′/<EM>F</EM><SUB>m </SUB>′及RuBPCase和FDPase活性等有不同程度提高,<EM>C</EM><SUB>i</SUB>、<EM>LCP</EM>和<EM>CCP</EM>有所降低。亚适宜温光下, 黄瓜幼苗光合速率下降的主要原因是非气孔限制, 外源ALA能明显提高其光合作用。

汪良驹, 姜卫兵, 黄保健 .

5-氨基乙酰丙酸对弱光下甜瓜幼苗光合作用和抗冷性的促进效应

园艺学报, 2004,31(3):321-326.

DOI:10.3321/j.issn:0513-353X.2004.03.007      URL     [本文引用: 2]

5 -氨基乙酰丙酸 (ALA) 10mg·L-1处理能够明显提高弱光下生长的甜瓜幼苗叶片净光合速率 (Pn) ,而低温胁迫明显降低甜瓜幼苗叶片光合能力。经ALA处理过的甜瓜幼苗光合表观量子效率、羧化效率、气孔开度、叶片叶绿素 (特别是叶绿素b)和可溶性糖含量均显著增加。在ALA处理 3d之后 ,将幼苗转移到 8℃低温条件下 4h ,对照叶片光合能力基本丧失 ,而ALA处理叶片仍能维持一定的光合能力 ,并在 2 0h内基本恢复至对照水平。若幼苗在低温下处理 6h ,对照植株完全死亡 ,而ALA预处理植株仅出现少量伤害症状。本研究结果表明 ,ALA处理可以提高植物耐弱光性和抗冷性

徐刚, 刘涛, 高文瑞 , .

ALA对低温胁迫下辣椒植株生长及光合特性的影响

江苏农业学报, 2011,27(3):612-616.

DOI:10.3969/j.issn.1000-4440.2011.03.027      URL     [本文引用: 1]

以辣椒品种超越5号为试材,利用加温玻璃温室模拟大棚的生产方法,探讨低温胁迫期间及常温恢 复过程中叶面喷施25 mg/L5-氨基乙酰丙酸(ALA)对辣椒花期植株生长及光合特性的影响。结果表明:叶面喷施25mg/L ALA提高了低温胁迫下植株的株高、茎粗、干鲜重、叶绿素含量以及净光合速率(Pn)、胞间CO2浓度(Ci)、气孔导度(Gs)和蒸腾速率(Tr)等光 合参数,降低了气孔限制值(Ls)和水分利用率(WUE);常温恢复后,低温+ALA处理的植株生长量和叶片中的叶绿素含量仍明显高于低温处理;低温处理 的叶片Pn、Gs和Tr显著低于常温对照,而低温+ALA处理的较低温度处理有明显提高,但仍低于常温对照。表明叶面喷施ALA能提高植株叶片中叶绿素的 含量,缓解低温胁迫对辣椒叶片气孔的限制,从而提高叶片的光合作用,促进植株生长。

张治平, 张丽丽 .

5-氨基乙酰丙酸对油菜幼苗抗冷性和抗氧化系统的影响

江苏农业科学, 2014,42(2):52-55.

DOI:10.3969/j.issn.1002-1302.2014.02.015      URL     [本文引用: 2]

以油菜品种沪油15为材料,叶面喷施30mg/L5-氨基乙酰丙酸(ALA)4d,再置于 4oC低温条件下处理8d,分析测定了低温处理0.4、8d及恢复4d后光合色素含量及SOD、CAT、APX、POD等抗氧化酶活性、脯氨酸含量、超氧 阴离子(O2-·)生成速率及MDA含量等生理指标,研究了ALA对油菜幼苗抗冷性的影响及作用机制。结果表明,低温胁迫下,喷施ALA明显缓解了油菜叶 片叶绿素a(Chla)、叶绿素b(Chla)和总叶绿素(Chl)含量的下降程度,增加了类胡萝卜素(Car)的含量,显著提高了SOD、CAT、 APX和POD的活性和脯氨酸的含量,降低了O2-·生成速率和MDA含量的上升幅度。低温胁迫时间越长,ALA的作用效果越明显,说明喷施ALA通过促 进叶绿素的合成,提高油菜植株的抗氧化能力,降低活性氧的产生,减少膜脂过氧化的伤害程度,从而提高了油菜幼苗的抗冷性。另外,喷施ALA可以促进油菜幼 苗在受到低温伤害后迅速恢复。

刘涛, 郭世荣, 徐刚 , .

