川半夏规范化种植技术和对不同温度的生理响应研究
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摘要
半夏(Pinellia ternata (Thunb.) Breit)为天南星科(Araceae)半夏属(Pinellia)多年生草本植物,具有燥湿化痰、除逆止呕、消痞散结、外用消肿止痛等功效。半夏是一味非常重要的中药,不但在我国新药、特药、中成药和方药饮片中有大量使用,同时还出口韩国、日本、东南亚等各国为汉方用药。半夏植物生长习性特殊,忌涝怕旱、耐阴惧晒,易“倒苗”,人工栽培规范化生产技术不完善,产量不高,同时野生资源破坏严重,市场缺口较大。四川为半夏主产区之一,其中以南充、武胜、安岳、达州等地产量大,质量较优。为保护川半夏野生资源和满足市场对川半夏的需求,本文从川半夏的规范化生产技术及其对温度的生理响应两方面进行了研究,研究结果如下：
1.采用9种不同贮藏方式对川半夏种茎进行越冬贮藏处理,结果表明川半夏在不同贮藏条件下,除叶绿素含量差异未达显著水平外(P>0.05),发芽率、发芽势、发芽指数、发根数、根长、根系活力、过氧化氢酶活力的差异均达显著水平(P<0.05)。川半夏种茎收获后,采用室内湿沙(土)贮藏或3℃(±1)冰箱贮藏,种茎的萌发和生长活力表现最好,为种茎适宜的贮藏方式。
2.将来自川半夏主产区的3份不同材料的种茎分别按大(d>1.5cm)、中(1.01.5cm者一般不适宜用做种茎,但良种繁育时可使用,此时珠芽产生量大,繁育系数高。
3.随机选取300粒川半夏种茎,单株考察其生育期内主要农艺性状,收获后计算其单株增重率和单株增殖率,进行K-模糊聚类分析。结果表明188粒有效种茎可分为3类,第Ⅰ类种茎平均直径为1.27cm,单株增重率为134.09%；第11类平均直径为1.40cm,单株增重率仅13.83%;第Ⅲ类平均直径为1.02cm,单株增重率为369.98%。建议生产中优先选择直径0.7-1.2cm(平均值1.Ocm)的块茎作为种茎,其次选择直径1.2-1.55cm者,直径1.55cm以上的一般不作为种茎使用。
4.分析了2份主产区川半夏材料地上茎叶和地下块茎不同生育时期氮、磷、钾含量变化情况。结果表明,川半夏不同生育时期各部分对氮、磷、钾的吸收量不同,其中对钾的吸收量最大,对磷的吸收量最少；在半夏大面积“倒苗”前后,各部分氮、磷和钾含量以及累积总量总体上高于非“倒苗”期。生长期间半夏地上和地下部分磷、钾吸收量间相关系数均分别达显著水平(P<0.05),且地下部分磷和氮吸收量间相关系数也达到显著水平(P<0.05)。半夏生长过程中,每生产100kg干品,大约需要吸收氮5.02-5.64kg,P2O52.44-2.56kg,K2011.62-13.49kg。川半夏在大田栽培中应当施足底肥,并在6月中旬和8月下旬前后及时追施氮、磷和钾肥,并适当提高钾的比例。追肥方式可采用地下浇施和叶面喷施相结合的方式,川半夏地下部分对磷肥和氮肥的吸收有相互促进效应。
5.在对川半夏氮磷钾营养元素吸收累积动态研究的基础上进一步探讨了不同施肥水平与其生长节律、主要农艺性状、产量和品质间的关系。结果表明,不同施肥水平下川半夏出苗、珠芽产生和抽薹等生长节律基本一致。生长期间平均出苗数间差异达显著水平(P<0.05),平均抽薹数、平均珠芽产生数、净产量、单株增殖率、β-谷甾醇以及生物碱含量间差异则达极显著水平(P<0.01)。氮肥对川半夏生物碱和β-谷甾酣含量均有极显著效应(P<0.01),对上述其他指标则无显著效应。磷肥和钾肥对各指标均无显著效应。川半夏生产栽培中最佳施肥组合为每亩施N、P2O5和K20分别为21kg、21kg和30kg或21kg、15kg和18kg。
6.本研究探讨了不同温度下川半夏叶片中叶绿素、脯氨酸、保护酶、丙二醛和膜质脂肪酸的变化规律,以及对根系活力和生物碱含量的影响。结果表明,在20-30℃下,川半夏不饱和脂肪酸(IUFA)、饱和脂肪酸／不饱和脂肪酸比值(SFA/USFA)、叶绿素a含量(Chla)、叶绿素b含量(Chlb)、总叶绿素含量(Chit)、过氧化氢酶活性(CAT)、过氧化物酶活性(POD)、超氧化歧化酶活性(SOD)、丙二醛含量(MDA)、脯氨酸含量变化规律和变幅基本一致,变幅相比35-40℃处理后较小,各指标含量更趋于对照,较为稳定。在35-40℃温度处理下,前3天内IUFA下降,SFA/USFA比值上升,Chla、Chlb、Chlt含量下降,CAT、POD、SOD酶活性上升,MDA和脯氨酸含量上升,这些高温胁迫的生理特征均得以迅速表现。但随着胁迫时间的延长各指标并非继续沿着一个方向持续变化,而是会逐步接近或回复到20-30℃水平。据此认为,川半夏最适生长温度为20-30℃,胁迫温度在30-35℃之间,而35-40℃下川半夏则会迅速“倒苗”。高温胁迫下川半夏会出现一系列的生理响应,这些响应指标间也存在密切的联系,共同促进了川半夏对高温胁迫的适应,但长期的高温胁迫会对其造成根系损伤,并最终导致植株“倒苗”,进而对川半夏生物碱的含量也有一定的影响。
Pinellia ternata (Thunb.) Breit is a perennial herb belongs to Pinellia Tenore of Araceae,which has the function of dry wet, convert phlegm, lowering athwart and stopping vomit, removing ruffian and dispeling vinculum.It is important as a part of Chinese traditional medicine or used as herbal pieces, and has an increasing demand for exporting to Korea, Japan and Southeast Asia. As a shade tolerant plant, P. ternata has a special growth habit, cannot grow well with drought and flood, and the part of the ground easily withered, which are commonly known as "sprout tumble". Sichuan is one of the main producing regions of P. ternata, which has the higher yield and quality in the regions of Nanchong, Wusheng, Anyue and Dazhou. It is in need to improve the cultivation technology for a higher yield and a better protection of the wild resource. Here we researched the standardized cultivation technology in P. ternata production as well as the physiological responses under different growth temperatures, in order to satisfy the market demand and protect the wild resource. The main results are summarized as follows:
