喀斯特地区顶坛花椒培育的生理生态特性研究
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摘要
“花椒”是花椒属植物的统称,顶坛花椒Zanthoxylum planispinum var. dintanensis Yu-Lin Tu是花椒属Zanthoxylum L.植物中竹叶椒Zanthoxylum planispinum Sieb. et Zucc的一个变种,是贵州喀斯特地区特有的,生态、经济价值很高的香料植物。在贵州喀斯特典型地区的花江,顶坛花椒已种植了10多年,但只是停留在传统和习惯的种植,研究工作做得少,它在生长及生产过程中的一些问题一直在困扰着人们,如播种出苗率不高、生长势较弱(叶片发黄)、产量不高、应用途径单一等,为了解决这些问题,对项坛花椒从培育的生理生态特性方面进行研究,以揭示影响项坛花椒培育的生理生态的特性及规律,优化顶坛花椒的培育。
利用目前相关生态学和生理学的方法,主要探讨了以下问题(1)培育的生理特性方面：种子形态结构与生理特点、种子萌发生理、幼苗水分胁迫、微量元素锌对干旱胁迫的影响、施肥对生长的影响、微量元素锌对生殖生理的影响；(2)培育的生态特性方面：苗期生长规律、海拔高度对果实生长的影响、幼茎营养成份分析、挥发油研究。
主要研究结果与结论(1)培育的生理特性方面：①顶坛花椒种子富含油脂,外种皮含蜡质,内种皮为坚硬的厚角质,厚度占种皮的一半以上；顶坛花椒种子内部胚乳残存,子叶还没有完全吸收胚乳中所含全部营养；有12%的种子具双胚现象；这些因素均不利于项坛花椒种子萌发。②立秋至白露中期为顶坛花椒种子采种的最佳时期；碱水处理是理想的项坛花椒种子脱脂处理方法；项坛花椒种子宜采用撒播的播种方式；磷肥作苗床底肥对种子萌发率有极显著影响,是提高顶坛花椒种子萌发率的一项有效措施。③顶坛花椒对水分胁迫的敏感性较强；幼苗随着水分胁迫加剧,叶绿素a、叶绿素b、叶绿素总量以及类胡萝卜素含量明显下降,丙二醛含量增加,游离脯氨酸含量增加；显示出项坛花椒具有一定程度的耐旱性,这也是项坛花椒能够在喀斯特地区生长的生理特性。④当ZnSO4浓度为0.06%-0.09%,顶坛花椒花粉活力和花粉寿命均能大幅度提高；当ZnSO4浓度为0.08%时,顶坛花椒种群开花率可达到最大值；ZnSO4浓度为0.09%时,顶坛花椒种群结果率达到最大值。当项坛花椒处于5-9a时它的生殖力出现峰值；在海拔为600m左右的地段,顶坛花椒具有最大的生殖力。⑤用模糊数学隶属函数法对抗旱性进行了综合评判,适宜浓度的微量元素Zn对顶坛花椒抗旱能力具有较强的促进作用,当ZnSO1预处理浓度为0.06%～0.12%时,能加强顶坛花椒对干旱的适应性和提高抗旱生产力。⑥顶坛花椒幼苗子叶属于木兰型Magnolia Type;幼苗地上和地下部分交替生长,田间管理要适时供应水分和养分；用模式方程Y=a+bX拟合了顶坛花椒幼苗的苗高、地径、主根长和根干重生长关系,可用拟合的模式方程和苗高、地径、主根长和根干重中任一指标估测该时期另三个生长指标。(2)培育生态特性方面：①在开花期到盛花期以沼肥做底肥,能促进新梢生长；以施用20 kg·株-1增产效果最为明显。顶坛花椒的最佳混料肥料配比为mN:mP:mK= (0.3981-0.4134)：(0.2981～0.3134)：(0.2864～0.2891)。②顶坛花椒果实发育生长呈“S”型生长曲线；花椒产量与品质在试验区有随着海拔降低而明显增加的趋势,花江峡谷的中低海拔地区(海拔<850m)属于南亚热带河谷气候,是种植顶坛花椒的优势地区。③顶坛花椒幼苗期叶的生物量占整个植株生物量的55.00%；蛋白质、脂肪、碳水化合物等营养成分含量高；膳食纤维含量为10种其他常见蔬菜嫩茎、叶的2～10倍；VB2含量为所有新鲜植物食品VB2含量之最；7种必需矿质元素Ca、Mg、Fe、K、Se、Cu、Mn含量均明显高于参与比较的蔬菜,顶坛花椒幼嫩茎可作优良蔬菜。④鉴定了贵州顶坛花椒和四川茂县大红袍的挥发油化学成分,顶坛花椒有29个化合物,茂县大红袍有38个化合物；其中顶坛花椒相对含量组分占挥发油总质量的99.417%,茂县大红袍占97.68%。顶坛花椒挥发性成分中的芳樟醇(Linalool L)含量尤其高,达到茂县大红袍的4.56倍,说明顶坛花椒香味好。
