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湖北贝母Fritillaria hupehensis为百合科Liliaceae贝母属Fritillaria植物,以干燥鳞茎入药,含多种异甾体、甾体类生物碱,具有化痰止咳、散结的功效,是恩施州大力发展药材之一[1]。由于野生资源日趋匮乏,目前多为人工栽培。如何提高湖北贝母产量和品质,成为湖北贝母种植亟需解决的问题。氮(N)、磷(P)、钾(K)是植物生长必需的矿质元素,对植物生长和品质形成有重要影响。研究表明:适量施用磷肥可增强浙贝母F. thunbergii抗逆性,有利于生物碱类物质质量分数及产量的提高[2]。不同品种钾肥配施可促进浙贝母的养分吸收,显著提高浙贝母鳞茎的生物量及其生物碱的积累量,且硫酸钾对生物碱质量分数的提升作用优于氯化钾[3]。陈天德等[4]研究发现:施用240 kg·hm−2氮、105 kg·hm−2五氧化二磷(P2O5)、135 kg·hm−2氧化钾 (K2O)为浙贝母最佳施肥量。陆中华等[5]认为:施用300 kg·hm−2氮、80 kg·hm−2五氧化二磷、180 kg·hm−2氧化钾为露地条件下浙贝母获得高产且贝母素总量较高的施肥方案。上述施肥量差异可能受土壤肥力、种植密度及作物品种等因素的影响。近年来有关施肥对贝母产量与品质影响的研究多以浙贝母为主,针对湖北贝母养分需求规律及有效成分积累的研究未见系统报道。
氮是植物体内蛋白质、代谢酶、核酸及生物碱等次生代谢产物的重要物质组成成分,氮肥通过影响药用植物的养分吸收与代谢转化等过程,成为限制中药材产量与品质的关键因素[6−7],并且药用植物在不同生育期对养分的需求及有效成分的积累均存在显著差异[8−9]。鉴于此,本研究以湖北贝母为供试材料,研究不同氮施用量对各生育期湖北贝母鳞茎氮磷钾吸收及生物碱动态积累的影响,以期明确湖北贝母不同生育期矿质元素的吸收规律及主要生物碱的积累特征,为湖北贝母规范化生产及科学施肥提供理论依据。
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由表1可知:不同氮肥用量对湖北贝母幼苗叶长、叶宽、株高、茎粗等生长指标影响差异显著。随氮肥用量增加,苗期、花期幼苗的叶长、叶宽、株高、茎粗总体呈增加趋势,且均在N2处理时达到峰值。各生育期湖北贝母鳞茎的鲜质量及干质量均随氮肥用量增加显著增加(P<0.05),且均在N2处理时达到峰值;但各生育期内不同氮肥处理时鳞茎的折干率总体差异不显著。相同氮肥处理下,苗期至花期湖北贝母鳞茎干质量的增长量为0.53~2.72 g·株−1,增长率为22.36%~83.44%;花期至收获期湖北贝母鳞茎干质量增长量为0.30~1.85 g·株−1,增长率为10.34%~50.82%。此结果表明:湖北贝母鳞茎干物质积累速率在苗期至花期较快,此后其产量增长量减缓。
表 1 不同氮肥施用量时湖北贝母的生长指标
Table 1. Growth indices of F. hupehensis under different nitrogen dosages
生育期 处理 叶长/cm 叶宽/cm 叶型指数 株高/cm 茎粗/cm 苗期 N0 11.15±0.12 b 1.03±0.06 c 10.88±0.56 a 19.70±0.92 c 0.50±0.01 b N1 11.33±0.12 b 1.33±0.05 b 8.55±0.29 b 29.30±1.00 b 0.53±0.02 ab N2 11.45±0.19 a 1.55±0.10 a 7.39±0.25 d 33.90±3.11 a 0.55±0.03 a N3 11.00±0.19 b 1.42±0.06 b 7.76±0.21 c 32.85±0.95 a 0.50±0.02 b 花期 N0 11.77±0.10 b 1.72±0.10 b 6.86±0.48 a 36.45±0.58 b 0.50±0.02 b N1 11.90±0.07 b 1.80±0.11 a 6.61±0.39 a 38.00±0.94 b 0.55±0.02 b N2 14.20±0.08 a 2.10±0.10 b 6.76±0.51 a 40.50±1.16 a 0.65±0.03 a N3 12.00±0.20 b 1.70±0.12 b 7.06±0.61 a 37.60±2.02 b 0.51±0.04 b 收获期 N0 - - - - - N1 - - - - - N2 - - - - - N3 - - - - - 生育期 处理 鳞茎鲜质量/(g·株−1) 鳞茎干质量/(g·株−1) 折干率/% 鳞茎干质量产量/(g·m−2) 苗期 N0 7.11±0.12 d 2.37±0.06 d 33.30±0.62 b - N1 7.60±0.22 c 2.63±0.09 c 34.65±1.51 