全 文 ：作物学报 ACTA AGRONOMICA SINICA 2010, 36(5): 856−862 http://www.chinacrops.org/zwxb/
ISSN 0496-3490; CODEN TSHPA9 E-mail: xbzw@chinajournal.net.cn

本研究由云南省烟草公司科技项目计划(07A30)资助。
*
通讯作者(Corresponding author): 赵正雄, E-mail: zhaozx0801@163.com; Tel: 0871-5220536
第一作者联系方式: E-mail: zuo-lijuan@163.com
Received(收稿日期): 2009-12-01; Accepted(接受日期): 2010-02-07.
DOI: 10.3724/SP.J.1006.2010.00856
增加施钾量对红花大金元烤烟部分生理生化参数及“两黑病”发生的
影响
左丽娟 1 赵正雄 1,* 杨焕文 1 段凤云 2 王德勋 2 徐发华 2 吕 芬 1
1 云南农业大学烟草学院, 云南昆明 650201; 2 云南省烟草公司大理州公司, 云南大理 671000
摘 要: 针对生产中红花大金元烤烟品种对黑胫病和根黑腐病(俗称“两黑病”)抗性较低的问题, 通过田间试验研究
了不打杀菌剂条件下增加施钾量(与目前生产推荐 187.5 kg K2O hm–2相比, 分别增加 75和 150 kg K2O hm–2)对红花大
金元烟株移栽后不同时间段健康烟株烟叶中的总氮、蛋白质、游离氨基酸、可溶性糖、总酚和类黄酮含量等生理生
化指标及烤烟“两黑病”发病情况的影响。结果表明, 随施钾量增加, 烟叶内总氮、蛋白质和游离氨基酸含量下降, 而
可溶性糖、总酚和类黄酮含量则相反; 同时“两黑病”发病情况也呈明显下降趋势。上述指标在对照与高钾处理间差
异显著, 且上述结果在烟株移栽后不同时间段规律一致。烤后烟叶产量、产值以中钾处理最大, 高钾处理次之, 对照
最小。中、高钾两处理产量比对照分别增加 4.49%和 1.55%, 产值分别增加 18.27%和 12.34%, 中钾处理与对照间差
异明显。说明目前红花大金元烤烟生产中的推荐施钾量偏低, 应适当增加施钾量以增强烟株抗性, 减少“两黑病”病害
的发生。就本试验而言, 当地施钾量以增加 75 kg K2O hm–2(即总量达 262.5 kg K2O hm–2)为宜。
关键词: 烤烟; 红花大金元; 钾; 抗性; 黑胫病; 根黑腐病
Effect of Increasing K Application Rate on Partial Physiological & Biochemical
Parameters and Occurrence of Black Shank and Black Root Rot in Tobacco
Variety Hongda
ZUO Li-Juan1, ZHAO Zheng-Xiong1,*, YANG Huan-Wen1, DUAN Feng-Yun2, WANG De-Xun2, XU
Fa-Hua2, and LÜ Fen1
1 College of Tobacco Science, Yunnan Agricultural University, Kunming 650201, China; 2 Dali Tobacco Company, Yunnan Tobacco Company, Dali
671000, China
Abstract: Flue-cured tobacco production plays an important role for the most farmers in Yunnan province. However, farmers are
not willing to plant Hongda variety, partly because the tobacco seedlings are much susceptible to black shank and black root rot
even when fertilized with the same ratio of potassium as for the resistant varieties such as K326 and Yun 85. In addition to the
genetic difference in resistance, whether potassium application rate recommended for Hongda variety in the production is
sufficient to enhance its resistance to diease is not clear. Therefore, three potassium application rates, including 187.5 (CK, rec-
ommended in the production), 262.5, and 337.5 kg K2O ha–1, were designed to compare the tobacco seedlings’ resistance to black
shank and black root rot in field experiment. The leaves were sampled from healthy tobacco plants (Nicotiana tabacum L. var.
Hongda) at 33, 41, 62, 69, 76 days after transplanting (DAT), and some physiological-biochemical parameters such as contents of
nitrogen, protein, free amino acid, soluble sugar, phenol and flavonoid were determined. Meanwhile, the severity of black shank
and black root rot on tobacco plants was recorded in the field at 10-day interval from 24 DAT to 114 DAT. The results showed that
the contents of N, protein and free amino acid reduced, the contents of flavonoid, phenol and soluble sugar in leaf increased when
potassium application rate was raised. In addition, decreased severity of black shank and black root rot was observed with in-
creasing potassium rate at 44, 54, 64, 74, 84, 94, 104, and 114 DAT. There was significant difference between the two treatments
with K2O application rate at 337.5 and 187.5 kg ha–1 respectively in the parameters mentioned above. However, the leaf yield and
economic value reached the peak when potassium was applied at 262.5 kg K2O ha–1. Compared with those of the control, the leaf
yield and economic value increased by 4.49% and 18.27% respectively in the treatment with potassium application rate at 262.5
第 5期 左丽娟等: 增加施钾量对红花大金元烤烟部分生理生化参数及“两黑病”发生的影响 857