本论文的创新点是：(1)从顶坛花椒的种子形态结构与生理学特性、种子萌发生理、幼苗水分胁迫和苗期生长规律方面进行研究,揭示了顶坛花椒种子萌发及苗期生长的特性、对其影响顶坛花椒出苗低、生长慢的原因进行了探索。(2)从微量元素锌对顶坛花椒生殖及水分胁迫的影响,提出了微量元素锌对顶坛花椒种群开花结果率有显著的影响；指出了用模糊数学隶属函数法对顶坛花椒抗旱性进行了综合评判,适宜浓度的微量元素Zn对顶坛花椒抗旱能力具有较强的促进作用,为顶坛花椒在培育方面使用微量元素的效果提供了理论依据。(3)从施肥对顶坛花椒的影响及海拔高度对顶坛花椒的影响,提出了施肥效果好的配方。说明了海拔变化对顶坛花椒产量影响的重要性。(4)对顶坛花椒幼茎叶营养成分的分析及果实挥发油的分析,给顶坛花椒的新用途提供了科学基础。
"prickly ash" is the general designation of Zanthoxylum L.. Zanthoxylum planispinum var. dintanensis Yu-Lin Tu is a variety of Zanthoxylum planispinum Sieb.et Zucc., and in Guizhou karst areas it is a specific perfume plant, with higher ecological and economic value. Huajiang is the typical area of Guizhou krast regions, Z. planispinum var. dintanensis has been planted more than 10 years there, but there are still some growth and production questions to puzzle the people continuously, such as low seedling emergence rate, weak growth vigor (yellow leaves), low yield, sole application ways and so on. In order to solve these problems, optimized the cultivation of Z planispinum var. dintanensis, the charictristis study of plant physiology and ecology are very important.
Using of the current methods related to ecology and physiology. In this paper, Z. planispinum var. dintanensis in karst areas was selected as the study object. Mainly has carried on the discussion from following several aspects:(1) Plant physiology characteristic:seed morphological structure and physiological characteristics, seed germination physiology, seedling in the Process of water stress, the influence of Zn under drought stress and the effect of applying fertilizer to the growth. (2) Ecological Characteristics:the grow regulation in seedling period, the effects of altitudes on fruit quality, the nutrient analysis of young stem and the studt of volatile oil.