ab - N2 11.28±0.17 a 4.00±0.13 a 35.43±1.75 ab - N3 8.85±0.16 b 3.26±0.17 b 36.86±2.50 a - 花期 N0 10.32±0.17 c 2.90±0.07 d 28.14±1.13 b - N1 10.52±0.29 c 3.64±0.22 c 35.24±1.19 a - N2 20.49±0.98 a 6.82±0.13 a 33.32±1.00 a - N3 16.75±0.78 b 5.98±0.14 b 35.78±2.53 a - 收获期 N0 11.72 ±0.59 d 3.20±0.20 d 27.31±0.83 c 64.00±3.91 c N1 16.67±0.69 c 5.49±0.27 c 32.93±0.47 b 109.89±5.88 b N2 21.83±1.02 a 8.26±0.27 a 37.93±3.02 a 165.20±4.69 a N3 18.25±0.48 b 7.22±0.18 b 39.57±0.08 a 144.38±4.00 a 说明:-表示无此项。数值为平均值±标准差(n=3)。不同小写字母表示同一生育期不同氮肥施用量间差异显著(P<0.05)。 -
由图1可知:随氮肥用量增加,苗期、花期及收获期湖北贝母鳞茎中氮质量分数较N0处理的增幅分别为2.49~3.28、1.13~2.59、0.52~1.34mg·g−1,均在N2处理时达到峰值,且贝母鳞茎中氮质量分数与其干质量显著正相关(P<0.01)。随氮肥用量增加,各生育期内贝母鳞茎中磷质量分数与N0处理相比总体呈降低趋势,且鳞茎中磷质量分数与其干质量显著负相关(P<0.05)。各生育期内湖北贝母鳞茎中钾质量分数均在N1处理时达到峰值。苗期时,N2、N3处理中贝母鳞茎钾质量分数均显著高于N0处理(P<0.05);花期和收获期,N2、N3处理中贝母鳞茎钾质量分数均小于N0处理,且均在N2处理时最小。相关性分析表明:湖北贝母鳞茎中氮与钾的质量分数显著正相关(P<0.05)。相同氮肥施用量时,湖北贝母鳞茎中氮、钾质量分数均随生育期延长逐渐增加,但磷质量分数均为苗期时最高,花期时最低。
图 1 不同氮肥施用量对湖北贝母鳞茎氮、磷、钾质量分数的影响
Figure 1. Effects of different nitrogen dosages on nitrogen, phosphorus and potassium in F. hupehensis bulbs under different nitrogen
由图2可知:在同一生育期内,随氮肥用量增加,湖北贝母鳞茎中氮、磷、钾的积累量显著增加(P<0.05),且均在N2处理时达到峰值。相关性分析表明:单株湖北贝母鳞茎干质量与氮、磷、钾积累量均显著正相关(P<0.05),相关系数分别为0.997、0.890、0.988。相同氮肥施用量时,苗期至花期,湖北贝母鳞茎中氮、钾积累量分别增长11.05~46.87、12.95~37.47 mg·株−1,增长率分别为44.86%~85.72%、59.93%~113.55%;花期至收获期,湖北贝母鳞茎中氮、钾积累量分别增长8.01~22.27、5.88~25.44 mg·株−1,增长率分别为21.93%~52.22%、17.01%~56.97%。总体而言,4种氮肥用量处理时苗期至花期湖北贝母鳞茎中氮、钾的积累量及增长率均高于花期至收获期,此变化趋势与湖北贝母鳞茎干质量的增长规律相同。这一结果表明:苗期至花期是湖北贝母增产的关键时期,重施氮、钾肥对于鳞茎生长至关重要。
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表2为不同氮肥用量时各生育期湖北贝母鳞茎中氮、磷、钾的化学计量比。总体而言,贝母鳞茎的氮磷比均小于14.0,苗期湖北贝母鳞茎中钾磷比小于3.4,花期及收获期鳞茎中钾磷比均大于3.4。在同一生育期内,随氮肥用量增加,湖北贝母鳞茎的氮磷比、钾磷比呈增加趋势。相同氮肥用量处理下,苗期至花期湖北贝母鳞茎的氮磷比、钾磷比均增加,花期至收获期湖北贝母鳞茎的氮磷比、钾磷比均呈降低趋势,且降幅减小。此结果可能与花期湖北贝母鳞茎中磷质量分数最低有关。在湖北贝母整个生育期内,鳞茎中氮钾比均小于2.1;在同一生育期内,随氮肥用量增加,湖北贝母鳞茎的氮钾比均先增加后降低,且在N2处理时达到峰值;相同氮肥用量处理下,随生育期延长,湖北贝母鳞茎中氮钾比总体呈降低趋势,说明湖北贝母由营养生长向生殖生长过渡时,鳞茎对钾的需求不断增强,可适量增施钾肥。
表 2 不同氮肥用量时湖北贝母鳞茎氮、磷、钾的化学计量比
Table 2. The stoichiometric ratio of nitrogen, phosphorous and potassium in F. hupehensis bulbs under different nitrogen dosages