kg K2O ha–1, while by 1.55% and 12.34% respectively in the treatment with K2O rate at 337.5 kg ha–1. It suggested that potassium
application rate recommended in the production of Nicotiana tabacum L. variety Hongda is inadequate, therefore a relatively
higher potassium application rate, i.e. 262.5 kg K2O ha–1 in the experiment, should be preferred to decrease the severity of black
shank and black root rot.
Keywords: Flue-cured tobacco; Variety Hongda; Potassium; Resistance; Black shank; Black root rot
烟草黑胫病(Phytophthora parasitica var. nico-
tianae)和根黑腐病(Thielaviopsis basicola)俗称“两黑
病”, 是世界性的烟草病害, 烟株苗期至成株期均易
被侵染, 在我国多数烟区危害较大[1-3]。红花大金元
是云南烤烟主要品种之一, 清香型风格突出, 卷烟
工业需求量大; 但与 K326、云烟 85 等烤烟品种相
比, 对“两黑病”等主要病害抗性较差[4-5], 且烘烤困
难, 种植规模难以扩大。施肥(尤其是施用钾肥)是影
响植物抗病性的一个重要因素。合理施用钾肥能明
显降低柠檬流胶病[6]、茶树炭疽病、轮斑病和云纹
叶枯病[7]、枇杷茎腐病[8]、玉米茎腐病[9]、小麦白粉
病[10]以及山茱萸炭疽病[11]等的发病率。就红花大金
元品种烤烟而言, 尽管目前生产中钾肥往往按 N∶
K2O为 1.0∶2.5~3.0的比例施用, 但其绝对用量明显
低于对 K326 和云烟 85 等品种。除遗传因素外, 是
否钾肥施用不足限制了红花大金元对“两黑病”的抗
性, 能否通过进一步增加施钾量以增强其抗性, 值
得研究。因此, 本试验以红花大金元烤烟生产中钾
肥推荐施用量为对照, 研究了提高施钾量对烟株部
分生理生化参数及“两黑病”发生的影响, 以为降低
病害提供理论依据和实践指导。
1 材料与方法
1.1 试验品种和试验地点
2008年在云南省大理州弥渡县新街乡西河新村
种植烤烟品种红花大金元。试验前土壤含有机质
34.5 g kg–1、碱解氮 149.5 mg kg–1、速效磷 29.8 mg
kg–1、速效钾 275 mg kg–1、pH 7.09。
1.2 试验设计
设 3个钾水平, 分别为 187.5(当地推荐)、262.5
kg K2O hm–2和 337.5 kg K2O hm–2。3次重复, 随机
区组排列。各处理氮、磷用量相同, 分别为 75 kg N
hm–2和 75 kg P2O5 hm–2。其中, 除纯氮按当地追肥
习惯于移栽后 7 d和 25 d采用硝酸钾兑水分别浇施
20%外, 其余氮、钾肥和全部的磷肥均于移栽前基施
(所用肥料种类为烤烟专用复合肥和硫酸钾), 即将
肥料放到移栽塘底(采用施肥器皿, 保证处理内各株
烟施肥一致), 拌土后栽烟。每小区栽烟 100 株, 株
行距 50 cm×110 cm。于 2008年 5月 4日移栽。大田
管理除不打防病药剂之外, 其余均按当地要求进行,
各处理一致。
1.3 烟叶生理生化参数的测定
以小区为单位, 分别于移栽后 33、41、62、69
和 76 d各选取两株具有代表性健康烟株的中部烟叶
(各处理一致), 作为混合样, 测定烟叶中总氮、蛋白
质、游离氨基酸、可溶性糖、总酚和类黄酮含量。
采用 H2SO4-H2O2消煮, 凯氏定氮法[12]测定氮含量。
参照 Pirie等[13]的方法测定总酚和类黄酮含量, 即取
0.5 g新鲜叶片剪成 1~2 mm小块后加含 1% HCl的
甲醇液 5 mL提取 24 h (4℃), 取 1 mL提取液稀释至
5 mL, 测定 OD280nm 和 OD325nm 值。类黄酮含量以
OD325nm g–1 FW 表示, 总酚含量以没食子酸标准曲
线进行计算。采用蒽酮比色法测定可溶性糖含量、
采用茚三酮溶液显色法测定游离性氨基酸含量、采
用考马斯亮蓝法测定蛋白质含量[14]。
1.4 病情调查
从移栽后 24 d开始, 每 10 d观察一次“两黑病”
的发生情况, 以株为单位, 调查全部烟株。按如下标
准分别进行黑胫病和根黑腐病病情分级。
1.4.1 烟草黑胫病 0级：全株无病; 1级：茎部
病斑不超过茎围的 1/2, 或半数以下叶片轻度凋萎,
或下部少数叶片出现病斑; 2级：茎部病斑超过茎围
的 1/2, 或半数以上叶片凋萎; 3 级：茎部病斑环绕
茎围, 或 2/3 以上凋萎; 4 级：病株全部叶片凋萎或
枯死。
1.4.2 烟草根黑腐病 0 级: 无病, 植株生长正
常; 1 级: 株高基本正常或稍有矮化, 中下部叶片褪
绿(或变色), 中午萎蔫夜间恢复, 半数以下根坏死;
2级: 比正常植株矮 1/4~1/3, 半数以上叶片萎蔫, 中
下部叶片稍有干尖干边, 半数以上根坏死; 3 级: 比
正常植株矮 2/5~1/2, 2/3以上叶片萎蔫, 明显干尖干
边或枯黄, 2/3以上根坏死; 4级: 植株严重矮化, 全
株叶片凋萎。根几乎全部坏死, 近地表的次生根明
显受害。
1.4.3 病情计算公式
发病率(%) = (发病株数/调查总株数)×100
858 作 物 学 报 第 36卷