The main results and conclusion are showed as follows:
(1) Plant physiology characteristics:①The seeds of the Z planispinum var. dintanensis contained a large amount of oil, the exopleura wax, the endotesta with hard cuticle layer, which take more than half of the total thickness of the seed coat. Endosperm remains in internal seed, cotyledons have not completely absorbed all the nutrition in the endosperm. There are 12% seeds have biovulate phenomena. All factors mentioned above are not propitious for the seed germination of Z. planispinum var. dintanensis.②The seed collecting time is better later than earlier. The late time from autumn begins to White Dew are the best time for seed collecting. Treating with aqueous alkali is the ideal method for seed derosination of Z. planispinum var. dintanensis. Compared with other regions, there is better quantity of heat in karst xerothermic valley, it is better planting methods to use broadcast for the seed of Z. planispinum var. dintanensis. Using phosphorus as base manure to the seedling bed has very obvious influence to seed germination. This is a effective method to improve the seed germination rate of Z. planispinum var. dintanensis.③With the aggravation of water stress, the chlorophyll a, chlorophyll b, Gross chlorophyll and carotenoid of Z planispinum var. dintanensis. seedlings decreased obviously, the MDA content and the POD activation increased. In the early stage of water stress, the seedlings were more damaged. The results showed that Z. planispinum var. dintanensis. has a certainty drought tolerance, it was the foundation of Z. planispinum var. dintanensis. adapting in karst mountainous country.④When ZnSO4 were between 0.06% and 0.09%, both pollen viability and life-span of Z. planispinum var. dintanensis can be boosted greatly. When ZnSO4 were between 0.08% and 0.09%, both blossoming percent and fruiting percent of Z planispinum var. dintanensis can be boosted greatly. The peak value of its reproductive age is during 5-9 years old, while at an altitude of 600m, the Z.planispinum var. dintanensis have the max fecundity.⑤The results by fuzzy comprehensive appraisal analysis indicate that drought resistances of Z planispinum var. dintanensis can be boosted greatly when 0.06%-0.12% ZnSO4 was used. The seedling cotyledon belongs to Magnolia Type. Aerial part and underground part growth alternate. We should supply water and nourishment according to the plant growth. We adopted model equation Y=a+bX to fit the relationships with height of seedling, ground diameter, top root length and root weight, the results have significant differences. With the model equation, we would use any index to estimate other index in the same time.
(2) Ecological Characteristics:①Combined with cultivation, used MFR between the time of floral initiation and florescence, which can promote the growth of new branch. The application of 20kg every plant has the best effect for increasing production. The best fertilizer composition for the Z. planispinum var. dintanensis is N:P:K= (0.3981-0.4134):(0.2981-0.3134):(0.2864-0.2891).②The fruit growth dynamics of Z. planispinum var. dintanensis showed "S" form. The region which altitude height below 850m is the most suitable area for planting Z. planispinum var. dintanensis.③The biomass of seedling leaf is about 55.00% of all the biomass of plant. It has higher proportion of protein, fats, carbohydrate and other nutritional ingredients. The content of prandial fibre is 2-10 times to other 10 common vegetables. Comparing with other fresh vegetables, Z.planispinum var. dintanensis has the largest proportion of VB2. The contents of seven indispensable mineral substances, which are Ca、Mg、Fe、K、Se、Cu、Mn, are obvious higher than other vegetables.④The chemical components of the volatile oil from Z planispinum var. dingtanensis and Z. bungeanum c."dahongpao" were analyzed by GC-MS.29 components from Z. planispinum var. dingtanensis, and 38 components from Z. bungeanum c."dahongpao". The identified constituents accounted for 99.417% of Z. planispinum var. dingtanensis and 97.685% of Z. bungeanum c."dahongpao". The chemical components in two kinds of volatile oil have significance differences. The maximum of Z. planispinum var. dingtanensis is Linalool L, which is 4.56 times than Z. bungeanum c. "dahongpao".
The innovations of this paper are:(1)In order to discover these reasons of the seed germination characteristic and grow in seedling period, lower germination rate and slow-growing of Z planispinum var. dintanensis, seed morphological structure and physiological characteristics, seed germination physiology, seedling in the Process of water stress, the grow regulation in seedling period being studied. (2) The influence of Zn under drought stress and to fecundity being studied, and then pointing out that the Zn have significant effect on blossoming percent and fruiting percent, comprehensive evaluation of Z. planispinum var. dintanensis under drought stress were evaluated by subordinate function. Drought resistances of Z. planispinum var. dintanensis can be boosted greatly when suitable concentration ZnSO4 were used, the results provided theoretical basis for using Zn when Z.planispinum var. dintanensis been cultivated. (3) The effects of applying fertilizer to the growth and altitude on fruit quality evinced the various effect difference when different fertilizers were used, then pointing out that various(4)The nutrient analysis of young stem and the studied of volatile oil provided scientific base for the new use of Z. planispinum var. dintanensis.