时期 处理 氮磷比 氮钾比 钾磷比 时期 处理 氮磷比 氮钾比 钾磷比 苗期 N0 2.58±0.07 d 1.14±0.03 d 2.27±0.03 c 花期 N2 4.79±0.08 b 1.32±0.02 a 3.63±0.05 c N1 3.29±0.10 c 1.23±0.03 c 2.68±0.04 b N3 6.31±0.17 a 1.14±0.02 b 5.53±0.22 a N2 3.75±0.07 b 1.38±0.02 a 2.71±0.02 b N3 3.94±0.09 a 1.31±0.01 b 3.01±0.07 a 收获期 N0 4.05±0.07 c 1.08±0.01 c 3.75±0.08 bc N1 4.23±0.06 c 1.11±0.01 c 3.81±0.08 b 花期 N0 3.70±0.02 c 1.00±0.01 c 3.59±0.03 c N2 4.66±0.02 b 1.30±0.03 a 3.57±0.09 c N1 4.86±0.10 b 1.14±0.01 b 4.25±0.08 b N3 6.30±0.17 a 1.16±0.01 b 5.42±0.20 a 说明:不同小写字母表示同一生育期不同氮肥施用量间差异显著(P<0.05)。 -
由图3可知:湖北贝母鳞茎中各生物碱质量分数由高到低依次为贝母素乙、贝母素甲、贝母辛、湖贝甲素。在湖北贝母各生育期内,鳞茎中总生物碱质量分数均随氮肥用量增加而降低,但4种生物碱质量分数的变化各异。随氮肥用量增加,苗期及收获期湖北贝母鳞茎中贝母素甲、贝母素乙、湖贝甲素质量分数均降低,花期鳞茎中贝母素甲、贝母素乙质量分数呈增加趋势。苗期及花期,鳞茎中贝母辛质量分数随氮肥用量增加而增加,收获期贝母辛质量分数随氮肥用量增加而降低。相同氮肥施用量时,随生育期延长,湖北贝母鳞茎中总生物碱、贝母素甲、贝母素乙、湖贝甲素质量分数从高到低依次均为收获期、苗期、花期,贝母辛质量分数为苗期较高,花期最低。
图 3 不同氮肥施用量对湖北贝母鳞茎生物碱的影响
Figure 3. Effects of different nitrogen dosages on concentration of alkaloids in F. hupehensis bulbs
由图4可知:在同一生育期内,随氮肥用量增加,贝母鳞茎中总生物碱、贝母素甲、贝母素乙、湖贝甲素、贝母辛的积累量显著增加(P<0.05),且均在N2处理时达到峰值。相关分析表明:单株湖北贝母鳞茎干质量与上述5种生物碱的积累量均显著正相关(P<0.05),相关系数分别为0.813、0.925、0.943、0.931、0.959。湖北贝母鳞茎中氮、磷、钾的积累量与上述5种生物碱的积累量也均显著正相关(P<0.05)。鳞茎中氮积累量与上述5种生物碱积累量的相关系数分别为0.825、0.925、0.945、0.931、0.964,磷积累量与上述5种生物碱积累量的相关系数分别为0.833、0.813、0.855、0.867、0.897,钾积累量与上述5种生物碱积累量的相关系数分别为0.844、0.944、0.949、0.936、0.930。氮肥用量相同时,苗期至花期鳞茎中总生物碱积累量仅增长0.47~9.96 mg·株−1,花期至收获期鳞茎中总生物碱积累量增长18.70~25.62 mg·株−1。此结果表明:花期至收获期为湖北贝母鳞茎总生物碱的快速积累时期。当氮施用量小于116.0 kg·hm−2(N2)时,花期至收获期鳞茎中贝母素甲、贝母素乙、湖贝甲素、贝母辛积累量的增幅高于苗期至花期的变化;当氮肥用量达到N2处理水平时,花期至收获期鳞茎中贝母素甲、贝母素乙、贝母辛积累量的增幅低于苗期至花期的变化。这种差异表明:氮肥用量对湖北贝母鳞茎中各生物碱的积累至关重要,过量施用氮肥不利于上述生物碱的合成。
Effects of nitrogen application on nitrogen, phosphorus and potassium absorption and alkaloid accumulation in Fritillaria hupehensis
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摘要:
目的 分析湖北贝母Fritillaria hupehensis鳞茎中氮、磷、钾质量分数及积累量的年变化规律,探讨不同氮肥用量对湖北贝母干物质及各生物碱积累的影响,为湖北贝母合理施肥提供科学依据。 方法 以湖北省恩施市新塘乡境内种植的湖北贝母为材料,在磷肥(五氧化二磷)、钾肥(氧化钾)用量均为102.4和86.4 kg·hm−2条件下,设置4个氮肥处理:不施氮肥(N0)、氮肥用量为58.0 kg·hm−2(N1)、氮肥用量为116.0 kg·hm−2(N2)、氮肥用量为174.0 kg·hm−2(N3),分别于苗期、花期、收获期取样,测定不同生育期湖北贝母鳞茎中氮、磷、钾及各生物碱质量分数。 结果 随氮肥用量增加,单株湖北贝母鳞茎的干物质,鳞茎中氮、磷、钾及总生物碱,贝母素甲,贝母素乙,湖贝甲素,贝母辛的积累量显著增加(P<0.05),且均在氮肥用量为116.0 kg·hm-2时达到峰值。随氮肥用量增加,湖北贝母苗期鳞茎中钾磷比小于3.4,花期及收获期贝母鳞茎中钾磷比均大于3.4;整个生育期内湖北贝母鳞茎中氮磷比小于14.0,氮钾比小于2.1。苗期及收获期,湖北贝母鳞茎中总生物碱、贝母素甲、贝母素乙、湖贝甲素质量分数均随氮肥用量增加而降低;花期湖北贝母鳞茎中贝母素甲、贝母素乙、贝母辛质量分数随氮肥用量增加呈上升趋势。氮肥用量为116.0 kg·hm−2时,湖北贝母鳞茎中总生物碱、贝母素甲、贝母素乙、湖贝甲素、贝母辛积累量的峰值分别为56.50、4.20、12.73、1.66、3.68 mg·株−1。相同氮肥用量时,随生育期延长,除贝母辛外,其余各生物碱质量分数均为收获期最高,花期最低。氮肥用量为116.0 kg·hm−2时,花期至收获期鳞茎中贝母素甲、贝母素乙、贝母辛的积累速率降低。苗期至花期,湖北贝母鳞茎的干物质积累速率较快,花期至收获期鳞茎中总生物碱的积累能力较强。 结论 氮肥用量对湖北贝母产量及品质影响显著,过量施用氮肥不利于鳞茎干物质及各生物碱的积累。湖北贝母在不同物候期鳞茎干物质及生物碱的积累是动态的过程,营养生长期以产量提升为主,生殖生长阶段以生物碱积累为主。图4表2参28 Abstract:Objective This study aims to analyze the annual variation of the concentration and accumulation of nitrogen, phosphorus, and potassium in Fritillaria hupehensis bulbs, and explore the impacts of different nitrogen dosages on the accumulation of dry matter and alkaloids of F. hupehensis, so as to provide scientific basis for rational fertilization of F. hupehensis. Method F. hupehensis obtained from Xintang township, Enshi City, Hubei Province was used as the material. Under the conditions of 102.4 kg·hm−2 phosphorus pentoxide and 86.4 kg·hm−2 potassium oxide for both phosphorus and potassium fertilizers, 4 nitrogen fertilizer treatments were set up: no nitrogen fertilizer (N0), 58.0 kg·hm−2 nitrogen fertilizer (N1), 116.0 kg·hm−2 nitrogen fertilizer (N2), and 174.0 kg·hm−2 nitrogen fertilizer (N3). Samples were collected at seedling, flowering and harvest stages to determine the concentration of nitrogen, phosphorus, potassium and various alkaloids in the bulbs of F. hupehensis at different growth stages. Result The accumulation of dry matter, nitrogen, phosphorus, potassium, total alkaloid, verticine, verticinone, peimissine and hupehenine in F. hupehensis bulbs significantly increased with increasing nitrogen application (P<0.05), and all reached their peak at a nitrogen fertilizer application of 116.0 kg·hm−2. With the increase of nitrogen application, potassium to phosphorus ratio in F. hupehensis bulbs