病情指数 = ∑(各级病株×该病级值)/(调查总株
×最高级值)×100
1.5 烟叶产量、产值的测定
以小区为单位 , 烟叶成熟时挂牌采收和烘烤 ,
按国家烟叶分级标准计算烟叶产量、产值及均价。
1.6 数据处理
采用 Microsoft Excel 和 SPSS 软件进行数据统
计分析。
2 结果与分析
2.1 施钾量对烟叶中总氮、蛋白质和游离氨基酸
含量的影响
表1表明, 烟叶内总氮、蛋白质、游离氨基酸含
量随生育期推移, 前期逐渐增加, 而从移栽后 41 d时
开始逐渐下降。不同施钾处理间比较而言, 上述指
标皆随施钾量增加而下降, 且低钾处理(K2O 187.5
kg hm–2)与高钾处理(K2O 337.5 kg hm–2)间差异显著;
这些差异在不同时间段规律一致。
2.2 施钾量对烟叶中可溶性糖、总酚和类黄酮含
量的影响
烟叶中可溶性糖、总酚和类黄酮含量因施钾量
不同而有明显差异(表 2), 不同时间段测定结果规律
一致 , 即随施钾量增加 , 上述指标上升 , 且低钾处
理与高钾处理之间差异显著。以上生理生化参数皆
随生育期发展逐渐增加, 到移栽后 69 d 达最大值,
之后则随烟叶的成熟而下降。
2.3 施钾量对烤烟“两黑病”发生的影响
表 3表明, 移栽后 24 d田间尚无黑胫病和根黑