利用目前相关生态学和生理学的方法,主要探讨了以下问题(1)培育的生理特性方面：种子形态结构与生理特点、种子萌发生理、幼苗水分胁迫、微量元素锌对干旱胁迫的影响、施肥对生长的影响、微量元素锌对生殖生理的影响；(2)培育的生态特性方面：苗期生长规律、海拔高度对果实生长的影响、幼茎营养成份分析、挥发油研究。
主要研究结果与结论(1)培育的生理特性方面：①顶坛花椒种子富含油脂,外种皮含蜡质,内种皮为坚硬的厚角质,厚度占种皮的一半以上；顶坛花椒种子内部胚乳残存,子叶还没有完全吸收胚乳中所含全部营养；有12%的种子具双胚现象；这些因素均不利于项坛花椒种子萌发。②立秋至白露中期为顶坛花椒种子采种的最佳时期；碱水处理是理想的项坛花椒种子脱脂处理方法；项坛花椒种子宜采用撒播的播种方式；磷肥作苗床底肥对种子萌发率有极显著影响,是提高顶坛花椒种子萌发率的一项有效措施。③顶坛花椒对水分胁迫的敏感性较强；幼苗随着水分胁迫加剧,叶绿素a、叶绿素b、叶绿素总量以及类胡萝卜素含量明显下降,丙二醛含量增加,游离脯氨酸含量增加；显示出项坛花椒具有一定程度的耐旱性,这也是项坛花椒能够在喀斯特地区生长的生理特性。④当ZnSO4浓度为0.06%-0.09%,顶坛花椒花粉活力和花粉寿命均能大幅度提高；当ZnSO4浓度为0.08%时,顶坛花椒种群开花率可达到最大值；ZnSO4浓度为0.09%时,顶坛花椒种群结果率达到最大值。当项坛花椒处于5-9a时它的生殖力出现峰值；在海拔为600m左右的地段,顶坛花椒具有最大的生殖力。⑤用模糊数学隶属函数法对抗旱性进行了综合评判,适宜浓度的微量元素Zn对顶坛花椒抗旱能力具有较强的促进作用,当ZnSO1预处理浓度为0.06%～0.12%时,能加强顶坛花椒对干旱的适应性和提高抗旱生产力。⑥顶坛花椒幼苗子叶属于木兰型Magnolia Type;幼苗地上和地下部分交替生长,田间管理要适时供应水分和养分；用模式方程Y=a+bX拟合了顶坛花椒幼苗的苗高、地径、主根长和根干重生长关系,可用拟合的模式方程和苗高、地径、主根长和根干重中任一指标估测该时期另三个生长指标。(2)培育生态特性方面：①在开花期到盛花期以沼肥做底肥,能促进新梢生长；以施用20 kg·株-1增产效果最为明显。顶坛花椒的最佳混料肥料配比为mN:mP:mK= (0.3981-0.4134)：(0.2981～0.3134)：(0.2864～0.2891)。②顶坛花椒果实发育生长呈“S”型生长曲线；花椒产量与品质在试验区有随着海拔降低而明显增加的趋势,花江峡谷的中低海拔地区(海拔<850m)属于南亚热带河谷气候,是种植顶坛花椒的优势地区。③顶坛花椒幼苗期叶的生物量占整个植株生物量的55.00%；蛋白质、脂肪、碳水化合物等营养成分含量高；膳食纤维含量为10种其他常见蔬菜嫩茎、叶的2～10倍；VB2含量为所有新鲜植物食品VB2含量之最；7种必需矿质元素Ca、Mg、Fe、K、Se、Cu、Mn含量均明显高于参与比较的蔬菜,顶坛花椒幼嫩茎可作优良蔬菜。④鉴定了贵州顶坛花椒和四川茂县大红袍的挥发油化学成分,顶坛花椒有29个化合物,茂县大红袍有38个化合物；其中顶坛花椒相对含量组分占挥发油总质量的99.417%,茂县大红袍占97.68%。顶坛花椒挥发性成分中的芳樟醇(Linalool L)含量尤其高,达到茂县大红袍的4.56倍,说明顶坛花椒香味好。
本论文的创新点是：(1)从顶坛花椒的种子形态结构与生理学特性、种子萌发生理、幼苗水分胁迫和苗期生长规律方面进行研究,揭示了顶坛花椒种子萌发及苗期生长的特性、对其影响顶坛花椒出苗低、生长慢的原因进行了探索。(2)从微量元素锌对顶坛花椒生殖及水分胁迫的影响,提出了微量元素锌对顶坛花椒种群开花结果率有显著的影响；指出了用模糊数学隶属函数法对顶坛花椒抗旱性进行了综合评判,适宜浓度的微量元素Zn对顶坛花椒抗旱能力具有较强的促进作用,为顶坛花椒在培育方面使用微量元素的效果提供了理论依据。(3)从施肥对顶坛花椒的影响及海拔高度对顶坛花椒的影响,提出了施肥效果好的配方。说明了海拔变化对顶坛花椒产量影响的重要性。(4)对顶坛花椒幼茎叶营养成分的分析及果实挥发油的分析,给顶坛花椒的新用途提供了科学基础。
"prickly ash" is the general designation of Zanthoxylum L.. Zanthoxylum planispinum var. dintanensis Yu-Lin Tu is a variety of Zanthoxylum planispinum Sieb.et Zucc., and in Guizhou karst areas it is a specific perfume plant, with higher ecological and economic value. Huajiang is the typical area of Guizhou krast regions, Z. planispinum var. dintanensis has been planted more than 10 years there, but there are still some growth and production questions to puzzle the people continuously, such as low seedling emergence rate, weak growth vigor (yellow leaves), low yield, sole application ways and so on. In order to solve these problems, optimized the cultivation of Z planispinum var. dintanensis, the charictristis study of plant physiology and ecology are very important.
Using of the current methods related to ecology and physiology. In this paper, Z. planispinum var. dintanensis in karst areas was selected as the study object. Mainly has carried on the discussion from following several aspects:(1) Plant physiology characteristic:seed morphological structure and physiological characteristics, seed germination physiology, seedling in the Process of water stress, the influence of Zn under drought stress and the effect of applying fertilizer to the growth. (2) Ecological Characteristics:the grow regulation in seedling period, the effects of altitudes on fruit quality, the nutrient analysis of young stem and the studt of volatile oil.