5-氨基乙酰丙酸对辣椒植株低温胁迫伤害的缓解效应

西北植物学报, 2010,30(10):2047-2053.

URL     [本文引用: 2]

以‘超越五号'辣椒品种为试材,研究了低温胁迫期间及随后的常温恢复过程中5-氨基乙酰丙酸(ALA 25 mg·L-1)处理对始花期辣椒植株生长量,叶片中脯氨酸、可溶性糖、可溶性蛋白含量,超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶 (CAT)、抗坏血酸过氧化物酶(APX)活性和电解质渗透率及丙二醛(MDA)含量的影响,以探讨ALA提高辣椒抗寒性的生理机制.结果表明,低温胁迫 下叶面喷施25 mg·L-1的ALA可显著提高辣椒植株生长量,增加叶片中脯氨酸、可溶性糖及可溶性蛋白含量,增强其POD、CAT及APX活性,并显著降低辣椒叶片中 SOD活性、电解质渗透率和MDA含量.叶面喷施ALA也显著降低了恢复过程中辣椒叶片中的渗透调节物质含量和抗氧化酶活性,使膜伤害基本恢复到对照水 平.可见,外源ALA处理可通过提高低温胁迫下辣椒叶片的渗透调节能力和抗氧化能力,促进植株生长,缓解低温胁迫对植株的伤害.

赵艳艳, 燕飞, 胡立盼 , .

5-氨基乙酰丙酸对NaCl胁迫下番茄幼苗光合特性的影响

应用生态学报, 2014,25(10):2919-2926.

URL     Magsci     [本文引用: 1]

<div style="line-height: 150%">为探讨5-氨基乙酰丙酸(ALA)对NaCl胁迫下番茄光合特性的调控作用,以&lsquo;金鹏一号&rsquo;番茄幼苗为试材,研究叶面喷施50 mg&middot;L<sup>-1</sup>或根施10 mg&middot;L<sup>-1</sup> ALA对100 mmol&middot;L<sup>-1</sup> NaCl胁迫下番茄幼苗光合及叶绿素荧光参数的影响.结果表明: NaCl胁迫下,番茄幼苗光合气体交换参数(净光合速率<em>P</em><sub>n</sub>、气孔导度<em>g</em><sub>s</sub>、胞间CO<sub>2</sub>浓度<em>C</em><sub>i</sub>、蒸腾速率<em>T</em><sub>r</sub>)及叶绿素荧光参数(实际光化学量子产量<em>F</em><sub>v</sub>&prime;/<em>F</em><sub>m</sub>&prime;、<em>F</em><sub>m</sub>&prime;、PSⅡ反应中心实际光化学效率<em>&Phi;</em><sub>PSⅡ</sub>、表观光合电子传递效率ETR、光化学淬灭<em>q</em><sub>P</sub>、光化学反应<em>Pc</em>)均显著降低,根施或叶施ALA均可以提高NaCl胁迫下番茄叶片的光合能力,但两种处理方式之间存在一定差异.叶面喷施50 mg&middot;L<sup>-1</sup>ALA或根施10 mg&middot;L<sup>-1</sup>ALA处理均显著提高了番茄叶片<em>P</em><sub>n</sub>、<em>T</em><sub>r</sub>、<em>g</em><sub>s</sub>和<em>C</em><sub>i</sub>,提高了水分利用效率(WUE),显著增加了NaCl胁迫下叶片的最大净光合速率,减轻了光抑制.根施ALA对叶绿素含量的作用效果较好,而叶施ALA对光合参数的作用效果较好,两处理叶绿素荧光参数差异不显著.叶面喷施或根施ALA可以提高番茄幼苗的耐盐性,其调控作用与促进叶绿素合成与稳定、维持正常气孔开闭、降低气孔限制,进而提高NaCl胁迫下番茄叶片的光合能力和PSⅡ光化学效率有关.</div><div style="line-height: 150%"></br>&nbsp;</div>

Korkmaz A, Korkmaz Y, Demirkiran A R .

Enhancing chilling stress tolerance of pepper seedlings by exogenous application of 5-aminolevulinic acid

Environmental and Experimental Botany, 2009,67(3):495-501.