表 1 施钾量对烟叶中总氮、蛋白质和游离氨基酸含量的影响
Table 1 Effect of K2O application rate on the contents of nitrogen, protein, and free amino acid in tobacco leaves
移栽后天数 Days after transplanting 项目
Item
施钾量
K2O 33 d 41 d 62 d 69 d 76 d
187.5 kg hm–2 2.43 a 2.78 a 1.96 a 1.70 a 1.51 a 总氮
Total nitrogen (%) 262.5 kg hm–2 2.27 ab 2.65 ab 1.75 ab 1.63 ab 1.42 ab
337.5 kg hm–2 2.11 b 2.50 b 1.61 b 1.50 b 1.24 b

187.5 kg hm–2 13.03 a 15.19 a 11.92 a 11.15 a 9.87 a 蛋白质
Protein (%) 262.5 kg hm–2 12.54 ab 14.75 ab 11.37 ab 10.65 ab 9.36 ab
337.5 kg hm–2 11.78 b 14.36 b 11.03 b 10.17 b 8.64 b

187.5 kg hm–2 0.32 a 0.35 a 0.29 a 0.26 a 0.24 a 游离氨基酸
Free amino acid (mg g–1) 262.5 kg hm–2 0.29 b 0.34 ab 0.27 ab 0.25 ab 0.23 ab
337.5 kg hm–2 0.28 b 0.33 b 0.26 b 0.24 b 0.22 b
各列中标以不同字母的数字在 0.05水平上差异显著。
Values in each column followed by different letters are significantly different at the 0.05 probability level.

表 2 施钾量对烟叶中可溶性糖、总酚和类黄酮含量的影响
Table 2 Effect of K2O application rate on the contents of soluble sugar, phenol and flavonoid in tobacco leaves
移栽后天数 Days after transplanting 项目
Item
施钾量
K2O 33 d 41 d 62 d 69 d 76 d
187.5 kg hm–2 1.70 b 6.09 b 14.13 b 18.65 b 17.00 b 可溶性糖
Soluble sugar (%) 262.5 kg hm–2 1.74 ab 6.20 ab 14.58 a 19.01 ab 17.41 ab
337.5 kg hm–2 1.76 a 6.25 a 14.69 a 19.25 a 17.62 a

187.5 kg hm–2 4.33 b 5.69 b 8.97 b 11.23 b 10.34 b 总酚
Phenol (mg g–1) 262.5 kg hm–2 4.51 ab 5.90 ab 9.12 ab 11.44 ab 10.58 ab
337.5 kg hm–2 4.69 a 6.01 a 9.30 a 11.55 a 10.64 a

187.5 kg hm–2 270.20 b 354.38 b 579.17 b 718.04 b 662.79 b 类黄酮
Flavonoid (OD325 g–1) 262.5 kg hm–2 288.63 ab 382.09 a 594.14 ab 735.73 a 683.87 a
337.5 kg hm–2 297.83 a 385.47 a 599.38 a 742.36 a 686.05 a
各列中标以不同字母的数字在 0.05水平上差异显著。
Values in each column followed by different letters are significantly different at the 0.05 probability level.


表 3 施钾量对烤烟“两黑病”发病情况的影响
Table 3 Effect of K2O application rate on occurrence of black shank and black root rot in flue-cured tobacco plant
移栽后天数 Days after transplanting 病害
Disease type
项目
Item
施钾量
K2O 24 d 34 d 44 d 54 d 64 d 74 d 84 d 94 d 104 d 114 d
187.5 kg hm–2 0 0 3.84 a 6.29 a 7.34 a 11.88 a 12.05 a 13.11 a 13.46 a 14.17 a 黑胫病
Black shank
disease
发病率
Incidence
(%) 262.5 kg hm–2 0 0 2.79 ab 4.54 ab 5.59 ab 10.13 ab 9.21 b 10.63 ab 11.69 ab 12.76 ab
337.5 kg hm–2 0 0 2.46 b 3.87 b 4.93 b 8.44 b 8.15 b 9.22 b 9.57 b 10.63 b