was less than 3.4 at seedling stage, while that during the flowering and harvesting periods was more than 3.4. Throughout the entire growth period, nitrogen to phosphorus ratio in F. hupehensis bulbs was less than 14.0, and nitrogen to potassium ratio was less than 2.1. During the seedling and harvest stages, the concentration of total alkaloid, verticine, verticinone, and hupehenine in F. hupehensis bulbs all decreased with increasing nitrogen application. During the flowering period, the concentration of verticine, verticinone and peimissine in F. hupehensis bulbs showed an increasing trend with the increase in nitrogen application. When the nitrogen fertilizer application amount was 116.0 kg·hm−2, the peak accumulation of total alkaloid, verticine, verticinone, hupehenine and peimissine in F. hupehensis bulbs were 56.50, 4.20, 12.73, 1.66, and 3.68 mg·plant−1, respectively. When the same amount of nitrogen fertilizer was applied, with the extension of growth period, except for peimissine, the concentration of all other alkaloids was the highest during the harvest period and the lowest during the flowering period. When the nitrogen fertilizer application amount was 116.0 kg·hm−2, the accumulation rate of verticine, verticinone and peimissine in bulbs decreased from flowering to harvest stage. The dry matter accumulation rate was relatively fast from seedling to flowering stage, and the accumulation ability of total alkaloids in the bulb during the flowering to harvest period was strong. Conclusion The amount of nitrogen fertilizer significantly affects the yield and quality of F. hupehensis. Excessive application of nitrogen fertilizer is not conducive to the accumulation of dry matter and alkaloids in bulbs. The accumulation of dry matter and alkaloids in bulbs of F. hupehensis at different growth stages is a dynamic process. Yield increase is predominant at vegetative growth stage, while alkaloids accumulation is dominant at productive growth stage. [Ch, 4 fig. 2 tab. 28 ref.] -
表 1 不同氮肥施用量时湖北贝母的生长指标
Table 1. Growth indices of F. hupehensis under different nitrogen dosages