DOI:10.1016/j.envexpbot.2009.07.009      URL     [本文引用: 2]

In this study, the possibility of enhancing chilling stress tolerance of pepper ( Capsicum annuum L.) during early growth stages by exogenous application of 5-aminolevulinic acid (ALA) was investigated. To improve chilling tolerance during seedling stage, ALA was applied in various concentrations (0, 1, 10, 25 and 50 ppm) through three different methods (seed soaking, foliar spray, or soil drench). After ALA applications, the plants were subjected to chilling stress at 3 C for 2 days. Although all ALA application methods improved chilling stress tolerance in pepper seedlings, seed soaking and foliar spray provided better protection against chilling stress compared to soil drench. Exogenous application of ALA provided significant protection against chilling stress compared to non-ALA-treated seedlings, significantly enhancing plant mass and chlorophyll, sucrose, and proline contents. ALA pre-treatment also increased relative water content, stomatal conductance and superoxide dismutase (SOD) enzyme activity and reduced membrane permeability. Of the ALA concentrations, the highest chilling tolerance was obtained with 25 ppm ALA pre-treatment. Results indicate that ALA which is considered as an endogenous plant growth regulator could be used effectively to protect pepper seedlings from damaging effects of chilling stress without any adverse effect on seedling growth.

魏中伟, 马国辉, 龙继锐 , .

5-氨基乙酰丙酸叶面肥对杂交晚稻光合作用和产量的影响

湖南农业科学, 2017(7):65-67,72.

DOI:10.3969/j.issn.1006-060X.2013.07.019      URL     [本文引用: 4]

以4个杂交晚稻为材料,研究了齐穗期叶面喷施1 mg/L的5-氨基乙酰丙酸叶面肥对杂交晚稻生育后期叶片SPAD值、光合作用、干物质积累和经济产量的影响。结果表明:5-氨基乙酰丙酸叶面肥能提高水稻功能叶SPAD值、光合作用,增加干物质积累量,产量增加3.2%~4.0%。

杨文平, 王春虎, 李恒昌 .

叶面喷施ALA对水稻豫粳6号干物质积累及运转的影响

湖北农业科学, 2011,50(7):1313-1314,1318.

DOI:10.3969/j.issn.0439-8114.2011.07.005      URL     [本文引用: 2]

以水稻品种豫粳6号为试验材 料,研究了在孕穗期喷施不同浓度(0、10、30、50、70mg/L)的ALA对水稻干物质积累、运转及产量的影响。结果表明,叶面喷施ALA后,能够 促进干物质的积累和运转;提高水稻的结实率和粒重;使产量增加0.37%~10.19%,其中以喷施30mg/L的ALA效果较好。

徐晓洁, 邹志荣, 乔飞 , .

ALA对NaCl胁迫下不同品种番茄植株光合作用、保护酶活性及果实产量的影响

干旱地区农业研究, 2008,26(4):131-135.

URL     [本文引用: 2]

以耐盐性不同的番茄品种金鹏超冠(耐盐性较强)和中杂9号(耐盐性较弱)为材料,研究了叶面喷施5-氨基乙酰丙酸(ALA)对150 mmol/L,250 mmol/L NaCl胁迫下番茄光合作用、保护酶系统及产量的影响。结果表明,ALA能使NaCl胁迫下番茄叶片的叶绿素含量、净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)升高,胞间二氧化碳浓度(Ci)降低;ALA处理显著提高了高盐胁迫下的SOD、POD和CAT活性,且降低了丙二醛含量,尤其是中杂9号更为显著,ALA处理可能是通过促进保护酶系统活性,保护细胞的光合性能,促进光合作用,从而促进番茄的生长发育,提高产量,缓解盐胁迫对番茄的抑制作用。

童金珠, 邹志荣 .

外源ALA对NaCl胁迫下不同品种西葫芦生理特性及产量的影响

干旱地区农业研究, 2009,27(4):116-120.

URL     [本文引用: 1]

以2个耐盐性不同的西葫芦品种翡翠二号(耐盐性较强)和新早青一代(耐盐性较弱)为材料,研究了叶面喷布5-氨基乙酰丙酸(ALA)对200 mmol/L,400 mmol/L NaCl胁迫下西葫芦生长某些生理特性及果实产量的影响。结果表明,外源ALA对200 mmol/L NaCl胁迫下盐敏感品种的生长及生理特性的缓解作用较好,而在400 mmol/LNaCl胁迫下ALA对耐盐性较强的翡翠二号的缓解作用显著。ALA处理可提高NaCl胁迫下西葫芦叶片的叶绿素、可溶性蛋白及脯氨酸含量,降低细胞膜透性,抑制MDA产生,增加西葫芦植株对盐胁迫的抵抗能力,显著提高盐胁迫下西葫芦产量。

鄢岩, 贺会强, 陈振东 , .