1. Using nine different storage methods to storage the tubles of P. ternata from Sichuan in winter, the results showed that all the indexes including the germination percentage,vigor of germination,germination index,rooting number,root length,root activity and catalase activity were significantly different among different storage methods (P<0.05), except for chlorophyll content(P>0.05). So after harvesting, the tubers should be preserved in icebox (3±1℃) or the wet sand in a greenhouse. Under these conditions, the germinating vigor and growth activity of tuber performed best, which are the optimal storage methods.
2. The tubers of three different populations of P. ternata from Sichuan were classified respectively as three different groups according to the diameters as follows: smaller group(0.5cm1.5cm), in order to explore the effect of the size of tuber on the yield and quality of P. ternata. The results showed that they all had the same growth rhythm. And the size of tubers with0.5cm1.5cm) while good strains breeding and propagation.
3. Randomly selected300tubles of Pinellia ternata from Sichuan, examine main agronomic traits to per plant yield during the growing stages, and Calculation individual weight gain rate and proliferation rate of each plant. The results of fuzzy cluster analysis showed that the188effective tubers could be classified as three different groups. The average tuber diameter of group Ⅰ was1.27cm, with growth rate per plant of134.09%, the average tuber diameter of group Ⅱ was1.40cm with growth rate per plant only13.83%, while the average tuber diameter of group III was1.02cm with growth rate per plant up to369.98%. The tubers which diameter between0.7-1.2cm (average1.0cm) should be prefer to be selected for production, followed by diameter between1.2-1.55cm. The tubers with diameter more than1.55cm were suggested not be used.
4. The contents of nitrogen, phosphorus and potassium of the stems, leaves and tubers of different growth stages of two populations of P. ternate from Sichuan were determined. The results showed that the absorption rate of aerial and underground parts of P. ternate were different in nitrogen, phosphorus, and potassium at different growing stages. Among them, the potassium uptake is the most, while the phosphorus uptake is the least. Totally, the uptakes of nitrogen, phosphorus and potassium of P. ternate from Sichuan during "sprout tumble" stage were comparatively higher than that not at "sprout tumble"stage. A significant correlation was observed between the phosphorus and potassium uptake of the aerial and underground parts, respectively (P <0.05). And there was also significant positive correlation between the nitrogen and phosphorus uptake at underground parts (P<0.05). During the growth stage of P. ternate, as producing100kg air-dried P. ternate,5.02-5.64kg N,2.44-2.56kg P2O5,11.62-13.49kg K2O need to be absorbed. P. ternate from Sichuan should be given enough base fertilizer in field cultivation, and then be supplied with phosphorus and potassium by combining foliar spray with field application during the middle ten days of June and last ten days of August. The ratios of potassium should be properly improved. There was promoting effect in the phosphorus and nitrogen assimilations of the underground parts of P. ternate from Sichuan.