187.5 kg hm–2 0 0 1.92 a 3.14 a 3.67 a 6.64 a 6.56 a 7.62 a 8.24 a 8.77 a 病情指数
Severity
262.5 kg hm
–2 0 0 1.05 ab 2.18 ab 2.71 ab 5.50 ab 5.14 b 6.29 ab 7.35 ab 7.53 ab

337.5 kg hm–2 0 0 0.79 b 1.76 b 2.29 b 4.57 b 4.70 b 5.76 b 6.02 b 6.47 b

187.5 kg hm–2 0 2.74 a 7.68 a 8.73 a 8.73 a 8.73 a 8.73 a 8.73 a 8.73 a 8.73 a 根黑腐病
Black root rot
发病率
Incidence
(%) 262.5 kg hm
–2 0 1.71 ab 5.59 ab 6.63 ab 6.63 ab 6.63 ab 6.63 ab 6.63 ab 6.63 ab 6.63 ab
337.5 kg hm–2 0 1.38 b 4.23 b 5.28 b 5.28 b 5.28 b 5.28 b 5.28 b 5.28 b 5.28 b

187.5 kg hm–2 0 1.11 a 4.10 a 4.63 a 4.63 a 4.63 a 4.63 a 4.63 a 4.63 a 4.63 a 病情指数
Severity
262.5 kg hm
–2 0 0.68 ab 2.70 ab 3.23 ab 3.23 ab 3.23 ab 3.23 ab 3.23 ab 3.23 ab 3.23 ab
337.5 kg hm–2 0 0.43 b 2.29 b 2.82 b 2.82 b 2.82 b 2.82 b 2.82 b 2.82 b 2.82 b
各列中标以不同字母的数字在 0.05水平上差异显著。
Values in each column followed by different letters are significantly different at the 0.05 probability level.