5-氨基乙酰丙酸和叶面肥对荒漠区设施番茄和辣椒生长发育、产量和品质的影响

西北农业学报, 2016,25(10):1515-1521.

DOI:10.7606/j.issn.1004-1389.2016.10.014      URL     [本文引用: 2]

为外源5-氨基乙酰丙酸(ALA)在荒漠地区设施园艺中的推广和应用,以番茄和辣椒为材料,设置清水、叶面肥和外源ALA(5-氨基乙酰丙酸)3个处理,研究外源ALA和叶面肥处理对荒漠区番茄和辣椒生长及产量品质的影响.结果表明,外源ALA和叶面肥均可促进荒漠区番茄和辣椒的生长,并显著改善果实相关品质.其中ALA处理的番茄株高、茎粗、叶片数和冠径与对照相比分别提高15.45%、8.33%、13.69%和8.58%,辣椒叶片数提高22.79%;番茄果实固酸比提高8.86%,辣椒可溶性固形物质量分数提高8.59%,并且还能够提高番茄和辣椒的产量.可见,ALA和叶面肥对荒漠区设施番茄和辣椒的生长都表现一定的促进作用,对品质有不同程度的改善,同时也提高了产量.

Han Y L, Li H, Miao Y H , et al.

Effects of illumination intensity,5-aminolevulinic acid concentration and their interaction on chlorophyll fluorescence parameters and yield of summer maize

Agricultural Science & Technology, 2013,14(5):757-762,805.

URL     [本文引用: 2]

[Objective] This study aimed to investigate the influence of illumination intensity, 5-aminolevulinic acid (ALA) concentration and their interaction on chlorophyll fluorescence parameters and yield of summer maize. [Method] Two illumination intensity levels and five ALA concentrations were applied in the experiment using 2 5 completely balanced program. The two illumination intensity levels were natural light (S0 ) and 60% shade (S1 ), and five ALA concentrations were0, 1, 10, 25 and 50 mg/L. [Result] The relative chlorophyll content of leaf (SPAD), the optimal/maximal quantum yield of PS (Fv/Fm), the photochemical quenching coefficient (qP), electron transport rate (ETR), grain number per cob and grain weight per cob in S1 treatment were significantly reduced compared with that in S0 . However, the non-photochemical quenching coefficient (qN) was significantly increased. The responses of these parameters to ALA were different under S0 and S1 treatments. The SPAD, Fv/Fm, qP, ETR, grain number and grain weight per cob were firstly increased, but then decreased following the raise of ALA concentration, ranging from0 to 50 mg/L, whereas qN showed opposite trend. The effect of the interaction of illumination intensity and ALA concentration on these parameters was significant (P0.05). Under natural light, summer maize could obtain higher SPAD, Fv/Fm, qP and ETR and lower qN combined with low concentration of ALA. However, high concentration of ALA was needed under shading to get the same results. [Conclusion] Soaking seed in suitable concentration of ALA can reduce the yield loss of summer maize caused by short-term shading in seedling stage.

Hotta Y, Tanaka T, Takaoka H , et al.

New physiological effects of 5-aminolevulinic acid in plants:the increase of photosynthesis,chlorophyll content and plant growth

Bioscience Biotechnology and Biochemistry, 1997,61(12):2025-2028.

DOI:10.1271/bbb.61.2025      URL     PMID:27396878      [本文引用: 1]

5-Aminolevulinic acid (ALA) promoted the growth and yield of several crops and vegetables at concentrations lower than those eliciting herbicidal responses, i.e., less than 1.8 mm by foliar spray and 60 0204m by root soaking. To evaluate the physiological action of ALA, the effects of ALA on plants were examined by several bioassay systems at 0.0006090009600 0204m. ALA at 0.060900096 0204m by root soaking increased the growth of rice seedlings in light, but did not affect this in darkness. In horseradish shoot primordia, promotion by ALA was not proportional among total chlorophyll content, chlorophyll concentration, and fresh weight. In the test using pothos, ALA at 0.06 0204m elicited the accumulation of chlorophyll, but the photosynthesis of the plants was promoted by treatment together with ALA and nutrients. These results suggest that ALA have a variety of plant physiological effects on chlorophyll synthesis, photosynthesis, and plant growth, and ALA acts as a growth regulator in plants at low concentrations. These effects of ALA were also assumed to be linked to light irradiation and an uptake of fertilizer by plants. However, excess ALA suppressed these effects.

Hotta Y, Tanaka H, Takaoka Y , et al.

Promotive effects of 5-aminolevulinic acid on the yield of several crops

Plant Growth Regulation, 1997,22(2):109-114.