5. Based on the absorption and accumulation of the dynamic study of nitrogen, phosphorus and potassium, the relationship between the level of fertilization and growth rhythm, major agronomic traits, yield and quality were explored. The results showed that, the P. ternata from Sichuan showed the same growth rhythm at different fertilizing levels, but there were significant difference (P<0.05)in the average emergence rate, and extremely significant difference(P<0.01) were detected in the average ratio of bolting, the average bulbils, individual growth rate, individual proliferation rate,β-sitosterol and alkaloid content under different fertilizing levels. Nitrogenous fertilizer had extremely significant effects on the content of alkaloids and β-sitosterol (P<0.01), whereas phosphata and potassium fertilizer had no significant effect. The effect of fertilizer factor and intereffects on β-sitosterol showed no significant effect, but have influence on other indexes.
6. The content of chlorophyll, proline, protective enzymes, malondialdehyde, membranous fatty acids of P. ternate from Sichuan were consecutively determined under different temperature per24h, the root activity and alkaloid were determined finally. The results showed that the variation and amplitude of that the index of unsaturated fatty acids (IUFA) and the ratio of saturated fatty acids and unsaturated fatty acids (SFA/USFA) in20-30℃are basically the same, and similar to the control, relatively stable. The same results also showed in the content of chlorophyll a (Chla), chlorophyll b (Chlb), total chlorophyll (Chit), proline, malonyldiadehyde (MDA), and the activity of catalase(CAT), peroxidase(POD) and superoxide dismutase(SOD). Under the temperture of35-40℃, the IUFA decreased while SFA/USFA increased. The contents of Chla, Chlb, Chit decreased. The activities of CAT, POD and SOD, and the content of MDA and proline increased at the first3days. These physiological characteristics under high temperature stress exhibited rapidly. But these indexs did not continue to follow a continuous change direction under the continuous high temperature stress, which will be close or return to the appropriate level. It was considered that the optimum growth temperature for P. ternate from Sichuan was between20-30℃, the stress temperature was between30-35℃.Under the stress temperature it would appeared a series of biochemical and physiological properties even "sprout tumble"quickly. These response indicators are closely linked, and jointly promote P. ternate to adapt to the growing conditions, but long-term heat stress would cause root damage and eventually lead to plants "sprout tumble". Furthermore, have a major impact on the accumulation of main chemical compositions of P. ternate from Sichuan.
1.采用9种不同贮藏方式对川半夏种茎进行越冬贮藏处理,结果表明川半夏在不同贮藏条件下,除叶绿素含量差异未达显著水平外(P>0.05),发芽率、发芽势、发芽指数、发根数、根长、根系活力、过氧化氢酶活力的差异均达显著水平(P<0.05)。川半夏种茎收获后,采用室内湿沙(土)贮藏或3℃(±1)冰箱贮藏,种茎的萌发和生长活力表现最好,为种茎适宜的贮藏方式。
2.将来自川半夏主产区的3份不同材料的种茎分别按大(d>1.5cm)、中(1.0