生育期 处理 叶长/cm 叶宽/cm 叶型指数 株高/cm 茎粗/cm 苗期 N0 11.15±0.12 b 1.03±0.06 c 10.88±0.56 a 19.70±0.92 c 0.50±0.01 b N1 11.33±0.12 b 1.33±0.05 b 8.55±0.29 b 29.30±1.00 b 0.53±0.02 ab N2 11.45±0.19 a 1.55±0.10 a 7.39±0.25 d 33.90±3.11 a 0.55±0.03 a N3 11.00±0.19 b 1.42±0.06 b 7.76±0.21 c 32.85±0.95 a 0.50±0.02 b 花期 N0 11.77±0.10 b 1.72±0.10 b 6.86±0.48 a 36.45±0.58 b 0.50±0.02 b N1 11.90±0.07 b 1.80±0.11 a 6.61±0.39 a 38.00±0.94 b 0.55±0.02 b N2 14.20±0.08 a 2.10±0.10 b 6.76±0.51 a 40.50±1.16 a 0.65±0.03 a N3 12.00±0.20 b 1.70±0.12 b 7.06±0.61 a 37.60±2.02 b 0.51±0.04 b 收获期 N0 - - - - - N1 - - - - - N2 - - - - - N3 - - - - - 生育期 处理 鳞茎鲜质量/(g·株−1) 鳞茎干质量/(g·株−1) 折干率/% 鳞茎干质量产量/(g·m−2) 苗期 N0 7.11±0.12 d 2.37±0.06 d 33.30±0.62 b - N1 7.60±0.22 c 2.63±0.09 c 34.65±1.51 ab - N2 11.28±0.17 a 4.00±0.13 a 35.43±1.75 ab - N3 8.85±0.16 b 3.26±0.17 b 36.86±2.50 a - 花期 N0 10.32±0.17 c 2.90±0.07 d 28.14±1.13 b - N1 10.52±0.29 c 3.64±0.22 c 35.24±1.19 a - N2 20.49±0.98 a 6.82±0.13 a 33.32±1.00 a - N3 16.75±0.78 b 5.98±0.14 b 35.78±2.53 a - 收获期 N0 11.72 ±0.59 d 3.20±0.20 d 27.31±0.83 c 64.00±3.91 c N1 16.67±0.69 c 5.49±0.27 c 32.93±0.47 b 109.89±5.88 b N2 21.83±1.02 a 8.26±0.27 a 37.93±3.02 a 165.20±4.69 a N3 18.25±0.48 b 7.22±0.18 b 39.57±0.08 a 144.38±4.00 a 说明:-表示无此项。数值为平均值±标准差(n=3)。不同小写字母表示同一生育期不同氮肥施用量间差异显著(P<0.05)。 表 2 不同氮肥用量时湖北贝母鳞茎氮、磷、钾的化学计量比
Table 2. The stoichiometric ratio of nitrogen, phosphorous and potassium in F. hupehensis bulbs under different nitrogen dosages
时期 处理 氮磷比 氮钾比 钾磷比 时期 处理 氮磷比 氮钾比 钾磷比 苗期 N0 2.58±0.07 d 1.14±0.03 d 2.27±0.03 c 花期 N2 4.79±0.08 b 1.32±0.02 a 3.63±0.05 c N1 3.29±0.10 c 1.23±0.03 c 2.68±0.04 b N3 6.31±0.17 a 1.14±0.02 b 5.53±0.22 a N2 3.75±0.07 b 1.38±0.02 a 2.71±0.02 b N3 3.94±0.09 a 1.31±0.01 b 3.01±0.07 a 收获期 N0 4.05±0.07 c 1.08±0.01 c 3.75±0.08 bc N1 4.23±0.06 c 1.11±0.01 c 3.81±0.08 b 花期 N0 3.70±0.02 c 1.00±0.01 c 3.59±0.03 c N2 4.66±0.02 b 1.30±0.03 a 3.57±0.09 c N1 4.86±0.10 b 1.14±0.01 b 4.25±0.08 b N3 6.30±0.17 a 1.16±0.01 b 5.42±0.20 a 说明:不同小写字母表示同一生育期不同氮肥施用量间差异显著(P<0.05)。 -
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