DOI:10.1023/A:1005883930727      URL     [本文引用: 2]

The role of 5-aminolevulinic acid (ALA) as a precursor of chlorophyll or a herbicide is well documented. In the present report, to elucidate the physiological effects of ALA, this compound was applied to the foliage of radish at 0.06, 0.18, 0.6, 1.8 and 6 mM. ALA at 0.06ndash;1.8 mM increased the dry weight of radish root (underground part), but injured radish seedlings at 6 mM. Also, the application of ALA at 0.18 and 0.6 mM increased fixation of CO 2 in light and decreased release of CO 2 in darkness. Furthermore, the effects of foliar treatment of ALA on the yield of several crops were also evaluated. The results showed that an application of ALA at low concentrations increased the growth and yield by 10ndash;60percnt; over the control on kidney bean, barley, potato and garlic. These results suggest that ALA has plant growth regulating properties at low concentrations and may enhance agricultural productivity.

要娟娟, 薛泽民, 赵萍萍 , .

施肥与种植密度对春玉米SPAD值的影响

山西农业科学, 2011,39(10):1060-1063.

DOI:10.3969/j.issn.1002-2481.2011.10.09      URL     [本文引用: 1]

有效的氮肥管理一直是农业可持续发展面临的重大挑战.通过田间试 验,采用裂区-裂区设计,研究了不同密度、不同氮肥用量和有机肥施用的不同措施对春玉米抽雄期穗位叶叶绿素含量(SPAD值)的影响.结果表明:种植密度 (60000,90000株/hm2)、氮肥施用量(N0,N225,N300)及有机肥的施用与否均显著影响玉米抽雄期的SPAD值;施用有机肥有助于 提高玉米SPAD值;种植密度增加,玉米SPAD值呈下降趋势;施氮量对玉米SPAD值的影响,在不施有机肥情况下呈直线型,而在施有机肥条件下呈抛物线 型.

Tanaka Y, Tanaka A, Tsuji H .

Effects of 5-aminolevulinic acid on the accumulation of chlorophyll b and apoproteins of the light-harvesting chlorophyll a/b-protein complex of photosystem Ⅱ

Plant and Cell Physiology, 1993,34(3):465-472.

DOI:10.1094/Phyto-83-456      URL     [本文引用: 1]

The effects were examined of 5-aminolevulinic acid (ALA) on the accumulation of Chl and apoproteins of light-harvesting Chl a/b-protein complex of photosystem II (LHCII) in cucumber cotyledons under intermittent light. A supply of ALA preferentially increased the accumulation of Chl a during intermittent illumination. However, when cotyledons were pretreated with a brief exposure to light or benzyladenine (BA), the stimulatory effect of ALA on the increase in the level of Chl b was greater than that in the level of Chl a, resulting in decreased ratios of Chl a/b. Time-course experiments with preilluminated cotyledons revealed that LHCII apoproteins accumulated rapidly within the first 30 min of intermittent illumination with a decline during sub-sequent incubation in darkness. A supply of ALA did not affect the accumulation of LHCII apoproteins during the intermittent light period, but it efficiently inhibited the decline in their levels during the subsequent darkness. After exposure to a single pulse of light of BA-treated cotyledons, the prompt increase in levels of LHCII apoproteins was not accompanied by the formation of Chl b, which began to accumulate later. The pattern of changes in levels of LHCII apoproteins was quite similar to that in levels of Chl a. These results suggest that LHCII apoproteins are first stabilized by binding with Chl a and that an increased supply of Chl a and the accumulation of LHCII apoproteins are prerequisites for the formation of Chl b.

Sasaki K, Marquez F J, Nishio N , et al.

Promotive effects of 5-aminolevulinic acid on the growth and photosynthesis of Spirulina platensis

Journal of Fermentation and Bioengineering, 1995,19(5):453-457.

DOI:10.1016/0922-338X(95)91261-3      URL     PMID:7737530      [本文引用: 1]

Photosynthetic activity in terms of O 2 evolution and the growth of Spirulina platensis was stimulated by adding 5-aminolevulinic acid (ALA, 500 mg/ l ) to photoautotrophically growing cells. After ALA was added to the medium, intracellular accumulations of phycocyanin and chlorophyll were stimulated simultaneously, followed by enhancement of the photosynthetic activities of photosystems I and II, and lastly, growth was promoted. ALA did not directly activate the photosynthetic electron transport system. However, during a 3-h incubation of intact cells with ALA, photosynthetic activity was enhanced.

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