The main results and conclusion are showed as follows:
(1) Plant physiology characteristics:①The seeds of the Z planispinum var. dintanensis contained a large amount of oil, the exopleura wax, the endotesta with hard cuticle layer, which take more than half of the total thickness of the seed coat. Endosperm remains in internal seed, cotyledons have not completely absorbed all the nutrition in the endosperm. There are 12% seeds have biovulate phenomena. All factors mentioned above are not propitious for the seed germination of Z. planispinum var. dintanensis.②The seed collecting time is better later than earlier. The late time from autumn begins to White Dew are the best time for seed collecting. Treating with aqueous alkali is the ideal method for seed derosination of Z. planispinum var. dintanensis. Compared with other regions, there is better quantity of heat in karst xerothermic valley, it is better planting methods to use broadcast for the seed of Z. planispinum var. dintanensis. Using phosphorus as base manure to the seedling bed has very obvious influence to seed germination. This is a effective method to improve the seed germination rate of Z. planispinum var. dintanensis.③With the aggravation of water stress, the chlorophyll a, chlorophyll b, Gross chlorophyll and carotenoid of Z planispinum var. dintanensis. seedlings decreased obviously, the MDA content and the POD activation increased. In the early stage of water stress, the seedlings were more damaged. The results showed that Z. planispinum var. dintanensis. has a certainty drought tolerance, it was the foundation of Z. planispinum var. dintanensis. adapting in karst mountainous country.④When ZnSO4 were between 0.06% and 0.09%, both pollen viability and life-span of Z. planispinum var. dintanensis can be boosted greatly. When ZnSO4 were between 0.08% and 0.09%, both blossoming percent and fruiting percent of Z planispinum var. dintanensis can be boosted greatly. The peak value of its reproductive age is during 5-9 years old, while at an altitude of 600m, the Z.planispinum var. dintanensis have the max fecundity.⑤The results by fuzzy comprehensive appraisal analysis indicate that drought resistances of Z planispinum var. dintanensis can be boosted greatly when 0.06%-0.12% ZnSO4 was used. The seedling cotyledon belongs to Magnolia Type. Aerial part and underground part growth alternate. We should supply water and nourishment according to the plant growth. We adopted model equation Y=a+bX to fit the relationships with height of seedling, ground diameter, top root length and root weight, the results have significant differences. With the model equation, we would use any index to estimate other index in the same time.
(2) Ecological Characteristics:①Combined with cultivation, used MFR between the time of floral initiation and florescence, which can promote the growth of new branch. The application of 20kg every plant has the best effect for increasing production. The best fertilizer composition for the Z. planispinum var. dintanensis is N:P:K= (0.3981-0.4134):(0.2981-0.3134):(0.2864-0.2891).②The fruit growth dynamics of Z. planispinum var. dintanensis showed "S" form. The region which altitude height below 850m is the most suitable area for planting Z. planispinum var. dintanensis.③The biomass of seedling leaf is about 55.00% of all the biomass of plant. It has higher proportion of protein, fats, carbohydrate and other nutritional ingredients. The content of prandial fibre is 2-10 times to other 10 common vegetables. Comparing with other fresh vegetables, Z.planispinum var. dintanensis has the largest proportion of VB2. The contents of seven indispensable mineral substances, which are Ca、Mg、Fe、K、Se、Cu、Mn, are obvious higher than other vegetables.④The chemical components of the volatile oil from Z planispinum var. dingtanensis and Z. bungeanum c."dahongpao" were analyzed by GC-MS.29 components from Z. planispinum var. dingtanensis, and 38 components from Z. bungeanum c."dahongpao". The identified constituents accounted for 99.417% of Z. planispinum var. dingtanensis and 97.685% of Z. bungeanum c."dahongpao". The chemical components in two kinds of volatile oil have significance differences. The maximum of Z. planispinum var. dingtanensis is Linalool L, which is 4.56 times than Z. bungeanum c. "dahongpao".
The innovations of this paper are:(1)In order to discover these reasons of the seed germination characteristic and grow in seedling period, lower germination rate and slow-growing of Z planispinum var. dintanensis, seed morphological structure and physiological characteristics, seed germination physiology, seedling in the Process of water stress, the grow regulation in seedling period being studied. (2) The influence of Zn under drought stress and to fecundity being studied, and then pointing out that the Zn have significant effect on blossoming percent and fruiting percent, comprehensive evaluation of Z. planispinum var. dintanensis under drought stress were evaluated by subordinate function. Drought resistances of Z. planispinum var. dintanensis can be boosted greatly when suitable concentration ZnSO4 were used, the results provided theoretical basis for using Zn when Z.planispinum var. dintanensis been cultivated. (3) The effects of applying fertilizer to the growth and altitude on fruit quality evinced the various effect difference when different fertilizers were used, then pointing out that various(4)The nutrient analysis of young stem and the studied of volatile oil provided scientific base for the new use of Z. planispinum var. dintanensis.
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