860 作 物 学 报 第 36卷

腐病的发生。而从移栽后 34 d和 44 d分别有根黑腐
病和黑胫病的出现, 且随着生育期发展, 病情趋向
加重。其中, 黑胫病发生一直持续到烟叶采收结束,
而根黑腐病于移栽后 54 d达到顶峰后保持稳定。不
同施钾量处理间“两黑病”发生差异明显, 两个病害
的发病率和病情指数均随施钾量增加而下降, 且低
钾处理与高钾处理之间差异显著; 上述差异在不同
时间段有一致的规律。说明钾肥施用数量对烤烟“两
黑病”发生影响很大, 增施钾肥能提高作物的抗病性。
2.4 施钾量对烤烟产量、产值的影响
表 4 表明, 在当地目前的施钾量(K2O 187.5 kg
hm–2)基础上, 适当增加施钾量可增加产量、产值和
均价, 但增加到一定程度后, 增产不明显甚至减产。
与对照相比, 中钾和高钾处理的产量、产值、均价
分别比对照增加 4.49%、18.27%、13.16%和 1.55%、
12.34%和 2.36%, 其中对照与中钾处理间差异显著。
3 讨论
植物体内总氮、游离氨基酸、可溶性糖、酚类
和类黄酮等含量与植物抗病性有密切的关系。多数
研究认为, 植物体内总氮[15-17]、氨基酸[17-19]等含量
高往往发病较重, 而糖[17,20-24]、酚类[17,25-29]以及类黄
酮[17,30-31]含量高则有助于抗病。钾营养状况直接或
间接影响着上述物质的含量 [27-29,32-35], 本试验结果
也进一步说明了这一点。烟株体内可溶性糖、总酚
和类黄酮的含量均随施钾量增加而明显增加, 与以
往研究结果[27-29,34-35]有相似的结论; 而体内总氮、蛋
白质和游离氨基酸含量则随施钾量增加而呈降低趋
势, 这与烟株处于较高的钾素供应状况进而影响了
氮代谢有关。有研究认为[36-37], 适量钾素可促进玉
米叶片对氮素的吸收和利用, 促进氨基酸合成蛋白
质或其他物质; 但超过一定范围之后, 高钾处理植
株氮代谢受到干扰, 叶片对氮的同化能力减弱, 从
而出现蛋白质含量下降。在中性和石灰性两种紫色
土上的烟草试验也表明 , 随着 K2SO4 施用量增高 ,
烟叶的水溶性总糖增加, 而总氮和蛋白质含量呈降
低趋势[33]。不同施钾量烟株体内上述物质含量的差
异在一定程度上导致了烤烟“两黑病”发病情况的差
异, 即随施钾量增加, 烤烟抗性增强, “两黑病”发病
率降低, 病情减弱, 这与以往报道类似[6-11,27-29,38-40]。
试验结果也表明, 与 K326、云烟 85等品种相比, 红
花大金元对“两黑病”抗性较低的原因除与品种间遗
传差异有关外, 在一定程度上应与该品种目前生产
中钾肥施用不足有直接关系。
从烤后烟叶产量、产值来看, 与对照(即当地目
前推荐施钾量)相比, 钾肥用量增加, 烟株“两黑病”
发病率降低, 病情减轻, 但其产量、产值并没有呈现
一直增加的趋势, 而以中钾处理(K2O 262.5 kg hm–2)
最大, 高钾处理(K2O 337.5 kg hm–2)尽管仍优于对照,
但与中钾处理相比有一定程度的下降。这与以往烟
草[41-43]、玉米[9,36,44-45]等的研究有相似的结论。高钾
处理烟株产量下降原因应在一定程度上与施钾量超
过一定范围后 , 烟株最大叶面积和叶面积指数下
降[42-43], 单叶重降低[41]有关。本试验中, 与对照烟株
相比 , 初烤前中钾处理烟株的叶面积系数增加了
7.47%, 而高钾处理则与对照相似 (为对照的
99.77%)(待发表)。高钾处理烟株光合面积的下降应
是其产量低于中钾处理烟株的可能原因之一。此外,
也有研究认为, 在含钾丰富的土壤中施用大量钾肥
所引起的 K-N、K-Mg离子间的拮抗作用也可能是影
响高钾烟株产量的一个原因[42]。试验中高钾处理烟叶
的蛋白质含量低于其他两处理也似乎说明了这一点。
4 结论
施钾量影响着烟株体内总氮、蛋白质、游离氨
基酸、可溶性糖、总酚和类黄酮等含量, 并在一定
程度上影响着烟株抗病性。生产中红花大金元烤烟
对“两黑病”抗性较低除遗传因素外, 在一定程度上
应与该品种现阶段钾肥推荐施用量偏低有直接关
系。与目前生产推荐相比, 适当增加施钾量可以提

表 4 施钾量对烤烟产值量的影响
Table 4 Effect of K2O application rate on yield and economic value of flue-cured tobacco
施钾量
K2O
产量
Yield (kg hm–2)
产值
Economic value (Yuan hm–2)
均价
Average price (Yuan hm–2)
187.5 kg hm–2 1842.87 b 22684.68 b 12.31 b
262.5 kg hm–2 1925.66 a 26828.71 a 13.93 a
337.5 kg hm–2 1871.40 ab 25484.29 ab 12.60 a
各列中标以不同字母的数字在 0.05水平上差异显著。
Values in each column followed by different letters are significantly different at the 0.05 probability level.
第 5期 左丽娟等: 增加施钾量对红花大金元烤烟部分生理生化参数及“两黑病”发生的影响 861


高红花大金元烤烟抗性, 降低“两黑病”的发生。综合
投入和产出因素, 试验当地红花大金元烤烟生产中
的施钾量以增加 75 kg K2O hm–2为宜, 既有较好控
病效果的同时, 又可显著增产。

致谢：本试验得到了云南农业大学资源与环境学院
植物营养专业刘智强、赵芳、张英和刘洋同学的大
力支持和帮助, 在此表示感谢。
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