3.随机选取300粒川半夏种茎,单株考察其生育期内主要农艺性状,收获后计算其单株增重率和单株增殖率,进行K-模糊聚类分析。结果表明188粒有效种茎可分为3类,第Ⅰ类种茎平均直径为1.27cm,单株增重率为134.09%；第11类平均直径为1.40cm,单株增重率仅13.83%;第Ⅲ类平均直径为1.02cm,单株增重率为369.98%。建议生产中优先选择直径0.7-1.2cm(平均值1.Ocm)的块茎作为种茎,其次选择直径1.2-1.55cm者,直径1.55cm以上的一般不作为种茎使用。
4.分析了2份主产区川半夏材料地上茎叶和地下块茎不同生育时期氮、磷、钾含量变化情况。结果表明,川半夏不同生育时期各部分对氮、磷、钾的吸收量不同,其中对钾的吸收量最大,对磷的吸收量最少；在半夏大面积“倒苗”前后,各部分氮、磷和钾含量以及累积总量总体上高于非“倒苗”期。生长期间半夏地上和地下部分磷、钾吸收量间相关系数均分别达显著水平(P<0.05),且地下部分磷和氮吸收量间相关系数也达到显著水平(P<0.05)。半夏生长过程中,每生产100kg干品,大约需要吸收氮5.02-5.64kg,P2O52.44-2.56kg,K2011.62-13.49kg。川半夏在大田栽培中应当施足底肥,并在6月中旬和8月下旬前后及时追施氮、磷和钾肥,并适当提高钾的比例。追肥方式可采用地下浇施和叶面喷施相结合的方式,川半夏地下部分对磷肥和氮肥的吸收有相互促进效应。
5.在对川半夏氮磷钾营养元素吸收累积动态研究的基础上进一步探讨了不同施肥水平与其生长节律、主要农艺性状、产量和品质间的关系。结果表明,不同施肥水平下川半夏出苗、珠芽产生和抽薹等生长节律基本一致。生长期间平均出苗数间差异达显著水平(P<0.05),平均抽薹数、平均珠芽产生数、净产量、单株增殖率、β-谷甾醇以及生物碱含量间差异则达极显著水平(P<0.01)。氮肥对川半夏生物碱和β-谷甾酣含量均有极显著效应(P<0.01),对上述其他指标则无显著效应。磷肥和钾肥对各指标均无显著效应。川半夏生产栽培中最佳施肥组合为每亩施N、P2O5和K20分别为21kg、21kg和30kg或21kg、15kg和18kg。
6.本研究探讨了不同温度下川半夏叶片中叶绿素、脯氨酸、保护酶、丙二醛和膜质脂肪酸的变化规律,以及对根系活力和生物碱含量的影响。结果表明,在20-30℃下,川半夏不饱和脂肪酸(IUFA)、饱和脂肪酸／不饱和脂肪酸比值(SFA/USFA)、叶绿素a含量(Chla)、叶绿素b含量(Chlb)、总叶绿素含量(Chit)、过氧化氢酶活性(CAT)、过氧化物酶活性(POD)、超氧化歧化酶活性(SOD)、丙二醛含量(MDA)、脯氨酸含量变化规律和变幅基本一致,变幅相比35-40℃处理后较小,各指标含量更趋于对照,较为稳定。在35-40℃温度处理下,前3天内IUFA下降,SFA/USFA比值上升,Chla、Chlb、Chlt含量下降,CAT、POD、SOD酶活性上升,MDA和脯氨酸含量上升,这些高温胁迫的生理特征均得以迅速表现。但随着胁迫时间的延长各指标并非继续沿着一个方向持续变化,而是会逐步接近或回复到20-30℃水平。据此认为,川半夏最适生长温度为20-30℃,胁迫温度在30-35℃之间,而35-40℃下川半夏则会迅速“倒苗”。高温胁迫下川半夏会出现一系列的生理响应,这些响应指标间也存在密切的联系,共同促进了川半夏对高温胁迫的适应,但长期的高温胁迫会对其造成根系损伤,并最终导致植株“倒苗”,进而对川半夏生物碱的含量也有一定的影响。
Pinellia ternata (Thunb.) Breit is a perennial herb belongs to Pinellia Tenore of Araceae,which has the function of dry wet, convert phlegm, lowering athwart and stopping vomit, removing ruffian and dispeling vinculum.It is important as a part of Chinese traditional medicine or used as herbal pieces, and has an increasing demand for exporting to Korea, Japan and Southeast Asia. As a shade tolerant plant, P. ternata has a special growth habit, cannot grow well with drought and flood, and the part of the ground easily withered, which are commonly known as "sprout tumble". Sichuan is one of the main producing regions of P. ternata, which has the higher yield and quality in the regions of Nanchong, Wusheng, Anyue and Dazhou. It is in need to improve the cultivation technology for a higher yield and a better protection of the wild resource. Here we researched the standardized cultivation technology in P. ternata production as well as the physiological responses under different growth temperatures, in order to satisfy the market demand and protect the wild resource. The main results are summarized as follows:
1. Using nine different storage methods to storage the tubles of P. ternata from Sichuan in winter, the results showed that all the indexes including the germination percentage,vigor of germination,germination index,rooting number,root length,root activity and catalase activity were significantly different among different storage methods (P<0.05), except for chlorophyll content(P>0.05). So after harvesting, the tubers should be preserved in icebox (3±1℃) or the wet sand in a greenhouse. Under these conditions, the germinating vigor and growth activity of tuber performed best, which are the optimal storage methods.
2. The tubers of three different populations of P. ternata from Sichuan were classified respectively as three different groups according to the diameters as follows: smaller group(0.5cm
3. Randomly selected300tubles of Pinellia ternata from Sichuan, examine main agronomic traits to per plant yield during the growing stages, and Calculation individual weight gain rate and proliferation rate of each plant. The results of fuzzy cluster analysis showed that the188effective tubers could be classified as three different groups. The average tuber diameter of group Ⅰ was1.27cm, with growth rate per plant of134.09%, the average tuber diameter of group Ⅱ was1.40cm with growth rate per plant only13.83%, while the average tuber diameter of group III was1.02cm with growth rate per plant up to369.98%. The tubers which diameter between0.7-1.2cm (average1.0cm) should be prefer to be selected for production, followed by diameter between1.2-1.55cm. The tubers with diameter more than1.55cm were suggested not be used.
4. The contents of nitrogen, phosphorus and potassium of the stems, leaves and tubers of different growth stages of two populations of P. ternate from Sichuan were determined. The results showed that the absorption rate of aerial and underground parts of P. ternate were different in nitrogen, phosphorus, and potassium at different growing stages. Among them, the potassium uptake is the most, while the phosphorus uptake is the least. Totally, the uptakes of nitrogen, phosphorus and potassium of P. ternate from Sichuan during "sprout tumble" stage were comparatively higher than that not at "sprout tumble"stage. A significant correlation was observed between the phosphorus and potassium uptake of the aerial and underground parts, respectively (P <0.05). And there was also significant positive correlation between the nitrogen and phosphorus uptake at underground parts (P<0.05). During the growth stage of P. ternate, as producing100kg air-dried P. ternate,5.02-5.64kg N,2.44-2.56kg P2O5,11.62-13.49kg K2O need to be absorbed. P. ternate from Sichuan should be given enough base fertilizer in field cultivation, and then be supplied with phosphorus and potassium by combining foliar spray with field application during the middle ten days of June and last ten days of August. The ratios of potassium should be properly improved. There was promoting effect in the phosphorus and nitrogen assimilations of the underground parts of P. ternate from Sichuan.
5. Based on the absorption and accumulation of the dynamic study of nitrogen, phosphorus and potassium, the relationship between the level of fertilization and growth rhythm, major agronomic traits, yield and quality were explored. The results showed that, the P. ternata from Sichuan showed the same growth rhythm at different fertilizing levels, but there were significant difference (P<0.05)in the average emergence rate, and extremely significant difference(P<0.01) were detected in the average ratio of bolting, the average bulbils, individual growth rate, individual proliferation rate,β-sitosterol and alkaloid content under different fertilizing levels. Nitrogenous fertilizer had extremely significant effects on the content of alkaloids and β-sitosterol (P<0.01), whereas phosphata and potassium fertilizer had no significant effect. The effect of fertilizer factor and intereffects on β-sitosterol showed no significant effect, but have influence on other indexes.
6. The content of chlorophyll, proline, protective enzymes, malondialdehyde, membranous fatty acids of P. ternate from Sichuan were consecutively determined under different temperature per24h, the root activity and alkaloid were determined finally. The results showed that the variation and amplitude of that the index of unsaturated fatty acids (IUFA) and the ratio of saturated fatty acids and unsaturated fatty acids (SFA/USFA) in20-30℃are basically the same, and similar to the control, relatively stable. The same results also showed in the content of chlorophyll a (Chla), chlorophyll b (Chlb), total chlorophyll (Chit), proline, malonyldiadehyde (MDA), and the activity of catalase(CAT), peroxidase(POD) and superoxide dismutase(SOD). Under the temperture of35-40℃, the IUFA decreased while SFA/USFA increased. The contents of Chla, Chlb, Chit decreased. The activities of CAT, POD and SOD, and the content of MDA and proline increased at the first3days. These physiological characteristics under high temperature stress exhibited rapidly. But these indexs did not continue to follow a continuous change direction under the continuous high temperature stress, which will be close or return to the appropriate level. It was considered that the optimum growth temperature for P. ternate from Sichuan was between20-30℃, the stress temperature was between30-35℃.Under the stress temperature it would appeared a series of biochemical and physiological properties even "sprout tumble"quickly. These response indicators are closely linked, and jointly promote P. ternate to adapt to the growing conditions, but long-term heat stress would cause root damage and eventually lead to plants "sprout tumble". Furthermore, have a major impact on the accumulation of main chemical compositions of P. ternate from Sichuan.
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